trvolution

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 $-19.95

Slrltttttittg evolutiolrrrv sclence

fiorrr its irrception to its lrrtest firrcl-
irrgs. tl-oni cliscovcrics lrncl cllrtlr to
philosophv :rncl historl', tliis book is
the ntost conrpletc, authoritlttir,'e,ancl
irrviting one volunte introcluction tct
solutionrrlv bioloi1' avrril:iblc.(lIe:rr,
i r r f i r r r r r r t t i v e r, r n d c o n r ; r r c h e n s i v e i n
s c o p c , E l t r l r r t i o r io p e n s u , ' i t h r r s e r i c s
of rrrrtjor css:rr..s
rlerrlinq u,ith the his-
tolv rrntl philosophl' oi er,'olutionnrv
biologr'. uith nrrrjor crrrpiricrrl :rrrci
t l r e o r e t i c r r lq u c s t i o n s i n t h e s c i c n c c ,
fioni s1-rccilrtiorrtc'r:rclrrptrrtion.firtrri
prllcor)toloqv to evolutiorirrv tlevel
opnrclrt (evo-tlcr,o). rrrrc'lconclurling

'#*
't sr
with essayson the social and political sienifi-
cance of evolutionary biology today.
A second, encyclopedic section travels the
spectrum of topics in evolution with concise,
informative, and accessibleentries on individ-
uals from Aristotle and Linnaeus to Louis
Leakey and Jean Lamarck; from T. H. Huxley
and Edward O. Wilson to Joseph Felsenstein
and Motoo Kimura; and on subjects fronr al-
truism and an-rphibiansto evolutionary psy-
chology and Piltdown Man to the Scopestrial
and social Darwinism. Readers will find the
latest word on the history and philosophy of
evolution, the nuancesof the scienceitself, and
the intricate interplay anong evolutionary
study,religion, philosophy, and sociery.
Appearing at the beginning of the Darwin
Year of 2oo9-the zooth anniversary of the
birth of Charles Darwin and the rjoth an-
niversary of the publication of the Origirt o-f
Species-this volume is a fitting tribute to the
s c i e n c eD a r w i n s e t i n n l o t i o n .

M TcHAEL RUSE i s L u c y l eT . W e r k n r e i s t e r
Professor of Philosophy and Director of the Pro-
granr in the History and Philosophy of Science,
Florida State (Jniversiry.He is the founder and edi-
tor ofthejournal Biologyand Philosophyand has ap-
peared on "Quirks and Quarks" and the Discovery
Channel. His books include Tlrc Euolution-Crcdtiort
Struggleand Danyin and Desigtrftoth from Harvard).

J oS EPH T R A V T S i s D e a n o f t h e C o l l e g eo f
Arts and Sciencesand Robert O. Lawton Distin-
guished Professor of Biological Science at Florida
State Universiry.

The BelknapPressof HarvardUniversiryPress

Cambridge,Nlassathusetts,
attd Lturdon,England

www.hup.harvard. edu

Jacket illustration: DEA Picturr Librarv/(letn Inrrqes

Jacket design: Jill Breitbarth

"One of the first and likely to be the most comprehensive in scope of the
many books occasionedby the zoog sesquicentennialofDarwin's Origin."

-Edward O.Wilson

FROM THE EDITORS' INTRODUCTION

This book presentsevolutionary science as a modern, dynamic discipline

and allows the reader to explore evolution according to his or her inter-
estsand background. The reader who wishes to be immersed in the sci-
ence can focus on the major scientific themes, while the reader who is
interested in the intellectual history and influence of the subject away from
science can enter through a separateset of essays.
A reader interested in
very specific topics or historical figures can find the appropriate entries,
along with essayson related topics. Our goal was to provide an exciting
and compelling introduction to evolution along with a basic reference
work that could point the way toward a deeper study of individual issues.
The essaysinclude bibliographies, which serve as guides to further and
deeper reading.The essaysin this book show not only how far we have
come but where the scientific horizons lie and how we might move to-
ward those horizons.

rsBN378-0-b7rr-03175-3
90000

llillfflilllilllllllllll
ilililil]tl
VoLUTloN
The First Four Billion Yers

EDlD BY

MlHAL RUsE
JosPH TRAVls

W|H A FoRWoRD BY

DWARD o. WlLsoN

H E E L K N A PP R s s o F
A R V A R DU N l V R s | YP R s s
A t ' 4 B R I D G
A. ssAHUss
L o N D o N , N G L A N D
2009
 ontnts

opyright o 2009 by th Prsidntand Fllows of arvrd Collg Frword

drd .Wils,
All rights rsrvd
Introdution
Printd ir-rth Unitd Stts Of Amri:r NIihl Ruse l
Librr of ogrss tlogig-i-Publition Dt Th Histoy of volu
vlution : th first flur illion yars / ditd by ihal Rus, Josph Trvis ; ihl Rs
with a forword dwrd O. Wilson.
p. m. Th origin of Lif
Inludsiliogphialrfrnesand indx' Jffr L' Bd c
I S B N 9 7 8 . 0 - 6 74 - 0 3 | 7 5 . ( a l k . p a p r )
1. volution (Biotogy) I. Rus, ihal. II. Travis,Josph, 1953- Palontolgy and th
QH366.2.86 z09 ihI Bnton
576.8-d22 2008030270
Adaptation
Josph Truis t

olulr volutin
FrnisoJ.A

volution of th Gnr
Brin hrles

Th Pattrn and Pro

rgrt B. Pt,

volution and Dvl

Grgor A. IVr

Soial Bhavior and S

Dil I. Rub

Human volution
Hnr . H

votutionyBilogy
ihl F' nto
 Contnts

nd Fllows of Hrvard olig Foword UTl

drd O' Wilson
srvd
Introdution i
'ttsof Amria
ihl Rus d Josph Truis
,g-i- Pub litkl
D t
Th History of volutinary Thought
d by Mihl Rus, Josph Travis ; ihl Rs
ward O. Wilson.
Th origin of Lif 49
frnsand ird.
'5-3 (alk. par) JffreL. Bd nd Atonio Lzno
hal. II. Travis,Josph' 1953-
3 2009
Palontologyand th History of Lif 80
20080270 ih,ll Bto

Adaptatin 105
Josph ruis nd Duid N. Rzik
olulavolution 12
Frnisol. l

vlution f th Gnom 152

Bri brlstuorthnd Dborh hrlsorth

Th Pattrn and Prossof Spiation 177

rgrt B, Ptk nd Sbl l. nktso

volution and Dvlopmnt 208

Crgor A' Wr

Soial Bhavir and Soibiolgy 27

DnielI,Rbnstein

Human volution 256

nr . nr

volutionaryBiolgy of Disasand Darwinian diin 281

ihlF. Anto|
Contts

Byond th Darwinian Paradigm:UndrstandingBiolgial Foms 299

Brin Goodtui

Philosphy of volutionary Thougt 31

Kim Strl

volution and Soity 0 Forwor

frd D. Lubiblr d J ishin

volution and Rligion 48 drd

Duid N' Liuigston

Amian Antivolutionism: Rtrospt and Prospt 70

ugi. Sott

Alphabtial Guid 401

Two nturis aftr i
ontritors 9S harls Darwin's ozz
important ook vr.
Illustrationrdits 949 te rigi hav not t
Ind ar not prahd fr
951
oath of offi with a
drssd th living wol
ity's pla within it, v
massiv sintifi vid
hav grown ontinuo
undrstanding and tht
So solidly hav th
of volution that it ml
(univrsal prinipls i:
first law is that all th
ultimatly obdint t
has n th driving f
half ntury. Th s
iology as wll as mu(
lmnts and pross
volution through nat
h two laws hav 1
for th rognition of
gr thy ar bing o
logial phnomna. T
th sondaddrsss
xampl, ar built and
not som othr. ah
lvIodrn biology as a l
proahs guidd y th
vn without th i
sam as th Darwin il
rstanding
BiologiaIForms 299

0 Forword
ishi

48 durd. Wilson

t nd Pospt 70

401
Two nturis aftr its author's irh and 150 yars aftr its puli:rtion,
935 hrls Darwin's the rigin of Speis an fairly rnkd as th most
important book vr writtn. Not th most widly rad, to sur. opis of
949 th origi hav not bn plad in hotl rooms aross Amria; its vritis
ar not prahdfrom pulpits alr Sunday; and no politial ladr taks an
951
oath of offi with a hnd on its ovr. It is th mastri that first ad.
drssdth living world and (with he Descnt of following) human-
ity pl within it, without rfrnto any rligion or idology and upon
massivsintifi vidn providd aross sussiv dds. Its argumnts
hav grown ontinuously in stm as th st foundation for hran slf-
undrstndingand th pilosophial guid for human rion.
So solidly hav th filds of iology uilt upn th Darwinian onption
of volution that it maks sns tday to rogniz it as on of th two laws
(univsalprinipls if you wish) that govrn ou undrstanding of lif. Th
firs law is that all th lmnts and prosss tht dfin living organisms ar
ultimatly odint to th laws of physis and hmistry. This formulation
has bn th diving for of molular and llular iology during th past
half ntury' Th sod law, th foundation :rnd produt of volutionary
iology as wll s muh of organismi and nvironmntal iology, is that all
lmnts and prosss dfining living organisms hav n gnratd y
volutionthrugh natural sltiln.
Th two laws hav provn to frrlly omplmntary, a prim rquirmnr
for th rognition of sintifi laws gnrally. Furthr' to an inrasing d-
grthy ar ing omind to ahiv samlssanalyss of partiular i-
logial phnomna. Th first law addsss how a phnonrnon ours and
th sond addrssswhy it ours. Mitohondria and othr organlls, for
xampl, ar built and work thus and so; and thy originatd in on way and
not som othr. ah dsription omplts th xplanation for th othr.
odrn biology as a whol has littl maning without th joining of both ap-
proahs guidd y th two lws.
vn without th imptus of th origin ssquintnnial-fortuitously th
samas th Drwin irth intnnil-2009 is an pproprit tim to prsnt

u
ultx Foreord

volutionary iology in its nompassing modrn form. I dout that any singl
writr, or any nsml of sveral writrs, ould summariz this urgoning
disiplin in a omprhnsiv and authoritativ form. Th multipliity of au-
thors in uolutio: The First For Billio Yers has gon far to*ard that goal.
Its array of prts provid stat of th art for ah sujt in turn. Th sin
is prsntd hr' ut also som of th main pratial onsquns and in.
vitably, with fundamntal rlvan, th onsquns of iologial volu- Introdu
tionary thought for philosophy and rligion. Th grat qustions-..Who ar
w?'' ...Whrdid w om from?'' and ...Whyar w hgrg}''-gan nswrd
only, if vr, in th light of sintifially asd volutionary thought.
NIihlR

h disovry of volr
.W.strn
thought, ranl
mong sintifi dis<
From sing natur aI
forvr hanging, w
drstanding volution
In sin th word
idas. First, thr is tl
ganisms, living and dt
natural pross of h
othr, diffrnt ons. l
of th anitlt Grks.
ganisms volvd rally
sidrd a asi elm
pulishd is on tbe
organisms on th plar
that opratd rlntls
th natral world. An
tology to moleular i
vidn that volutiol
Sond, uolution is
glo. This an rfr t
volution of orhids,
suh as th volution <
reptil iaw. volution:
vidn that rangs fi
to similaritis and diff
tin of som faturs.
stood, whil th volu
organlls of lls, is l
h third ida is th
that volution has not <
ldrn form. I dout that any singl
ould summaizthis urgoning
tiv form. Th multipliity of au-
,rs has gon far toward that goal.
lr ah sujtin turn. Th sin
in pratial onsqunsand in-
:onsqunsof iologial volu-
Introdution
. Th grat qustions-cGWho ar
Y arewe h?''_an answrd ihl Rse d JosphTruis
d volutionarythought.

Th disovry of evolution is on of th gratst intlltual ahivmnts of

.Wstrn
thought, rnking with th alulus and gnrl and spifi rlativity
among sintifi disovris that hangd indlily how w s our world.
From sing natur as fid torvr in form and omposition to sing it as
forvr hanging, w hav n transformd uttrly y disovring and un-
drstanding volution.
In sin th word euoltion is usd to prss thr diffrnt ut rlatd
ids. Fist, thr is re ft of volution. This is th ralization that all or-
ganisms, living and dad' irrluding humans, ar th nd produts of a long
natural pross of hang through whih ah spis is dsndd from
othr, diffrnt ons. Although protovolutionary idas dat ak to the tim
of th nint Grks, only in th ightnth ntury did th laim that or-
ganisms volvd rally start to gain urrny. Howvr' th ida was not on-
sidred a asi lmnt of sintifi knowldg until aftr harls Darwin
pulishd his ozl the rigi of Spis in 1859. Darwin argud that all th
organisms on th plant mrgd through a singl, straightforward pross
that opratd rlntlssly from th dawn of lif and that ontinus to shap
th natural world. An normous amount of sintifirsarh,from palon-
tology to molular biology' has providd th omplling and ovrwhlming
vidnthat volution is indd a fat.
Sond, uoltion is oftn usd to rfr to the pth of lif's history on this
glo.This an ref to th history of a group of isting spis,as in th
volutin of orhids' or th path through whih partiular traits mrgd,
suh as th volution of th middl ar of mammals from small ons in th
ptil |w. volutionary historis ar onstrutd from a wid varity of
vidnthat rangs from squns of spis and forms in th fossil rord
to similaritis and diffrns in th DNA of isting organisms. Th volu-
tion of som faturs,suh as th hoovs of ungulats,is vry wll undr-
stood, whil th volution of othrs, partiularly vry old fatures lik th
organllsof lls, is lss wll undrstood.
Th third ida is the theor of volution. This rfrs not to a suggstion
that volution hs not orrd or that th historv of ortiular fatursannot
Introdution

ronstrutd ut to orrr idas aout th fors that driv volutionar bhavior. Thr ar f
hang.In the rigi Darwin propsed what today is almost univrsallyon- whih rval that ds1
sidrd th major for in organi hang, natural sltion.But natural s- ovrd. Not that th.
ltion is not th only importnt volutionary for; natural sltion oprats rathr' thr is muh
on th variation in shaps, sizs, and iorrs of organisms atd y muta- spis or faturs' pa
tions in thir gns' and th rt and mirgnitud of th hangs that ar la lvls' and to rfinr
drivn y natural sltinare govndy hw thos gnsontrol dvlop- stonishing divrsity
ment. An enormous amount of osrvation and eprimnt has doumntd ook show ot only h
th ation of naturl sltionand som of th gnti rsPonssto sltion. li and how w might
But modrn disovrisin gntisand dvlopmnt have raisd many nw This ook is organi
qustions and guidd us toward sorsurprising answrs out how all ths on rh varhing tt
fors omin to driv volution. thought to th ontro
Outsid sin th word uolution rfers to a natural pross onsidrd origin of lif to th p
the inspiration for host of soiologial, litrary, politial' philosophial, m a j r f a t u r so f v o
and rligious ids. From writs xploring th animal natur of humans to ras lik molular .
psyhlogistssarhirrgfor th origirrof humn havior in volutionary his. Th sond pat onti
tory to thologians grappling with th impliations of volution for ou un- topis. Ths inlud t
drstanding of th Divin, th influn of volution outsid sin f p o p l a f m i l i a r ,t
ds that f any othr sintifi disovry vr mad. And as modrn on- urs in th disovy
trovrsis in th shools illustrat, partiularl in th Unitd Stats' it pro. oks that rount tht
voks rtiolls lmost s stong as issussurrounding th eginningand nd a disiplin'
of hnran lif. This organizat\on a
In this ook w xplor all ths fts of volution. Our uthors prsnt or hr intstsand ;
th vidn for volution as fat from th fossil rord to gnomis, illus- th sin an fous <
trat th histry f groups from atri to irds, and dsri our urrnt intrstd in th intlI
idas about how hs historis and tlris vidn am to b. Our uthors sin an ntr throl
also xplor tlr influn of voltrtion on plrilosopy, rligion, soiology, spifi topis or hist
psyhology, and many othr aras and addrss th urrnt ontovrsis with ssays on rlatd
aout th tahingof volution in shools.Th ook ovrs th historial dis- plling introdution to
overis, suh as homology, that opnd th path for disovring volution; point th way towad
th phnomena' suh s industrial mlanism, that playd maior rols in in- biliogaphisw , hih
spiring primntal studis of volution; th appliations of volutionary Although w hop t
prinipls to aras smingly far fild, suh as omputr sin;and th volum to inform,
ontributions f th major figurswho shaped th history <lfvlutionary si- sholars. W find vol
n and th disiplin as it xists today. intrsting, and ind
Prhas most important' this book prsnts voltionary sin as a mod- mrg with th sam
rn, dynami disiplin. All too oftn disussions of volution outsid th si- t i o n , i t i s i m p o t a n tt .
n itslf giv th imprssion tht th subjt am fssilizd in th hallng of antiiot
nintnthntuy' This is partiularly so whn th suitaility of volution as tious disass,it shor
a sujt for sondary-shoolstudntsis ing argud.Th irony is tht th a l t r a t i n so f h a i r a t
disovriesof modrn sin,from molular iology to otl1putationalin- planations of who '
novation' hav providd onrplling vidn that Darwin ould only dram m a n y o f o u r l o s l yh
of sing. From thos disovrishav also om appliations that Darwin h last of ths r
might nvr hav imagind, suh as Darwinian mdiin. Prinipls of volu- Provn so ontrovrs
tion ar ing applid to a wid vrity of sintifi prolems, from undr- popl in th Unitd S
stnding snsn to stratgis for onsrvation to plaining hunran of lif on arth. lt is u

th lrs tht div volutionary
hat today is almost univrsallyon-
:, natural sltion'But natural s-
ar for;ntulsltionopats
ms of organisms rated y muta-
ragnitud of th lrngs that ar
y how thosgnsontrol dvlop-
ln and primnthas doumntd
f the gntirsponssto sltion.
vlopmnt hav raiso many nw
lrising answrsabotrthow all ths
:rs to naturl Pross onsidrd
l, litrary, politial, philosophial,
lg th animal natur of humans to
mn bhavior in volutionary his-
pliationsof volution for our un-
f volution outsid sin far
ry vr mad. And as modrn on-
r l a r l y i n t h U n i t d S t a t s 'i t r o -
;urroundingth ginning and nd
of volution.our authors prsnt
r fossil rord to gnomis, illus-
to irds, and dsriOur urrnt
:vidnam to . our authors
n philosophy, rligion, soiology,
addrss th urrn ontrovrsis
Th ook ovsth historial dis-
h path for disovring volution;
sm' that playd major rols in in-
th appliations of volutionary
rh as omputr sin; nd th
ld th history of volutionarysi-
nts volutionarysieneas a mod-
ssionsof volution outsid th si-
subjt am fossilizd in th
lhn th suitailityof volution as
ringargud.Th irny is tht th
ular iology to omputational in-
that Darwin ould only dram
c om appliations that Darwin
Lianmdiin.Prinipls of volu-
.sintifiprolms,
from und-
]srvation to plaining human
ltroductio i
havior. Thr ar fw topis in sin with so many xiting nw fats'
whih rval that dspit all w hav larnd, thr is muh mor to dis-
ovrd. Not that thr is futhr ned to sustantiat volution as fat;
rathr' thr is muh mor to don to tra th volution of groups of
spisor faturs, partiularly omplx faturs t th llular and molu-
lar lvls, and to rfin our thory of volution to aont mor fully for th
astonishing divrsity of lif as w ontinu to disovr it. Th ssays in this
book show not onl how far w hav om but whr th sintifi horizons
li and how w might mov toward thos horizons.
This ook is organizd into two parts. Th first part ontains long ssays
on th ovrarhing thms in volution. From th history of volutionary
thought to th <rntrovrsisovr duation, from th fossil rord and th
origin of lif to th pross of adaptation, th long ssays offr primrs on th
majo faturs of volution. Many of thm off los looks at partiular
aras lik molular volution' gnomi volution' and Darwinian mdiin.
Th sond part ontains a larg numbr of shortr ssays on mor spifi
topis.Ths inlud ssayson malor groups of organisms with whih most
popl ar familiar' topis that a important fats of volution, major fig-
us in th disovry and shaping of volutionary sin' and th ritial
ooks that rount th history of disovry, dvlopmnt, and maturation of
a disiplin.
This organization allows th adr to xplor volution arding to his
or hr intrsts and bakground. h radr who wishes to b immrsd in
th sin an fous on th mjor sintifi thms, whil th adr who is
intrstdin th intlltual history and influn of th sujt away from
sinan ntr through a sparat st of ssays. A radr intrstd in vry
spifi topis o historial figurs an find th appropriat ntris, along
with ssays on rlatd topis. Ou goal was to provid an iting and om-
plling introdution to volution along with a basi rfernwork that ould
point th way toward a dpr study of individual issus. Th ssays inlud
ibliographis' whih srv as guids to furthr and dpr rading.
Although w hop that our fllow sholars njoy ths ssays' W want this
volum to infom, duat' and it radrs who ar not profssional
sholars.\w find voltion in all of th word's manings to a provoativ,
intersting,and indd aw-inspiring topi' and we want our radrs to
mrgwith th sam fling. But yond th itemnt of studying volu-
tion, it is important to undrstand and appriat it. voltion plains th
hallng of antibioti.rsistant pathogns and th sourg of novl inf-
tious disass, it shows us wht W an xpt limt hang and our own
altrationsof haitt to do to th natural world, and it may offr profound
xplanations of who w ar and why we hav as w do. It also hallngs
many of our losly hld lifs about man's, and woman's, pla in natur.
Th last of ths issus is prhaps th fondation of why volution has
provn so ontrovrsial. Survy aftr survy shows that fwr than 50% of
poplin th Unitd Statsapt that volution is rsponsil for th divrsity
of lif on arth. It is unlar how muh of that opposition is asd on a poor
Introdution

undrstanding of volution and th vidn aout it and how muh is

asd
on th hallngs that it offrs. Yt opposition to volution y rligious po-
pl, spialiy y hristians' is y no mns invital; som of th'most
dis-
tinguishd livrs hav aptd volution without many qualms. N
ar
onvind that th hristian-with th Jw and uslim and other rligious
blivrs-should at th had of th quu of thos who welom evolu-
tionary idas. Finding th ida, uilding and laorating upon it, and looking Th Hir
at its influn and importan ar' as th ontriutors to this volum show
again and again, truly th st of proofs that w ar mad in th imag
of
Thougt
God, lrating ration in all its wondrful manifstations.
This ook is a tstamnt to on of mankind's
8ratst disovris, a disov-
ry that offrs unparallld insight into what ration rally is. I/lov
NIiblR
vo-
lution, and if w and our fllow ontriutors an inspir you vn in a
small
way with our nthusiasm' thn w shall happy indd.

Th ida that all org

mans from othr forl
(fifth ntury ...) tr
ightnth ntrrry an(
ida of natural dvlo
portrs. hr ar ra
gan to gain momnl

Th arly Days

Th Grks had no grt

tur did not hav a plz
Spifially, th Grk
going, inrmntal o
Plato and Aristotl-l
isms) showd ordr
simply hav appard
tainly was not somtt
th omplity of tod;
and dsign: ..Sinw,sl
and tos. Th mitur
of a singl stuff, a pi
on to put things in ol

Now thse Wer ml

nnt aus of its pr
gnrations: our I
ralm of wild ast
ular thy knw that
and laws or hoofs
 aout it and how muh is asd
tion to volution y rligious po-
rs invitabl;som of th most dis-
.!7
on without many qualms. ar
w and uslim and othr rligious
ruof thos who wlom volu-
d laorating upon it, and looking
Th History of volutionary
ontributors to this volum show
that we ar md in th imag of
Thought
[ul manifstations.
ind gratstdisovris,a disov-
lat ration rally is.
.s7 ihel Ruse
lov vo.
)rs an inspir you vn in a small
happy indd.

Th ida that all organisms (inluding humans) ar gnratd by natural

mans from oth forms has anint roots. Aristotl tlls us that mpdols
(fifthntury ...) toyd with suh thoughts.Howvr, it was not until th
ightnthntury and th nlightnmnt that uolutio (as w now all this
ida of natural dvlopmnt) rally startd to gain a srious numr of sup.
ports. Thr a rasons oth for th long dlay and why th ida finally
ganto gain momntum.

Th arly Days

Th Grks had no grat rligious ojtion to volution, ut thir world pi-

tur did not hav a pla for any kind of signifiantdvlopmntalprosses.
Spifially,th Grks thought that thy had irrftal rasons to rjt on-
going, inrmntal organi hang. Thy-partiularly th philosophrs
Plto and Aristotl-thought that th world (spially th world of organ-
isms)shwd ord and intntion and, as suh, was not somthing that ould
simply hav appard thrugh lind, ungovrnd prosss of law. It r-
tainly was not somthing that ould hav grown from simpl ginnings to
th omplity of today. Plato usd human fingrnails as an ampl of ordr
and dsign: ..SinW,skin and on wr intrwovn at th nds of our fingrs
and tos. Th mitur of ths thr was drid out' rsulting in th formation
of a singl stuff' a pi of hard skin, th sam in vry s.'' Plato tn wnt
on to put things in ontt.

Now ths wr mrly auxiliary ass in its formation-the prmi-

nnt us of its prodution was th purpos that took aount of futur
gnrations: our rators knw that on day womn and th whol
rlm of wild astswould on day om to from mn, and in parti-
ular thy knw that many of ths offspring would nd th us of nails
and laws or hofs for many purposs. This is why thy took ar to
The Histor of uolutior Thought

inlud nils formd in a rudimntay Way in thir dsign for hu- humn thought and
mnkind, right at th start. This was thir raso, thn, and ths th pogrss.
profssd aims that guidd thm in making skin, hair and nails grow at Th British physii;
th xtrmitis of our |ims. (Timus, 76d_e, in oopr 1997, 1277) harls) Was a para
mankind:
In an inrdily influntial disssion, Aristotl in D prtibs nimlium
Imp:
(1,984\identifid th fators at work hre as ..final auss.'' Ths ar auss
Of la
that our not just to produ or do somthing (th fingr parts dry and mak With
ails) ut fo the sak of som kind of purpos (th nails prott th fingr And r
nds). Thy show som kind of forethought or intntion. For this rason, fi.
Arost
nal auss annot rdued to lind, unguidd law, as is dmandd in vo- An r
lution. Th world, partiularly th world of orgnisms' must in som sns
hav n dsignd rathr than just produd undr its own stam tly natual (.D
prosss.(Sdly2008 is th dfinitivstudy.)
This is all rh nd pr.
Th Jws, and following thm th arly Christians, had religious rasons
gn' whih auss
for th rjtion of volution. It gos against th ration storis of th arly
haptrs of Gnsis, whih portray a world ratd miulously y God and Hw lo
thn popld y him through divin fiat ovr a short tim span. But do not From .
think that rligion as suh was thn and always an asolut ar to volution- ow fr.
ism. Th hurh fathrs (th major hristian thologians of th arly n- h pilt
turis) workd toward an undrstanding f th ilial txt tht wuld allow
intprtation, partiularly in th fa of advans of sin. Sint Augustin
was agr not to lt anint ration aounts stand in th way of modrn Similar idas wr t
thought. H himslf, living that God stands outsid tim, spulatd in a Pbilosophi zoologiq
kind of protovolutionary fashion that th Divin had fomd sds of lif produd th first ful
that thn sprang into full ing whn thy wr plad hr on arth. How- to our own spis fr
vr' on should not ad too muh into any of this. Lik th Grks, th n produd from l
Jws and Christians wr simply not looking in th dirtion of volution and ity and othr natural
would hav thought final auss an unnswrabl oition to signifiant d- Although th mta1
vlopmntalism. As is wll known, ths kinds of auss am a founda' ris of volutionary i<
tion of on of th major proofs of God's istn, th Argumnt from vidn. Aristotl ha
Dsign, whih movs frm dsign her on arth to th xistn of th divin sha ommon pattrl
artifir. and th volutionists
.!hy,
thn, did volution start its rise in the ightnthentury? Th an. tris (Figur 1). Likr
swr is simpl. It was at this time that popl startd to hallng th 8o unnotid (Figur
Christin pitur of world histry-a providntial pitu of a world r. sition. Th fossil r
atd y God, whr humans ar mad in his imag ut hav falln and ar would invok first as
abl to ahiv salvation only through his undsrvd gra. Som gan ful. As a systmati p
to argu that prhaps humans hld thir fats in thir own hnds and were magr inded.
ould progrssivly improv thir own lots. It Was this ida of pOgrss- fat that no on hal l
th lif that th world and its dnizns ar on a trajtory upward nd ltion. ost assm
that this upward ris is mad possil y (and only y) th unaidd fforts fors, ut littl mor
of th world's human inhabitants-that gav ris to th ida of organi that haratristisa
volution (Rus 1996). nthusiasts for progrss tndd thi thinking ately to th futur g
into natur and dvlopd th id of volution-progrssiv hang up- that th inhritan
ward from th simpl to th ompl. They thn rad this ida ak into Lmrkism*ut bt
 Th Histor fuolutior Thougbt 3

f wa in thir dsign for hu- human thought and soial prati as onfirmation of thir lifs about
thir rason, thn' and ths th progrss.
king skin, hair and nails grow at Th British physiian and man of sin rasmus Darwin (grandfathr of
76d-r, in Coopr 1997,|277) harls) was a paradigm who hymnd in vrs lif's upward ris to hu-
mankind:
'ristotl i D prtibs imlium
Imprious man, who ruls th stil rowd,
as ..final auses.''Ths ar auss
of languag'rason' arrd rfltionproud,
:hing (th firrgrparts dry and mak .With
brow rt who sorns this arthy sod,
urpos (th nails prott th finger
And stlshimslf th imag of his God;
;ht or intntion. For this rason, fi-
Aros from rudimnts of form and Sns,
guidd law, as is dmandd in vo-
An mryon point' or mirosopi ns!
of organisms)must in som sns
d undr its own stam by natural (. Darwin 180' 1: anto I,lins 09-14)
udv.)
This is all th nd produt of th progrssivdvlopmnt of human intlli-
y hristians,had rliglous rasons
gn,whih aussarrd is rfltdin humans' sintifiahivmnts:
nst th ration storis of th arly
d atdmiraulously y God and How lovs nd tasts,and svmpathisommn
vr short tim spn. But do ot From vansntnotis of sns;
lwys an absolut ar to volution- H<lw from yilding touh and rolling ys
stian thologians of th arly n- Th pils immns of humn sinris!
'f th ilial txt that would allow
(anto III,lins 4346\
dvans of sin.Saint Augustin
ounts stand in th way of modrn Similar idas wr to found lswh,most notaly in Fran. In his
tands outsid tim, spulatd in a Philosophie zoologiqu (1809), th taonomist Jan.Baptist d Lamark
r Divin had formd sds of lif produd th first full-blown volutionay thory-a pitur of upward ris
F w plad hr on ath. How. to ou own spisfrom th most primitiv forms of lif, whih in turn had
, any of this. Lik th Grks, th nprodud from mud and slim through th ations of hat and ltri-
ng in th dirtion of volution and ity and othr natural fors.
wralojtion to signifiant d- Aithough th mtaphysial ida of progss was the main fator bhind th
kinds of auss am a founda- ris of volutionary idas, it is not tru that thr Was no prtinnt mpirial
j's istn, th Argumnt vidn.Aristotl had notd that organisms of vry diffrnt spissm to
from
arth to th istn of th divin sharommon pattrns or struturs-wht today ar known as iromologis-
and th volutionists wr rady to intrprt ths as signs of ommon ans-
n th ightnthntury? T an- tris (Figur 1). Likwis, th susssof animal nd plnt brdrsdid not
: popl statd to hallng th go unnotid(Figur2). But gnrallyth vidntook vry sondary po-
ovidntial pitur of a world r- sition. Th fossil rord, somthing that today many (if not most) popl
his img ut hav falin and are would invok first as th proof of dvlopnrntalorigins, was lss than hlp-
is undsrvdgra. Som gan ful. As a systmati prof of progssiv hang, th glanings from th roks
ir fats in thir own hands and Wmagrindd. In any s, ounting against th mpirial sid was th
)ts.It was this ida of progrss- fat tht no on had any grat undrstandingf wht might hav ausd vo-
i ar on a trajetory upward and lution. ost assumd som kind of vagu' upwardly thrusting for or
(and only y) th unaidd fforts fors, but littl mor. Gnrally, vryon was ommittd to th folk blif
gv ris to th ida of organi that haratristisquird irr on gnration ould transmittd imrndi-
progrss xtndd thir thinking atly to th futur gnrations-Lamark was so nthusd y this pross
.olution-pogrssiv hng up- that th inhritrrnof aquird haratristishas sin om known as
1y thn rad this ida ak into Lamarkism-ut yond this was siln.
4 The istor of uolutior Thought

A DARWlnl
tl8D li llr

l, '!? or

Figur 1. Homologis. Thse re th forelims of (fro lft to right) the at (its

wing), the porpois(itspaddl),th hors (itsleg),and th human (its arm). Th
onesar oviouslysimi[ar,s if modledon a singlrhetyp'and yt th
uss to whih thy ar Put ar ver diffrnt.It was th British anatomlst Figur 2'Prize pig. Al
Rihard own who alld them ..homologis'''lthough it ws not until fter ountrysid and go to
Darwin's on th origi of Spis(1859)that thy amuniversallyaptd on th land to produ
as proof of desntfrom a ommon nestor. rvolution, and at th
mjor ky to suessl
hirir shp,and ig

Th idology of progrss was wht ountd, and it was for this rason that
most popl around 1800 would hav rgardd volutionism lss as a ral
sin and mo as a ptnd,smwhat lik animal magntism (ms- divin industialist a
mrism) and th rading of haratr from skull shap (phrnology). vn of th world througt
!udgd by th standards of that tim, volution was what may fairly alld Critis of voluti
a psudosin. Oviously, hristian opponnts of volution dislikd in- Gorgs uvir, m;
tnsly the antiprovidntial undrpinnings of th dotrin. But volution was mummifid bodis o
not assoiatd with total nonlif, athism, or vn what latr in th nin. from gypt. Althou1
tnth ntury Thomas Hnry Huxly was to all agnostiism. ost volu- tmporary forms an,
tionists wr dists who livd in God as unmovd movr' a ing who had with th positiv at
st th world in motion and now lt it unfurl without nd lf miraulous in- that rally ountd.
trvntion. For th dist' inded, volution was proof of God,s powr and in- hatrd of a dotrin
tntion rathr than disproof. vrything was plannd forhnd and wnt than thy ould d z
into fft through th lws of natur. In rasmus Darwin's words: ..\)hat lution, whn oppon.
magnifint ida of th infinit powr of GRAT ARITT! T us provd tru and saft
oF AUss! PARNT oF PARNTS!Ns NTIU!' (1801' 2: 247|. Thr is a as for vry othr o
link hr, spially in ngland, with th Industrial Rvolution. Popl wr final aus. Th Fr
harnssing th fors of natur-watr, ol' and othrs-to produ goods not ignord; indr
through mahins rathr than y hand. Th god of th dist was th ultimat lif. Th ky to un

os of (from left to right) th at (its
s lg),and th humn (itsarm). Th
r a singlearhtyp,and yet th
l w a sr h B r i i s h n a t o m i s r
s,''althoughit ws not until after
at thy eameunivrsallyaptd
td, and it was for this rasn that
4addevolutionism lss as a ral
hat lik animal magntism (ms-
m skull shap (phrnology). vn
tion was what may fairly b alld
|Ponntsf volution dislikd in-
of th dotrin. But volution was
m' or vn what latr in th nin-
s to all agnostiism.ost volu-
unmovd movr' a ing who had
] without nd of miraulous in-
was proof of God's powr and in-
las plannd forhand and wnt
lrasmus Darwin's words: "What a
GRATARHITT! T us
tu!'' (1801,2: 247).Thr is
ndustrialRvolution. Popl wr
al, and othrs-to produ goods
gd of th dist was th ultimat
The llistor of uolutior Thought 5
A Dlnwl*tA lDA.
t.lfitD lr 1n. .rlt ilor.
l' i!' o l
' tt a' t.l,! t! 'r?ttlx, .!]}',l'
'!! !r1 '\'r.titoi
lt otl t t'.l,t rt l!l!
Figure2. Priz pig. An industrial rvolution means that pople lav th
ountrysidand go t th fatorisin th itis. n, far fwer peopl left
on the land to produ th nation's food. Thr has to an agriultural
revolution,and at th nd of th ightenthntury popl disovrdtht a
major key to suesswas th redingof ttr-qualitylivstok-fattr pigs'
hairirshep,and iggrattl.(From Punh, |865.)
divin industrialist as h harnssd th fors of natur to produ th goods
of th world through law rathr than miraulously or y hand.
itis of volution, notly th grt Frnh omprativ antomist
Gorgs uvir, mad rfrn to mpirial prolms. Cuvir itd th
mummifid odis of humans and animals rought bak y Napolon's army
from gypt. Although thy wr vry old, thir forms wr idntial to n-
tmPorary foms and hn ountd against ongoing rgani hang' But as
with th positiv as for volution' in th ngativ as it was th idology
that rally ountd. Aov all, for uvir, thr was hatrd of progrss-
hatrd of a dotrin that had ld humans to think that thy ould do mor
than thy ould do and had ld ultimatly to th trrors of th Frnh Rvo-
lution, whn opponnts of th isting stat had trid to hang all that had
provd tru and saf for mny gnrations. omind with this, for Cuvir,
as for vry othr opponnt of volution, was th still-nsolvd problm of
final aus. Th Fnhman mphasized that this was somthing that ould
not ignord; indd, it was th most important distinguishing fatur of
lif. Th ky t undrstanding th organism lay in th ft that it was not
 Th Histor of uolutior Thought

simply sujt to th physial laws of natur ut was organizd. Th parts in human natur, t
wr dirtd to th nd of th funtioning whol, and ah individual fa. Now all of this is i
tur playd its rol in th ovrall, purpos-dirtd shm of things: of organi ration
th hour hand is t
Natural history nvrthlss has a rational prinipl that is lusiv to
w do of that latt
it and whih it mploys with grat advantag on many oasions; it is
rtrogrssion of th.
the oditios of eistee or' popular|,
fil uss. As nothing may missil, but a forl
ist whih dos not inlud th onditions whih mad its xlstn
rs 1846, 40040
possibl, th diffrnt parts of ah ratur must oordinatd in suh
a Way as to mak possil th whol organism, not only in itslf ut in
On th oth sid
its rlationship to thos whih surround it, and th analysis of ths on-
aout th signifian
ditions often lads to gnral laws s wll foundd as thos of alula-
modifid ut thinkin1
tion or xprimnt. (uvir 1817' 1: 6\
ral man of Sottish
uvir's point simply was that th organism is far too intgratd-organizd grs: ..It would augl
and ompl-to allow signifiant hang in any dirtion. It is rtainly too wr inftd with th
intgratd to allow hang from on spis to anothr. organisms at mid. taintd with Matria
point would litrally nithr fish nor fowl and hn would simply un- rfltions that still fi
a]to xist or surviv. volution was in som snsa thortial impossiility, th slaknss of sho
as wll as mpirially unfoundd. diating th unsoun<
Rligion was involvd too. Progrss gos against th hristin dotrin of startd into puli fa.
Providn. Nvrthlss, although Cuvir thought that thr was vidn n, and of sapping
of Noh's flood, nithr h nor othr serious sintists wantd to mak th trstingly' this kind
as by simpl rfrn to Gnsis. Indd, y th ginning of th nin- viws. Although, fro
tnth ntury' all wr starting to ralize that th rth's history must far flt th nd to makr
oldr than th traditional 6,000 yars that on an work out from th g- aov is from th fifl
nalogisgivn in th Bil. It is not that th Bil is fals, ut rathr that it mor rdit to th Cr
nds intrprtation. Som solved th prolm y thinking of th si days of of ours, nithr
ration as si long priods of tim; othrs solvd it y supposing that thr ral viden, and fin
wr long, unmntiond gaps twn th ilial days. God's ration thre- all to miral may
for was a long, drawn-ut pross' ut it was not volutionay. good sintifi soluti<
Th ontrovrsywas at an impass,and not muh had hangd y th mid-
dl of th nintnth entury. on the on sid wr th volutionists' om-
mittd to progrss and ardnt in thir lif that organi dvlopmnt was Charls Darwin
th perft omplmnt to this idology, with nthusiasm outstripping mpir-
iai knowldg. onfirming this pattrn' ]n 1844 th Sottish pulishr harls Robrt Darl
Robrt hamrs wrot (anonymously)a highly popular work on volution, train in th family
Vstiges of tb Ntrl Histor of rtio, in whih h argud that vry- droppd out, ord r
thing was in a stat f upward oming and that wht happnd in th so- rady Darwin had st
ial world mirrord what happnd in th iologial world: std in th living w
anatomist and an -v
Th qustion whthr th human ra will vr advane far yond its work (th young h'
prsnt position in intllt and morals, is on whih has ngagdmuh tion was an ida to r
attntion. Judging from th past' w annot rasnaly dout that grat nvrthlss, that thl
advans ar yt to mad; ut if th prinipl of dvlopmnt ad- whol of hristianit;
mittd' ths ar rtain, whatvr may th spa of tim rquird for was a fator in his r
thir ralization. A progrssion rsmling dvlopmnt may trad th stalishd Chur'

Lturut was organizd.Th parts
ng whol, and ah individual fa-
'-dirtdshmof things:
lnal prinipl that is xlusiv to
/antagon may osions;it is
|, fil uss.As nothing may
litions whih mad its istn
tur must b oordinatd in suh
ganism'not only in itslf ut in
Lit, and th analysisof thson-
vll foundd as thos of alula-
;m is fr too intgrtd-organizd
, in any dirtion. It is rtainly
to
;isto anothr. organisms at mid-
lwl and hn would simply un-
m snsa thotialimpossiility,
s aginst th Christian dotrin of
lr thought that thr was vidn
ious sintists wantd to mak th
d, y th ginning of th nin-
tht th rth history must far
at on an work out from th g-
th Bil is fals' ut rthr that it
lm y thinking of th si days of
.s solvd it y
supposing that thr
ilial days. God's ration thr-
Was not volutionary.
not muh hd hangd y th mid-
: s i d w r t h v o l u t i o n i s t s' o m -
lif that ogani dvlopmnt was
i t h n t h u s i s mo u t s t r i p p i r r g m p i r -
,, in \844 th Sottish pulishr
highly popular work on volution,
lz, in whih h argud that vy-
and that what happnd in th so-
iologial worid:
will vr advan fa eyond its
is on whih has ngagd muh
Lnotrasonalydoubt that grat
prinipl of dvlopmnt ad-
th spa of tim rquird for
ing dvlopmnt may trad
b Histor of uolutior Thought
in human natur' oth in th individual and in larg grouPs of mn . . .
Now all of this is in onformity with what w hav sn of th progrss
of orgai ration. It sms ut th minut hand of a wath, of whih
th ho hand is th transition from spis to spiS. nowing what
w do of that lattr transition' th possiilit of a didd and gnral
rtrogrssionof th highst spis towards a manr typ is sa ad-
missil, ut a forward movmnt sms anything ut unlikly. (Cham-
s 1846.400402|
on th thr sid wr th opponnts' ommittd to Providn, vol
aout th signifian of final aus' and apting that Gnsis must
modifid but thinking that this ould radily don. David Brwster' gn-
ral man of Sottish sin and iographr of Nwton, ould s th dan.
grs: ..It would augr ill for th rising gnration, if th mothrs of ngland
wr inftd with th rrors of Phnology: it would augr Wors wr thy
taintdwith atrialism.'' Th prolm, Brwstr gloomily onludd with
rfltionsthat still find muh support in many irls today, stmmd from
th slaknss of shools and univrsitis: ..Prophti of infidl times, and in-
diating th unsoundnss of our gnral duation, .Th Vstigs . . .' has
statdinto puli favour with a fair han of poisoning the fountains of si-
n)and of sapping th foundations of rligion', (BrWstr1844' 503). (In-
trstingly, this kind of attak sms to hav onfirmd hamers in his
viws. Although, from th start, progrss was th litmotif of his ook' h
flt th nd to mak his point vr mor xpliit. Th passag just quotd
av is from th fifth dition of Vstigs and rplas a passag that givs
mor rdit to th rator. [Figur 3])
of ours, nithr sid was ally satisfatory. Idology is no substitut for
ral vidn, and final aus an b ignord ut dos not go away. Rfring
all to mial may soially and psyhologially omforting, ut it is not a
good sintifi solution. Th tim had om for a signifiant stP forward.
Charls Darwin
hls Robrt Darwin (1809_1882) was snt to dinburgh Univrsity to
train in th family tradition of mdiin (Figur 4). Aftr two yars h
droppd out, ord with th lturs and voltd y th oprations. Yt al-
ady Darwin had startd to mi with sintists, spially naturalists intr-
stdin th living world. On of his aquaintanswas Rort Grant, an
anatomist and an avowd volutionist. So, quit apart from his grandfathr's
wok (th young harls rad rasmus's major tratise,Zooomia), volu-
tion was an ida to whih Darwin was introdud at an arly ag. It sms'
nevrthlss,that th youthful Darwin aptd in a fairly litral form th
whl of hristianity, inluding th arly haptrs of Gnsis, and that this
was a fator in his rdirtd hoi of arr: to an ordaind ministr in
th stablishdhurh of ngland. To ahiv this nd on ndd a dgr
8 The Histor of uolutionar Tbought

Figur 3. hamrs'svolution.This is the piturof volutiontht hmrs

gve in Vestiges of th Nturl Histor of retio. Apparntly' for no
apparnt aus' a primitiv organism wnt on dvloping in th wom nd
tuned into a fish; thn a fish Went on dvlopingand turnd intl a rptil;thn
a ird; and finally a mammal. Note th intertwining of thinking abouj the
history of th individual and the history of th goup, somthi that ws to
om ommonpla in evolr'tionarythinking latr in the nininthntury.
(From Vestigs of the NturI Histor of retio.|

from an nglish univrsity, and so, in 1828, Darwin was pakd off to
hrist's Collg, amridg.
In 1831 Darwin (who ontinud mixing with sintists)got his ig rak.
Aftr h graduatd, his arr as a lrgyman Was put on hold through th of-
fr of a lngthy voyag on HS Begle,just about to start on a survying trip
around South Ameria (Figur 5). A major influnt this point was (viar- Figur 4. harls Darl
iously) th Sottish gologist Charls Lyll, who at th eginning of th d- 1 8 3 9 j u s t f o r h e m l
ad egan pulishing his massiv Priipls of Golr>g(thr wr thr liknss(no surpris's
volums; Darwin took th first with him and had th othrs snt out). AI- portraitist in ngland
though h was no volutionist, Lyll insistd that th physial world must from a very rih, midd
.Wd
xplaind in trms of natural auss of a kind now still working (Figur 6). wif' was Josia
mass prodution.Alth
This had a great fft on Darwin, whos first systmti work was in gol-
wok for a living. Th
ogy, nd prpad him to think aout th ogani world likwis lss in bi-
plains w n th |
lial trms and mor in trms of natural auses. In 1835, laving South frindly fashion. Dar
Amria, the Begl saild into th Paifi oan and visitd th Galipagos Wdgwood family.
Arhiplgo' a 8rop of islands on th quator, far from land. Thanks to th
governor' who pointd out that th giant tortoiss indignous to th arhi-
plago wr diffrent from island to island, Darwin am to s that this hld plaind was throu[
for th Galipagos fauna gnrally-th irds in partiular, th finhs and radin arly 1837
th mokingirds, wr puliar to thir spifi homs (Figur 7). Th dif- li thinkrs, what w
frns had to b signifiant, and th only way th signifian ould .Without
volution.
 The Histor of uolutior Thought 9

-/B

---B

_---F

litu of volution that hamers

"etio. Apparntly, for no
,n devloping in th wol and
rping and turned into a eptil; thn
twining of thinking out th
h group, something that was to
'ng latr in the nintenth nturY.
tio.)

[828, Darwin was pakd off to

;with sintists)got his ig rak.

an was put on hold through th of-
;tbour to start on survying rrip
.influn
at this point ws (viar- Figur4. halsDarwirr'This is wddingportrait of Drwin, paintdin
l, who at th ginning of th d- 1839just forh marridhis first ousin mma.Wedgwood.It is a vry good
)les of ceolog (thr wr thre liknss(no surpris,sinth paintrwas GorgRihmond, th st
and had th othrs snt out). Al- portraitistin ngland at th tir),whih points to th fat that Darwin am
ld that th physial world must from a vryrih, middl-iirss family. On of his grandfathrs,shardy his
<indnow still working (Figur 6). wif, ws Josih Wdgwood, rsponsilfor ringing th pottry trad lnto
massprodution.Although h was a full-timsintist,Darwin nvrad to
first systmati work was in gol-
work for a living. Th fat that h was living off th family fortun prhaps
gani world likwis lss in bi- eplainsw tbe rigitt of Spiswas writtn in a vry opn nd user-
l auss. In 1835, laving South friendlyfshion.Darwin ws witing ir-rprt for his sponsors,th Darwin-
.!ildgwood
oan and visitd th Galipagos family.
ltor' far from land. Thanks to th
tortoiss indignous to th arhi-
Darwin am to s that this hld plaindws through ..dsntwith modifiation,',an ida that Darwin m-
rds in partiular, th finhs and bad in arly 187, shortl after th Bgl voyag (Figur 8). Unlik ar-
ifi homs (Figur 7). The dif- lir thinkrs, wht was of grat onrn to Darwin was th aus or auss of
ly way th signifian ould volution.\/ithoutauss'h ws no mor than on among many volutionists.
10 The Histor of uolutior bought

iii,i:.:.'.:' P
-i

AN ATTE1I

Figur 5. }{S Begle.The Bgle was aptind y Rort Fitzroy, thn only
2 earsold. Bing a aptainon warship would a very lonly Post aus
on would st asid from th rw. Fitzroy thforlookd for somon
who ould b his ompnionon th trip, somonnot in the navy ut II
gntlmanabl to pay his own bills. Darwin fit th jo prftly,rapidly
volving from aptin's frind to full-tim ship's naturalist.Th rw usd to all
him ..Philos,''short for Natural Philosophr_the trm thn usd for sintists.

.sithout
allss' h ould nvr th Nwton of iology. Darwin wantd to
find a biologial for that would explain th volution of lif and was uni-
vrsal in th world and th ky hind all organi motions' and to this nd h
workd fvrishlyfor th nxt 18 months, spulatingnd ding an tnsiv
litratur rviw.
Darwin soon ralizd that th proal ky to hang ly in somthing
paralll to th way in whih animal and plant rdrsfft hang, namly,
th sltin of th dsil and th rjrion of all othrs. Brrt how was oD
to gt a natural quivalnt to th rdrs' artifiial sltion? At th nd of JOII]
Sptmr 1838 Darwin rad n ss on th Prinipl of Popltioz (sixth
dition, 1826| th Rvrnd Thomas Rort Malthus. In his work
althus argud that food supplis would always b outstrippd y potntial Figur 6. Priniples o
population growth. Hn' unlss thr was ..prudntial rstraint,'' thr his mthodolgialin
wr bound to ongoing struggles for xistn as popl omptd for th
availl food and supplis,inluding living spa (Figur 9). Darwin turnd
althus's rasoning on its had. H pointd out that thr ould no
prudntial rstraint in th animl and plant wrld' and hn,bauspopu-
lation growth rtainly Was not rstritd to humankind, thr would an
ongoing' organi-wid struggl for xistn. or than this, suss and
failur in th struggl would (on avrag) a funtion of th diffrnt
 Tbe Histor of uolutior Thought 11

PRINCIPLBS

G o Lo GY,
ElIiG

AN ATlP To EPLAIN TII FORMR IIANGES

oF II DARII'S sURFAc'

Y REaEBNoE o Auss Now lN oPEBAloN.

lind y Rort Fitzroy' thn only

lould a vry lonly post eause
y thrforlookd for someon
nonnot in the nvy ut a IIARLEs LYLL, s Q., F.R.s.
fit th jo prftly,rapidly FOe. 9J8C. TO TflD QEOI" SOC' !c,
ip's naturalist.Th rw usd to all
--th rrmthn usdfor sintists.

lton of biology. Darwin wantd to IN wo vol.us.

th volution of ]if and was uni-
Yol. I.
'rgani motions' and to this nd h
; p u l a t i n agn d d o i n g a t n s i v

: ky to hang lay in somthing

Lntrdrsfft hang, namly,
ion of all othrs. But how was on LONDON:
'
artifiial sltion? At th nd of JoIIN lvIURRAY' ALBE},IARLE.SRET.
tb Priiple of opultioz (sixth
s Robrt Malthus. In his work "o**.
rlways b outstrippd y potntial Figur6. Principles of Golog' The full titl of Lyell's work tells you all aout
was ..prudntial rstraint,', thr his mthodologialintntions. ventually, thr wre thr volumes.
stnas popl omptd for th
8 spa (Figur 9). Darwin turnd
ntd out that thr ould no
: world, and hn, aus popu-
o humankind, thr would an
n. or than this, suss and
:) a funtion of th diffrnt
12 be Histor' of uolutior Thougbt

Abingdon
\
indIoo
owel
A|bemr|o e hr south meri tofuis
fudo dbingdonii shldo bul
fiom Abidon l5lnd

fudo vi.in
fom Almr| |sland

e*udo ephiwiu
from Dunn lnd

Figur 7. Th Gal;ipagostortoiss.Darwin did not omean volutionistuntil

h rtrrndhom and h ould show his olltionof irds to John Gould' a
ladingornithologist.It was whn h ws told that threwr unmiguously
diffrnt spistt he movd ross to volution. Not that vn efor h
aman volutionist,Darwin was moving in irlswhr h ould all on
th tp minds fo supPort and informatio. He mad powrful frinds t
amridgand thy spottdhis talntarly.

Figur 8. Th tr of lif. This littl skth,drawn inJuly 187, is from a private F i g u r 9 . T h a l t h s

notbook that Drwin was kping on th sPisprolm. Lmark had tr- l m ki n 1 8 5 l . o r
of-[ifpiturand in th ightnthnturyon an find thm in th writings f tl{ th truth of voltrti
nonvolutionistswho wr nvthlsstrying to shorth odr in whih God on idas tlrat wr wi
hd rtd.
 h istor of uoltionr Thougbt 1

on

BindIo
Iow l
t^
\,J he south merin toise
studobulfu

s
GALAPAGOS
ISLANDS
l..s-y
J

| > Indefstigable
\*-/ / J /
/'2
.rigton <-'/,/ h.ath.am

harIes

/> ood

lid nt oman volutionistuntil

ltionf irds to John Gould, a
ld that thrwre unamiguously
lution. Not that vn fr h
in ir|eswhre h ould all on
H made powrful frinds at

rawn in July 1837' is from a privat Figur 9. Th althusian xpltlsion. This arton appard in the omi
lisprolem.Lamark had a tr. lmk in 1851. On of tlr rasons tht l)rwin was al to onvin popl
)nan find thm in th writings of of th truth of volrrtion Was' apart frtlrn lris frindly styl, th fat that h rlid
g to show th ordr in whih God on idas tlrt wr widlv rltd in Vitorian Britirin.
I+ Th Histor of uolutionr Thought

hartistispossssdv organisms. n tl-rrwould b an ongoing

natural sltion in whih th winnrs would pss on faturs to th nt gn-
rtion and th losrs would gt nothing. Ovr tim' this sltion would lad
to full-lown organi hang. But this organi hang would of a partiular
kind' for th faturs that mk for suss,known as ..adaptti6n5''-f61 g;- . Rut with rgerd to th mlbil wold,
vnts sre bug
*

fr 8 tiFwc lrive thot

ampl, ars' ys' tth, hands, lgs, lavs, rk, and roots-would show that in6utail iokrl6itio!! of Dine
oi
Iw . , ex l kd
gut iw.''
ir

8, ut by thc t,Iiment
purposful natur that pitoizs final irus. \Y. Wvrl : Ba

Darwin sat on his volutionary idas for 15 yars' during whih tim h
's7' ..l. l t no lt of s
turnd to a massiv study of arnals. ar not quit sur why this dlay nclud, thrcfor'
ruritY. or ill.aPli mdl"tioD, thiDk or:
vll sfudi.'d io t
ourrd, although y this tim Darwin hd falln sik with a mystriousail. ..*o.u'' .h @ lar o i

wo.t, or i the l} of God's orks; divinity or

or pr
mnt that W21sto plagu him for th rst of his lif, and so undoutdly h rAtI lot ruD cldvour n ri.lles gr$

Bo; .4dvlre

Was not rlishing th hug dattlrat his idas wr ound to aus. Also a
major fator must hav n his rlutan to upst powful sin-
stalishmntfigurs (inluding mridg mntors) who had nouragd
th young Darwin in his work. On of th things h did during th paus was
to ntwrk with youngr sinrists'who ould rally around him whn h did
go publi. Finally, howvr, Dzrrwin was pushd into ation whn, in th
.Walla, Do' Bfolq, K|'
middl of 1858, h rivd short ssay y Alfrd Russl a oll- o.ros' l!l' 1859.

tor in th Malay pninsula' that had virtually th sam prmiss and onlu-
sion that h had disovrdsom 20 yars arlir.

oN oRlGlN oF sPls

Darwin quikly wrot up his idas, and ()n th rigin of Spies b es Figur 10. Th titl pag
of Ntrl Seltion, or th Preserutio of Fulred Res i th Struggle hky in quoting th p
for Lif apprd in th fall of 1859 (Figur 10). Th work ginswith a dis- th topi of th law-gor
ussion of artifiial sltion rrd animal and plant rdrs'grat susss. Drwin's mntorswhr
This is partly huristi,t prpar th rdr by following th path tht Dar. ontinud in this r<llir
win himslf had takn to disovry, and partly justifiatory,suggstingthat known opponnt of v
Darwin snt a omplir
what w an do, natur an surly do ttr.Following is a disussion of th
y anning the ook fr
variation tht lways ours in th wild, somthing that Darwin oviously
h ws astr (had).
ndd as th building loks of hang. If vry organism Wr xatly lik
its parnts, thn thr ould no volution' At this point Dawirr was rady
to introdu th strugglfor istennd' following on this, th mhanisnr
of natural sltion: Not that vn mor
for rprodution. lt is
A struggl for istn invitaly follows fr<lmth high rat t whih ll sxal dsirsof a phi
organi bings tnd to inrils. vry ing, whih during its natural way' givn naturally
liftim produs svral ggs or sds,must suffr dstrution during
som piod of its lif, and during som sason or oasionl yar' oth- Canit...thou1
rwise, on th prinipl of geomtrial inras, its nurrrl-lrswould hav undoutdl1' .
quikly om so inordinatly grat that no ountry ould support th ah bing in th g
produt. Hn, as mor individuals ar produd than an possily sur- ur in th ours o
viv, thr mllst in vry as a struggl ftlr istn,ithr on indi- doubt (rmmbri
vidual with an()tof tlr sm spis,or lvith th individuirls of distint sily surviv) that
spis,or with th physil onditions of lif. (. Dawin 1859' 63) ovr othrs' would
 Tbe istor of uolutior Thought 15

. n thr would b an ongoing

ould pass on fatursto th nxt gn-
;. ovr tim, this sltion would lad T 0RIGIN 0F SPcIs
gani hangwould of a partiular
ss,knwn as ..adaptation5''-f91 91- ..ut with rcgst.l to th mbrjal wold, w t least go o BY }lE^Ns0 NATURAL8EL0TI0N'
flr N this*we privr that eveDt ale bougt aout not by
s' bark, and roots-Would show that iutd iDtrp6itios of Die wer, eerhd in sh artiulsl
e, ut by t 6blishmrnt o[ gonl lsw.''
:aus. w. wEvlLL : nffitr Trti*.

for 15 yrs, during whih tim h PltEsIivAloN o!' FAVOURD RAcs rN T TIiUGGL0
! ar not quit sur why this dlay ..'l nlud' thefe{orc, lt no msn out
thik
of o wk
o mritain'
oncit
ttt
of
t
F'o'r LJ .
liety, o ill-spplied modati0n,

had falln sik with a mystrious ail- lrrn u sr fu f o s t ell fudi d in the mk of God's
worJ, o in thr k f Gorl's ltork; diviniiy or pllilsolit; ut

st of his lif' and so undoutdly h rth0r let meD eDdvoul nn ldles grss r ploficine in both.''

| d'turnt o! Iain(]'
Lisidas wr ound to as.Also a
B^
Ry OHARLDS DR1r!TN' 1{'A.,
-ltan to upst powrful sin- lL o trE
lJoURN^l
Y^L' GEoIDGI^L' LlNNt^n, .' sor8;

^UR o! o i8Dt DlG . . . ErGl voYto

idg mntors) who had nouragd 0ND weD'.

t things h did during th paus was

ould rally around him whn h did
as pushd into ation whn, in th
y y Alfrd Russl
.salla,
a oll-
o"*'"'r,i*Iil'',,,,.
:uallyth sam prmissand onlu-
rs arlir. "*i:}^""' *TRET
,o,*,o,'u*oo'l

.ticfu@n

o the rigi of Speies b ens

n of Fuourd Res i the Struggle
ur 10). Th work ginswith a dis-
and plant rdrs' grat susss.
adr y following th path that Dar-
I partly justifiatory, suggsting that
ttr. Following is a disussion of th
l, somthing that Darwin oviously
If vry organism Wr atly lik
ion. At this point Darwin was rady
Ld,following on this' th mhanism
Figur 10. Th titl pag of n tb rigin of Spis.Darwin ws ing a littl
Whwll, oppositth titl pag' on
hkyin quotingth philosophr.!illiam
.!hwll
th topi of th law-govrndnatur of th univrs. had n on of
Darwin'smntoswhn th lattrWas a studntat mridg,nd h
ontinudin this ro1in th arly yars aftr te Bgl voyag.Ll was a wll-
known opponntof volutionand was sathingon th sujtof Vstigs'
Darwin snt a omplimntary opy of the rigi to.Whwll, who rspondd
by anningth ook from th shlvsof th lirry of Trinity ollg' of whih
h was Mastr (had)

Not that vn mor than a struggl for xistn,Darwin ndd a struggl

lws from th high rat at whih all
y ing' whih during its natural
s' must suff dstrution during
1 sason or osionl yar' oth-
al inras, its numrs would
hat no ountry ould supprt th
produd than an possily sur.
ggl for xistn,ithr on indi-
, or with th individuals of distint
of lif. (. Darwin 1859' 6)
for rprodution. It is no good having th physiqu of Tarzan if you hav th
sxualdsirsof a philosophr. But with th struggl undrstood in this sort of
Way' givn naturally ourring variain, natural sltion follows at on:
an it . . . thought improal' sing that variations usful to man
hav undoutdly ourrd, that othr vriations usful in som way to
ah ing in th grat and ompl attl of lif, should somtimso-
ur in th ours of thousands of gnrations?If suh do our' an w
dout (rmmringthat many mor individuals ar born than an pos-
sibly surviv) that individuals having any advantag' howvr slight,
ovr othrs, wuld hav th st han of surviving nd of prorating
tb The Histor of uolutior brgbt

tlrir kind? On tlr othr hnd W Iyfl sur that any r,rizrtionin th ltion partiularly pr
last dg injurious worrld rigidly dstrlyd.This prsrvatiorrof th sam spis:sl
favouralvariations and th rjtionof ilrjurious variations,I all Nt. istis rthr thn sp
u r l S l t i t l n(. 8 0 _ 8 1 ) s sltion (inludin
aving thus prs
In latr ditilrrsDawin intodtrd th ltrrtivtrm gtruiul of th fittest, dud th famous m
wlrih was prhps a littl unfotulrat,for it has ld to ndlss laims that s- sam lass hav soml
ltion is a tutollgy, rduing simply to th lairn tht thos who surviv ar il largly spaks th
thos who surviv. But althouglr this is oviously tru, sltionmans mor spis xtant today.
than tlrat. It laims, ior ttr lr flr wors' th21ton vrag tlros who surviv volutionary paths of
a diffrnt frr-nthos whtl do not, and thilt sussin tlr struggl is a fun- vry first shard orig
tiorr of thos diffrns.This may ()r may not tru, ut it is not a tautology. and ths, if vigorou
Darwin inludd a sondary sltiv mhanism. H argud that th ranh, so y gnrz
stugglis not always for food irnd spa ut an dirtly fo mats. This whih fills with its d
mhanisrn, stll seleti>tl, Darwirr dividd into twtl Prts. Thr was s- rs th surfa with i
ltion through lal omat (flrf;rrals). Tlr antlrs of th dr would win 1859, 129-130)
th produt of this. hn thr was slti<lnthrugh fmal hoi (of th Now, with som -
most dsiral m:rls).Th tail fathrs lr gnral plumg of th paok prsnt th sond
-l
or sirrrilarird wol] pr<ldr"rt lf this (Figtrr 1). Noti hrwsual s- Darwin took th ra<
instint, palontology
and showd that ph
through natural sl
support th mhan

Th anlogywith anirnalrdingld Drwin ro

Figur 11. Sxual slti<ln.
distinguishsualsltilnfrlmntur1sltirln.Nturl sltion
orrspondstl sltionfor tti[.'uts
tht hlp sttrviva[,lik tlrikroats,
whassulsltiOno[rsP()nds frlr attri[ruts
to sltirlrr that giv pla.
suto th rd"rs,lik prttirfathrsr mOfroiouslightingqulitis.
This pituris takn |romTb I)stttof . (Busthe rigi ws writtn Figur 12. Th tr <
in a hurry, thrw:rsnl tim for fotnltingor il[ustrting.Most of Darwin's this piture is lss or
othr rvorksar riLrllyfootnltdnd opiouslr'illustratd.) sprads.

fl sur that any variation in th
r"dstroyd.This prsrvation of
of injuriousvaritions'I all Nat-
trnativ tJm suruiul lf the fittest,
,r it has ld to ndlsslairns that s-
l i n lt h t t h r l s w h o s t t r v i v
bviously tru, sltion mns mor
' that on avragthos who surviv
tht strssin th strtrgglis fun-
not tru' ut it is not a tautology'
mhanism. argud that th
bllt an dlrtly for mts. This
idd into two parts. Th was s_
;). Th antlrsof th dr would
:tion through fr-nal hoi (of th
or gnral plurr-rag of th paok
; (Figur11). Noti how sual s-
h animalrdingld Drr,vrnto
:tion.Naturl sltion
rlpsurvival,lik thikroars,
tion fr ttriutsthat giv pla-
mor foiousfightingqulitis.
[a.(Bustl.t()rigin ws writtn
or illustrating.Most of Darwin's
iouslyillrrstratd.)
Th Histor o|.uolutillzrlT1',,,''' 17
ltion partiulalyputs th mphasis on omptition btwn mmrs of
th samspis:sltionis dirtd toward individual-nfitingharatr-
istisrathr than spis-nfiting haratristis.Darwin tndd always to
ssltion (inluding natural sltion)ating in this way.
Having thus psntdth min Inhanisms of hang, Darwin intro.
dudth famous mtaphor of a tr: ..Th affinitis of all th ings of th
samlasshav somtimsn rprsntdy a grat tr. I livthis sim-
il largly spaks th truth.'' Th lavs and twigs at th top reprsnt th
spisxtant today. Thn, as w go down th branhs' w hav th grat
volutionary pths of ystrday. All th way down w go until w rh th
.,As
vry fist shard origins of lif. uds giv ris y growth to frs uds,
and ths,if vigorous, ranh out and ovtop on ll sids many a fblr
ranh, so y gnration I liv it has n with th grat Tr of Lif,
whih fills with its dad and rokrrranhsth rust of th th, and ov-
rs th surfa with its vr branhing and autiful ramifiations'' (. Dar-
win 1859, 129_130) (Figu 12).
Now, with sol minor problms brr.rshdaway, Drrvin was rady to
prsnt th sond part f his thory. For a good two-thirds of th rigi,
Darwin took th radr through th vaious ranhs of iologial sin-
instint,palontology,iogography,lassifition,m<lrphology,mrvol.,gy-
and showd tht phnornna in ths ranhs ar plained y volution
thrughnatural sltion,and, onvrsly,thsvarious branhs point to and
support th mhanisnr of volrrtion through sltion (Figur 13). hr
Figur 12. Th tof lif s given in t rigi. Noti that th mphasis in
this pituris lss on showing lif rahing up and mor on th way in whih it
spreds'
18 The Histor of uolutilrl Thought

would all th nr
p l a i n ds i t n l s e
n o l o n g r t h r , r .
rought out all of D
whr now th Wrr
not find it. vn if w
dotrtftll that w wou
prssd and mtamor1
o s t o t r r a so f

-l -l _l -l l
trir-rtion(iogograph
is that on finds th va

L__ltt ll tt II tl
Why, fo instan,do

L__lL__lII L__l
that ar xhiitd y tl

lt
lnstint Paleontology cographial lssifiation orphology mryology
simply that th found
th mainlnd, nd on.
sify undr th nw s
Distribution
Figur 7). Systmatis
Figur 1.3.Th struturof Darwin's rgumnt.l)arwin did not hit y hanon volution. As Linnau
this kind of argumnttion,whr a unifyir-rg hyplthesisplainsin many ally in nstdsts,thu
'Whwll,
diffrntarasand in turn is justifidor suportdl.thsras. who opology lrrsthos
alld it a ..onsilin
of indutilns,''mad muh of it whn disr.rssing
diffrnt spis.ovi.
sussfulthtlrisof physis,spiall1' Nwttlnianmlrnisand th wav
thory of light.Whwll idntifidth irussat th ntrof onsilirr ryology likwis was .
s
..truauss,''wht Nwttln hd alld Urd(l|ts. Showinga onsilinwas mbryos of sor diff-
thrforn ssntilstfatgyfor worrld-bNwtlt-lof iology. whras th adults r
fat tht in th wonr
simila (thywould nor
th rwo adults would l
Wr som prolmati isstts.Soial hlvior, prtitrllyth:rt shown by Hr' as lways thou
th soial insts (th hymnoptra-th ants' th s, and th wasps), analogy with th worl
would asily plial if sltion favlrdth group. But frrm an indi- ..Fanirsslt
hand:
vidualisti prsptir,,why slror"rldol1 gt stril lr,orkrs lvlro smirrgly ar nrly 8rown up:
spnd thir whol livs dvotdto th lvll-ingof othrs irl th nst or hiv? struturs lrv n a
How an natural sltion,with its mphasis on th importan of rprodu- posssssthm'' (. D'
tion, rodu organisms that an<rtrprodu? }.inirlly,Darwin' wo al- So w ar ld to tt
ways favord individulisti pproahs ovr grLlp ppr(lhsaus h hristianity lrd volr
thought th lattr too opn to hting and ploitation, didd tht in suh mmrs of th \/d
ass th group mmrs ar so ll,ll intgatdthirt it is prmissilto trat ld that h Was W
th ll,hol hiv as kind of suprorganismin lr,hith individual instsar unslf-onsiousrfr
parts of th whol. Just as sltiltltr work oll th v' for ampl, whih sm to fully satis
xists for th nfitof th whol rganism, so th workr ists for th n- dntly ratd.''This r
fit of th whol hiv. tr with what w kno
Palontology also raisd diifiultis. It lrad its good points-for instan, th prodution and
that s W go down tlr rorl,inrasingly w fin1orgnisrs that srnto shuld hv n du
hav faturs midwy twn fatr-rrsof xtrrt <lrganismsthat ar widly and dath of th indi
diffrnt.owvr, against th positiv, tr wr gps in th fossil rord, Wht God dos not d
and' vn wors' th fossil rord gan atlptl-vt t start of whr w nss in supposing oth

-l
L_l
__lL__l
]lassifition Morphology
lnt. Dwin did not hit y han on
lg hyplthsisl:rins in many
portd by ths :rrzrs.Whwll, who
n l t l h r l f i t r rh n d i s u s s i n g
:wtonin mhnis and th wav
s at th tr of a onsilin as
us. Showing a onsilin was
Nwton of iology.
,havior,partir-rlrlvthat shown y
t ants, th s, and th wsps)'
vord th group. But from an indi-
gt stril ,orks who smingly
l-ingof othrsirr th nst or hiv?
asis on th importanof rprodu-
lrodu? Finally, Drwin, who al-
ovr group approahs aus h
Ldploitation,diddthat in suh
gratdthat it is prnrissilto trat
r in whih th individalinstsar
lork on th v, ft>rampl, whih
m' so th u.orkr ists fr th n-
had its good points-for instan,
gly w find orgarrismsthat sm to
)f tant organisrs that ar widly
.hrwr gaps irr th fossil rord,
r aruptly at th star of what w
h istor of ulltilrThogbt 19
would ll th zrnrrin;thr Was l1()r()rdfor that' Th gaps Darwin
plairrdsirlpl as a produt of ilrtlmpltfrrssilization.T.hrmains might
no longr thr,ut th linking anstorsdid xist. Th arupt ginning
rought out all of Darwin's invntiv Powrs. arly lif proaly xistd
whrnow thwr oans;this would xplain why w on dry land ould
not find it. vn if w wr al to drill nathth oans, howvr, it was
doutfulthat . would find lif; th prssur from abov had surly om-
prssdand mtirmrphosdth fossil rmains lrlow (Figur 14).
ost oth ars of iology fll int<-l pla witlr mor s.Gographi dis-
l
triution(iogogrplry) rvas triumph ttsD:rrwin plaind iust why it
is tht on fin]sth variolts pattrns of arrirland plant lif aourrd th glo.
dos on hav th strng sorts of distriutions and pattrns
!hy, for instt-l,
that ar hiitd y th Gal5pagos Arhiplago and othr island groups? It is
simply that th ftrundrsof ths isolatd islirnd dnizns am y han from
th mainland, alld on thy wr stalishl,thy startd to volv and divr-
mryoIogy sify undr th nw sltiv prssurs to wlrih thy wr now sujt (s
Figur 7). Systrirtislikwis hiits th kind of pattrns on pts from
volution. As Lirrrrattsdmonstratd, organisnrs an b lassifid hirarhi-
allyin nstdsts,tlrusslrowinga pttnthat vlspast history (Figr 15).
Morpholog1,hirs thos similaritis,lld lromologis,btwn orgnisms of
diffrntspis.oviously thsspk to slrardanstry(sFigur 1). m-
ryologylikr,l,isWas a partiular point of ridfor Drwin. Why is it tlrt th
mryosof sotldiffrnt spis,suh as man and th dog, ar vry similar,
whrasth adults ar vry diffnt? Darwin argud tht this follows frorn th
fat tht in th wom th sltiv fors on th two mryos would b vry
similar (thywlr"rldnot thrfor torn apart)' whras th sltivfors on
th rwo adults wold vry diffrnt(thy wor"rld torn apart) (Figur 16).
r, as always tlrrouglror-rt Darwin turnd to th
his disussions lf vllr-trion,
anabg,vr,r,ithtlr world of rdrsin tlrdr to larify and support tlr point t
hand: ..Fiu-rirs sltthir horss, dogs, rld pigons, for rding' wlrn thy
ar narly gro\vn up: thy ar indiffrrrt rvhthr th dsird qualitis nd
struturshav bn aquird arlir or lat in lif, if th full-grown animal
posssss thm', (. Darwin 1859,446).
So w ar ld to th onluding passgsof the rigi. Darwin, whos
hristianityhad r,olvdinto a form of disrn (Lyll, a Unitarin lik many
mmrsof th Wlgwood family' Was mjor influn hr), nvr on'
ald that h w,:rsr,vlrkingin a God_kl mod, and thr ar frqunt
unslf-onsiot'ts frnsto th rttl' ..Authors of th lrighstnrinn
smto fulh. satisfidwith th virv tlrlrt ah spishas n indpn-
,.To my mind it
dntlyratl'''This was not Darwin's ptlsition. aords t-
tr with wht w know of th laws irnprssdon mattr y th rator' that
thprodutionalrd tintion of th past and prsntinhaitantsof th world
should hav n du to sondary auss' lik thos dtrmining th birth
and dath of th irrdividual'' (. )rwin 18's9' 488). God aks th world.
What God dos Irot 1ois gt involvd in th world, and sinhas no busi-
n s si n s u p p t l s i n gr l r h r r r ' i s .
20 h Hist<.lr lf uolutior Tbought

ABL o STRATA ttu Ops or' APAR,ANr Arr, I,rr

UPoN IlE l!AR.

Tray.
o Shetl-arl.
N Glacial Drift. i fl y Wapon.
o Brick Ert. E8
z
Nowih 1
Red }Crg.
Comllin )
o

Flun. 3

s lase.
"" .'"-"-.. . . . ;I E
F Gypr.
A,
UJ
e
t- !9o{9n
rltl,
}t"",.
fgu]ta. Canivr
l
astricht' Cyloid. l _ osaun.
Uppe Chalk. Ci"ooid. l !rg,g' olyptyodo'
Iwe Chalk'. s Brs, Bone.
Uppe leensand Prrlian Coodilia.
Iver Grensand
o
N 'lYeld
la1..
N IatingSan.l Igunodn.
'l Puck Bd. _l\Iasupial' - Cheloni y Bone.
5 Kimmeidgian. P]iosauu'
ofodian.
ellovian.

"E- * .f:{
ltoest ble.
Bath-Stone. EE
o
z StonofieldSlt' o
asupial.
o Grat oolit.
o
Li. Ithoptegia.
"{-*-";:":"w
lrl
U' BonoBed.-Ar vrAL|
Figue 15. Th Linnaa
w i t h t h m m r so f | o
o m P h n s i v s) r sa t l
s e t ( k n o w n a s t a a ,s i n 8
shard past' althoughas
without rgardto volu
interprtationgavit n
hypothsisof shrdd
.

l1 DEscEN |

!

What out humans?

that humans a part i
on th Bgle voyag
Figur 14. h history of lif as snin 1860. his is takn trom Rihard irra dl Fugians' h
own's Pleot<llog.Aftr th rigi, Own aman implal nmy of
ally Darwin didd l
th Darwinians,thinking thir theoryto matrialisti;ut, s young mn
want to onal his th
Own and Darwin had nfrindly.own probly bama
transmuttionistof a kind (with God guidingth oursof history),ut on points aout volution
should not nssarilyintprtpiturslik thsas implyingvoIution.As mnts of all tim, h si
with th tesof lif, popIworkd out th fossil rord thinking that it might and his history.' (. D
simply tll of th ordr in whih God squntillyrtdOrgnisms. th topi and dal wi
two-volume work, .
 he istlr of uolutior Thougbt 21

#;dse
)F APPAANOE or ANIA, ,rFE
l'

rffi
>1.0o0,oo0speies (nim|s)

.";;i;.^-"6.
a--;.;"; ;.
di

"*Wehf?
&
I
6

..-;;H'r't"+$:s

d

5,,l60 seies (songbirds)
trl

ld
,d.l. osauu.l
id. r l8lrs' PoIptyodn.
I
' y Bone. l
lin Coodilia. l
I

odo.
pial,- Cheloniy Bon
u8.

,hyopteygi. "-t#
i, by Foot-pfints.
totygia.
Figure15. Th Linnaanhirarhy.organisms n all plad in a hirarhy
itodontia. with th mmrsof lowr-lvlsts ing groupd into vr fWr(ut mor
omprhnsiv) stst highrlvls.All organismslongto on and only on
st(knownas taa, singulartaon) at ah lvl (tgory).This refltsa
'ia, by Toot.pints. sharedpast, although as With th fossil rord' muh of this was workd out
lL|A ganoh. Ineta without regardto volution. Hn, giving th hirarhy an volutionary
interprtationgav it nw maning and at th sam tim onfirmd th
-si hypothsisof shard dsnt.
Ei
.o
I ganOld. ."f
I plao.ganoid.+-
ig.{ plaoid.-

$1'$:'*:w
Fuoids. zoopl\I,os.
60. This is takn from Rihard
wn aman implalnemy of
matrialisti; ut, as young mn
n proalyama
ing th ourseof history)'ut on
<thesas implying volution. As
re fossil rord thinking that it might
entially ratdorganisms.
l] DEscN oF MAN
What aout humans? From th first, Darwin was always ston.old rtain
that humans ar part and parl of th volutionary pross. His nountr
on te Bgl oagewith th nativs at th ottom of South Amria, th
Tirra dl Fugians' had onvind him of this. In the origi, howvr, tati-
ally Darwin didd not to mak muh of th human qustion. H did not
wnt to onal his thinking; h simply did not want it to swamp his gnral
pointsaout volution and its pross.Hn, in on of th grat undrstat.
mnts of all tim, h said mrly' ..Light will thrown on th origin of man
and his history'' (. Darwin 1859' 488). Darwin fully intndd to gt ak to
th topi and dal with it at lngth. This h did in 1871, offring a major
two-volum work,The Desent of n.
22 Th Histor of uoltilr T'htlugbt

naturlisti prsPtiv
tl ssay on how humal
with spiritualism and .
natural vnt. Thr ml
off of this onlusion,
tristis that ould not
human hairlssnssan
ally livd with nativs,
powr unusd by pril
prodd y sltion.
th Thor of Nturl '
Suh a onlusion w
good ar8umnts. Natul
tiv human faturs,
that is haratristi o
aury and dsirabilit;
.Wlla
wr not suh
rtainly not out of th(
sion of sxual sltio
.Why
ar mn ig and s
stors who wr got tl
domsti? Baus th
st jo with th famil;
anstors found sua
womn (using Darwin'
th standard of ut1
x p t d l y ,a l l o f t h i s i
timnts abot th lat
}.lg.l. Uppl ffgrlrultr.'rJ}*3f;Ji:kr'
didd minn ov
l,owefi$tre thlt of s dog,
mntal powr in man
326-27).
o. For.broln, et.l lrtisphers,
t.
;f. Fist visal arb'
g. solrdvislarh.
No argumnt is n
r,. ]ld.brin. .,rln,8 (llt'lign'is. I{. vetbsl olurnns and multl his thinking wr star
. Ilind.lrrai. erlllunt, medulla in pe:or d0vlor(.ut.
obloug8t. may hav n a rvol
d. e. it.**'ltli]l.
'il }tnri
ties.
. I. I,. o ti yx. tri. It is littl wondt
By univrsal alaim,
Figur 16. Similarityof th rnryoof humirnand dog. Again, W sth
that nglish Valhalla,
strngthof the rigi in r.r.raking
sof fts lik rlrs'wll known to all
iologists.(rom e Dstrl|-.|

Aft Darwin

Bfor the origi was

\/hnon looks at this ook, it is rally rathr puliar. Most of th ook
of progrss, it was op1
is not on th lrun-ranspist all ut onsists lf an tndd ssay on th to that of msmris
seondary mlrnism of su,rlsltion.Tlr puliarity of tlr topi yilds Darwin hang thing
th lu to its gnsis. Alfr] Russl !all startd y shaing Darwin's was rtainly a ultur

f ally livd with nativs, maintaind that thr ws a vast mount of rain.
J stjo with th family.
nok. Iwe figrrr that ol dog'
jf. First visal s}t.
9. Sond viseal h.
H. Vrtbal lumns and
in r8{ ot dvlonrt,ut.
mult:!
d. Antri l
I{. osti Jxtenrllle.
Il. il tr o oyx.
tan and dog. Again, W s th
:ts lik ths,wll known to all
ly rathr puliar.Most of th ook
:onsistsof an tnddssay on th
:r.Th pulirityof th topi yilds
alla startd y sharing Darwin's
The Histor of uolutionr Thought 1)
z-)
naturalistiprsptiv on human origins and vn pnnd a stimulating lit-
tl ssayon how humans might hav volvd. But thn h am namord
with spiritualismand onvind that humankind ould no han, purly
natural vnt. Thr must hav n a Mind guiding human gnsis. In sup-
.salla
port of this onlusion' argud tht thr Wr many human hara-
trististhat ould not hav n th produt of natural Sltion, suh as
human hairlssnssnd humans' grat intllign. \Wall,who had atu-
powr unusd by primitiv man, and thrfor it ould not hav n
produd y sltion. (\7alla's ssays ar olltd tnts otribtios to
th Thor of Nturl Seltio, 1870.)
.Walla
Suh a onlusion Was anathma to Darwin, ut h flt that had
good argumnts.Natural sltion unaidd ould not produ many distin-
tiv human faturs, ut sual seltion ould. Darwin argud that muh
that is haratristi of humans am aout through varying standards of
auty and dsiraility. Hn, although distintiv' th faturs itd y
.!alla
W not suh as to tk mttrs out of th rang of volution and
rtainly not out f th rang of sin. Thus Darwin Wrot a long disus-
sion of sxual sltion in gnral and thn its appliation to humankind.
10fhyar mn big and strong and powrful and intllignt? Baus thir an-
storswho wr got th st womn. \7hy ar Womn soft and yilding and
domsti? Baus thir anstors who wr got th st mn and did th
.Why
ar humans hairlss? Baus that is wht thir
anstorsfound sually iting. Why (t gt mor spifi) do Hottntot
Womn(usingDarwin's languag)hav suh ig aksids?Baus this was
th standard of auty of th tri, and th ravst ravs got first hoi.
xptdly,all of this is wrappd up with som fairly standard Vitorian sn-
timntsabout th rlativailitis of mn and wmn: ..If mn ar apal of
didd minne ovr Womn in many sujts' th avrage standard of
mntalPowr in man must aov that of woman'' (. Darwin |871,2,:
26-27\.
No argumnt is ndd to alrt us to th fat that by th 1870s Darwin and
his thinking wr starting to sttl into som fairly onvntional mods. H
may hav n a rvolutionary, ut h Was not always that far ahad of his
tri. It is littl wondr that Darwin am a major sour of national prid.
By univrsal alaim, whn h did, h was urid next to Isaa Nwton in
that nglish Valhalla, I/stminstrAy (Figur 17).
Aftr Darwin
Bfor th rigi was pulishd, volution rod on th ak of th dotrin
of progrss,it was opposd y th ida of final aus, and its status was akin
to that of msmrism and phrnology-it was a psudosin. ow did
Darwin hang things? As far aS progrss was onrnd, Darwin himslf
was rtinlya ultural progrssivistand saw volution itslf as progrssiv.
24 Th istor of uoltttior Tblught
Rmmr that as f
ntural sltion spal
blind.law planation
fo th nd-ditd n.
thisws no mino mt
th organi world is it
H was rought up oI
sntd in th lassi dl
Tholog (1802). Hov
Darwin may hav l
ny kind had no pla
plaind without roul
T h s t t u so f v o l u t i
and surprising answr:
tionary thought up to
n, funtioning wit
pratitionrs using n:rt
organi istn and
might all a profssio
man' so th fat of hi
Thomas ny Hul1
in Grmny. Unfortun
all am ardrrtvoll
rigin, whih showd
gthr undr th umr
Figur 17. artoon of Drwin as n ap. T hr-lmorwas lw.avsgntl and vry kn on natural s
nvr svag. Th nglish Wr vr}' pr<ludlf Drwit.t, nl thy still r. H is
that th whol issu o
on th ak of today's {10 not. Now, as tn, Darwin is as muh a prt of
was unndd, and ot
ultur s of sin. (Pulishd in l871 |n t|111or|1rmgazin.)
and that sltion ou
othr putativ auss'
today might ll mar
owvr, h lso saw that th link twn progrss and volution Was
v o l u t i o nt h r o u g h a k i
somthing that rought down th sttus <lftlr Itt,and h ralizd tht
Admimdly' thr w'
natural sltion is a mhanism that dnis th invitaility of iolgial
thory. on was tht t
progrss. In othr ,ords' lnptuallyDwin rok th link twen th
s how suffiint nw
two' althouglr, having done this, h rgud thar natural sletionould lad
tions that sltion or
to progrSsas th nd rstrltof what today's rlr-rtioists rvuld ll a kind
unfortunatly Darwin
of arms ra, with vr ttr ftursoming from omptition with rivals.
..If w tht whn two orgni
tak as tlr stand:rrdof high orgaisation' th amount of diffrnti.
turs blnd with thos
tion and spialisationof t svrirloflnsirr alr eing whrr adult (and
variation, within a g
this will inlud th advanmntof th rain for intlltualpurposs),nt-
What was ndd was
ural sltionlrly lds tlwards this standrd:for all physiologists admit
nw fturs from g
that th spialisationof organs, inaslnuh as in tlris stat rhy prform thir
Was that on th bsis t
funtions ttr, is n advanrilgtO ah ing;and hrrth ar'rmultion
ing physiists (notal
of variations tnding towards spialisation is within th sop of natural
suh a lisurly pro
sltion'' (. Drwin 1959,222; this is trlm th third dition of te origin
But in a way th sir
of 1861).

]hhumor was alwavsgntland
l of Darwin, and thy still ar. H is
:hn,Darwin is as muh part of
t teHornt mgazin.)
]twn progrss and volution Was
s of th lattr, and h ralizd that
lnis th invitaility of iologial
Darwin rok th link twn th thory.on was that h had no working thory of hrdity.Popl ould not
ld that natural sltionould lad
lay's volutionistswould all a kind
lming from olptition with rivals.
Lnisation,th amount of diffrntia-
3ans in ah ing whn adult (and
rain for intlltualpurposs),nat-
;tndard:for all physiologistsadmit
h as in this stat thy prform thir
. ing;and hnth aumulation Wasthat on th asis of lulations aout the ag of th arth mad y lad-
tion is within th slp of natural
frm th third dition of the origi
Th Histor of uolutior Tbougbt 25
Rmmrthat s far s {inl aus Was onrnd' Darwin thought that
naturalsltionspaks to th dsignlik aspt of living ings. It givs a
blind.law planation: sltion lds to adaptation, and this is th rson
for th nd-dirtdnatur f suh faturs as ys and hands. For Darwin
this was no minor mattr. H always rognizd that th dsignlik natur of
th rgani wrld is its most distintiv fatur nd should not ignord.
ws rought up On th Argumnt from Dsign' spially as it was pr-
sntdin th lassi disussion of Arhdaon \Villiam Paly in ls Nturl
Tholog(1802). Howvr' what was no lss inrportant was that although
Darwin may hav n a dist (latr h am agnosti)' for him a god of
any kind had no pla in a sintifi planation. vrything must .
plaind without rours to mirls or othr spially guidd forms of for.
Th statusof volutionary thought raisspartiularly intrstingqustions
and surprisinganswrs.Darwin himsl{ wantd to rais the status of volu-
tionary thought up to th lvl of physis and hmistry-s a matur si-
en, funtining within univrsitis and rsarh ntrs, with skilld
pratitionrsusing natural sltion as a toll to ut through th prolms of
organi xistn and funtion. H wantd volution to what W today
might all a profssional sin. But it was not to . Darwin was sik
man' so th fat of his idas had to ntrustd to othrs-followrs lik
Thomas Hnry Huly in ngland, Asa Gray in Amria, and rnst Hakl
in Grmany.Unfortunatly, thy had aims othr than thos of Darwin. Thy
all amardnt volutionists,nd all agrdthat Darwin's rgumnt in th
rigin, whih showd how many aras of iologial inquiry ould tid to-
gthrundr th umrlla of volution, was dfinitiv. But fw, if any' wr
vry kn on natural sltion.No on dnid it outright, but som thought
that th whol issu of finl aus Was ovrlown nd hn that sltion
was unndd,and othrs thought that final aus still ndd xplanation
and that sltionould not really do th jo. ithr way, popl turnd to
othr putativ auss' suh s Lamarkism, volution y jumps or what w
today might all mromutations (th thory is known as saltationism)' or
volutionthrough a kind of intrnal momntum (orthognsis).
Admittdly,thr wr som signifiantiolgil polms with Darwin's
show suffiintnw variation ould aris and maintaind within popula-
tions that sltion ould hav a lasting fft (Vorzimrnr 1970). Somwhat
unfortunatlyDarwin ompoundd his prolms in this rspty ssuming
that whn two organisms rd, th usual fft is that thir distintiv fa-
turs lnd with thos of thir mats. n, if only on has a nw' valual
variatin, within a gnration or two it is lndd into virtually noning.
!hatwas nddwas a ,.partiulat''thory of hrdity,that prsrvdgood
nw faturs from gnration ro gnration. Anothr prolm of Darwin's
ing physiists (notaly th futur Llrd Klvin), thr Was too littl tim for
suh lisurly pross s naturl sltion to tak fft (Burhfre|d |975).
But in a Way th sintifi prolrns Wr not th diding fator. lfhat was
1
:
26 h Histor of uolutioltr Thoght
4
i

signifiant ws that vn Darwin's most ardnt followrs wr gnrally not

that intrstd in turning volution into a profssional sin' at last not as
Drwin nyisioned it. This ws th tim of grat fossil disovries in th Nw
\world-all of thos dinosur rlonstrsof th past-and naturally thr was
grat intrst in woking out th tru pths of lif's history (Figur 18). In
ombination with not-ltogthr_rlilmthodologis liftd from mryol-
ogy' most famously Hakl's so.alld iognti law tht ontogny (th his-
tory of th individual) rpitulatsphylogny (th history of th group),
muh ffort was xpndd on working out th full dtails of th volutionry
tr of lif. But as a ausal thory of origins, volution was somwhat of a
flop. Its nthusists Wr ontnt to kp it mor t th lvl of a ..popular si.
n,'' whr it ould hv a diffrnt rol.
Why? By th sond half of th nintnth ntlrry, mn lik Huxly wr
working had to ring thir soitis rut of th ightnth ntury and into
th twntith (Dsmond 7994, 1997)' Thy trid to form th military, md-
iin, th ivil srvi'duation, and othr institutions' A strjtlik pysi-
ology oviously hs pragmati payoff in suh ars as mdiin (F.igur19).

TtI
. . G tS t i
I . i g u r l 9 . h
uninhaitlusth
<rfvryn'saW sWag
Vitorians th nd for tr
nginring.(Fr om P ul

volrrtion sms to hav

ind with th idolog
th nw rfornld soi
C h i s t i a n i t v ,a k i n d o f s
origins, story of hurn
they wr going arrd wh
bsilly this ws how l
rus of th day, of whol
nr rbrt Spnr. J
anything from th igh
Figur 18. Dinosaur modls'It ws Rihard own, in th rly 1840s,who from th undiffrntiat
llda major grou lf rptilsas dinosaurs.From th firstthy aptud
th homognous to th
popl'simaginations'as thy still do. Thsonrt,paintedmodlswr
th arth, in th dvlt
mad fo th Grat hiition of London in 1851' whrBritain showdits
prowss to th world. Thy ar now prsrvdin park in south London. Not Soity, of Govrnmn
that thy show dinosaursas rthrhavy,lursyruts'Tldy,w think thm atur, sin' Art, tl
muh mor ativ. though sussivdif]
 h istor of uolutilr Tbogbt 27

ardnt followrs wr gnrally not

a profssional sin,at last not as
of grat fossil disovris in th Nw
lf th past-and naturally thr was
,aths f lif history (Figur 18). In
mthodologisliftd from mryol-
,iogntilaw that ontogny (th his-
ylogny (tlr histry of th group),
ut th full dtails of th volutionary
'igins, volution was somwhat
of a
it mor at th lvl of a ..popular si-

nth ntury' mn lik Huly wr

t of th ightnthntury and into
iy trid to rform the military, md-
rr institutions. A sujt lik physi-
r suh arasas mdiin (Figur 19).

T ..SILNT }IIGWAY''.MAN

Figur19. Th ..GratStink.''In th surmrof 1858 London mnrly

uninhabital bausth Thams had orsuh a vil, odifrousrptal
of vryon'sraw swag.Fw things rught hom mor vividly to th
Vitoriansth ndfor trainingin modrn thnolgialsr.rjts,
lik ivil
enginring.(From uch' 1858.)

volution sms to hav no suh immdiat funtion. But' spially om-

l own, in th arly 1840s,who
rrs. From th first thy apturd
e onrte'paintedmodls wr
n 1851,whrBritainshowdits
',din a park in south London. Not
lumsy ruts.Today, w think thm
ind with th idology of progrss, it ould funtion as th philosophy of
th nw rformd soity-it ould hav a rol as kind of altrntiv to
histianity,a kind of sular rligion in its own right. It ould a story of
origins,a story of humans' xaltd pl in th pross, a vision of whr
thywr going and what thy should do to nsursussand triumph. And
basiallythis was how voluti-ln Was prolnotd, spiallyy th lading gu-
rus of th day' of whom non Was mor voal than th nglish man of si-
n H r rS p n r .P l y i n g u p v t l I u t i t l l lh, i s v i s i t l W a s a s p r l g r s s i v a s
anythingfrom the ightnthntury. Spnr saw progrss as ing a mov
from th undiffrntiatd to th diffrntiatd, or what h alld a mov from
th homognousto th htrognous:..Whthrit in th dvlopmntof
th arth, in th dvlopmntof Lif upon its surfae' in tlre dvelopment of
Soity,of Govrnmnt,of Manufaturs, of ommr, of Languag, Litr-
atur, Sin,Art, this sm volution of th simpl into th ompl'
throughsussivdiffrntiatilns,holds throughout'' (Sprrr1857,2_).
2 8 Tbe Histor of uolutior hought

vrything oys this law. Humans ar mor omplx or htrognousthan th individual and th d.
just-quotd ssay on prl
othr animals, uopans ar mor omplx or htrognous than savags'
and th nglish languag is mo omplx o htrognousthan th tongus trm for both. In th
..dsntwith modi
of othr popls (Figur 20). was
..volvd.''Darwin alwa
Inidntlly, it was rrt Spnr who popularizd th word euolutio,
Until th middl of th nintnth ntury' th trm was gnrally rsrvd him a woolly and loos t
for th dvlopmnt of th individual (ontogny). For evolution in th mod- in the rigiz, a work tl
rn sns (phylogny), most popl usd words lik
..trnsformation'' tndd as a maior ontr
(Rihards 1'992). Sing no ssntil diffrn twn th dvlopmnt of us a trm tht appad
to philosophy, litratur
tim of Tbe Dscnt of 1
day had om univrs
and as omfortaly as a
A good rligion' su
of volution' th artiul;
winism. Th asi patt
nss of volution-sir
ai

;2
> mond to man. This is
vry last, not impd i
with th volutionary p
should promot it and <
y volution. Start wit}
nv ours-nd it is :
R
soioonomi soial p<
i6
ptition, and th wak
f3
supportd amlioration
th fat that undr th r
its unhalthy' imbil
ing,thoughwll-mani
th purifying pross br
a th multipliation of tt
Sqtl failing provision, and ,
*
providnt by hightni
z'tt2
*i*
si
( S p n r| 8 5 1 , 2 3 _ 3 2
r: Christians diffr, soI
manings of thir rlig
from anything ls, ma:
world than in th iolo1
. wr lirtarian; othrs
<:
volution; and still oth'
.a !.
appaling to a kind f
i.. low mmbrs of a sp
hism. Som wr fmi
Som wr paifists (S
Figur 20. Th tr of lif as drawn y rnst Hakl. Noti just how th
All usd volution lss
ida of progrssinflundhis pitur of lif's history. (From HekeL1897,
2:188.) living world-as a full-
0 Th Histlr of ulltilrThoght

for th nW soity. No longr:r psudlsitl,

vlutit)nws ainlyrro
mor than a populr sin.

l"{
\
Th Synthti Thory N-\.J
'\\
Things prsistdthis way into th arly dlsof th twtith ntu nd
in som rsptsam vn wors whn ontinntal thirrkrs,notaly th
mbryologist Hans Drish in Grmany and th philosophr ni Bgson
in Fran, gan pLrshinga kind of no-Aistorlianthory of lif tht szrw
r
spil, nonphysial final fors guiding tlr path of volution. Dris's nt-
lhisnd Brgson,sls uitu' th foundatins of ..vitalisr,''wr iust
{
not th lmnts of forward-looking modrn sin.Rlif finally am
with th dvlopmntof Mndlin gntisnl its mlding rvith Darwiniurn
sltiorr,although vtrhr th pross took til-r-r nd lvirs not stri]ishtfor-
ward (rovine |97I). Grgor ndl, Morvian mtlk who livd irr th
Austro-Hungian mpir, disovrdth ssntialprinipls of hrdity in
th 1860s. Unlik Darwin, h sa'ur, th trnsmissionof hartristisas ..pzlr-
tiulat,'' living tht faturslik olor nd siz n p.rssdon without
th thrat f ing lndd wy (Figur 2l ). Howvr, it was not Llntil th
bginning of th twntith nttrrythat popl am to ppitth impor.
tan of Mndl's work and saw tht it providd ky lmntin th story of
volution. You miglrt think tht this was a trril missd opportunity, and
that, hd Dwin rad Mndl's ky papr, h would hllv rlizd tht tow
h had th answrs to all of tlr issus of hrdity tht his thory of ntural
sltion dmandd. Hen, th thory would hav mlvd forward mor
quikly' muh rlir. This is proaly not sO. vn whn th work of
Mndl was disovrd, it to<lk tim to ssinriltit. Marrt. -loplat first
thought it an altrnativto Drrvin rathr thn a omplllnt (slo.)'So
F i g u r 2 l . T h p a p l a n
vn if Darwin lrad rad Mndl-nd th monk's work rvas pulishd in a ndl to work out th
journal wll-nough known that, if h sarhd,Darwin would hav found many diffrentvariti
it-thr is no rason to think tlrt thr wold hav n il ..urka'' mo- so forth. ndl ould
mnt. ndl inidntallydid rd th origi' Ir-rrrstinglv, h trvrthought passdfrom on gnf
of his own work as prtinnt to Darwin,s prtlln-rs. Judgirrgfrom th anno. rsultsw a littl too l
tations that ndl mad in t margins of th Grmn transltion of th he was only rordingr
ovrzalouslytrying to
rigi' h was fr mor intrstdin th thllogialimpliations of volr-r-
sevntnth-ntury pl
tion than in th troules of hrdity that Drwirr fad.Th is rally no strr-
pris hr sit.t,:rftrall' N,Indlwas first nd formost mn of God :rnd
o n l y s o n d a r i I ya p l a n t s i n t i s t .
Thr wr fairly straightforward rasons r,vhyat first, whn Mndl was was rvald, and it
rdisovrdat the ginning of th twntih ntury, no ()l1thought tlrirt Finally, around 1930,
h was spaking to Drwinin issus'Natr'rrll,rly ntiistsfr.rsd on taly R' A. Fishr (19
ig vriations and so tndd to favor a kind of saltat thory of ovrall Wright (197' t9z)
hang, that is volution y lag jumps. Slowly' howvr' thanks prtiu- slvs and ar transmi
larly to work in th sond dd of th ntury y Thlmas Hunt organ works nly in th grou
and his studntst olumia Unir,rsityin Nrv York, th ntllr of th gn work to rh mplti
 Tbe Histor rlf uolutionr bought 1

osin'volution was rtainly no

ddslf th twntithntury and

Lnlntintalthinkrs, notably th
'and th phiiosophr Hnri Brgson
l-Aristotlianthory of lif that saw
th path of volution.Drish's nt-
foundtionsof ..vitalism'''wr just
modrn sin.Rlif finally am
tisand its mldirrgwith Darwinian
s took tim and ws not straightfor-
a ravian monk who livd in th
h ssntialprinipls of hrdity in
rnsmissionof haratristisas ..par-
r and siz an passdon without
21). Howvr. it was not until th
lopl am to appritth impor-
lrovidd a ky lmnt in the story of
's a trrilmissd opportunity' and
lr, h would hav ralizd tht now
lf hrditythat his thory of natural
. would hav mlvd forward mor
y not so. vn whn th work of
l assimilatit. Many popl at first
)r than a lmplment(slow). So
Figur21. Th pa plant.This lowly grdnvgtlWas th organismusd
h monk's work was pulishd in a
ndlto wrk out th priniplsof hrdity.lt ws asyto grow nd am in
;arhd,Darwin would hav found manydifferntvaritisand forms-sd nd pod shapand olor, hight'and
: would hav n a ..urka'' m<l- so forth.ndl ould thrfordistinguishnd rord hw ftursWr
|rigi' Interstingly,h nvr thought passdfrom on gnrationto th nt. It now smsrtainthat ndl's
's rsultswr littl too good to tru. It ould hav ntht suonsiously
prolms.Judging from th anno-
ns of th Grman trnslation of the hwas only rordingrsultsthat fit his hypothsis,or that th gardnrwas
r thllogialimpliations of volu- ovrzlouslytryingto plashis mastr.This stylizdpituris from a lat.
seventnth-ntury lant ook.
Darwin fad.Thr is rally no sur-
rst and formost a man of God and

sns why at fist,whn Mndl was
)ntith ntury' n() on thought that
aturally,arly gntiistsfousd on
kind of saltatory thory of ovrll
s. Slowly' howvr, thnks partiu-
ntury y Thomas Hunt Morgan
in Nw Yrk, th natur of th gn
was rvald,and it ould sn as th omplmnt tO natural sltion.
Finally,aound 1930, a numr of highly giftd mathmatial biologists,no-
t a l yR . A . F i s h r ( 1 9 3 0 )a n d J . B . S . H l d a n ( \ 9 2 \ i n n g l a n d a n d S e w a l l
!right (|91, I92) in Amri, showd how ndlian gns sort thm-
slvsand ar transmittd irr groups (arrssntial finding, givn that sltion
works only in th group situation),and thn it ws possilto bring Darwin's
work to th ompltion that it ndd.Along with mutation (th oming of
2 h Histor of uolutionr Thought

new variation, ausd y spontanous hangs in gns),ntural sletion

an truly a signifiant for for hang.
Sin y th 1930s th qustion of th ag of th rth was no longr prss.
ing (th disovry of th warming ffts of radiotiv day showd that
th rth is quit old nor-rghfo th slow workings of sltion),iologists
movd rapidly forward with nw idas (known as popltion gentis) tat
wold put mpirial flsh on th mathmatial skltons. In Britain a highly
voal supportr of h thoy was Thomas nry Huxly's grandson Julian
Huly (th oldr brothr of novlist Aldous Huxly)' who produd a major
work that pulld idas togther:uolution: T|e odr Snt|lesis(|942).
Sintifially, aftr Fishr th most important figur was th oxford iologist
. B. Ford (|964), who did groundraking studis of sltion in popul.
tions of uttrflis and who gthrd around himslf in shool of ..ologi.
al gntis''a numr of youngr rsarhrslikwis intersrdin sltion
and its ffts in natur. Notworthy wr Athur ain (1954) and Philip
Shppard (1958), who workd on anding pattrns in snails, and Bernard
t t l w l l1l 9 7 3 ) ' w h o s i n t r s w
s r i n i n l u s t r i am
l lanisnl-rh wys in
whih uttrflis hang adptiv olor pattrns as thir haitats ar hangd
y th fftsof smok and pollLrtion.[n Amria th influntial figur was Figu 22. ThodosiusDo
th Russian-orn gntiistThodosius Dozhansky' whos Getics d th fourth from left) at onf
rigi of Spies |\97) ws an inspiration to zrwhol gnrtion of volu- sienmns having stud
.s7orking
tionists (Figur 22). longsid him wr othrs' notaly th o- Hrvard gnriistRihr
nithologist/taonomist rnst ayr (author of Sstemtis d the rigin of ornll gentiistBru V
was a Spnishprist for
Spis, 1942), th palontologist Gorg Gaylod Simpson (Tmpo d
right in th front row. l
od i uolutio, 1944), and th otanist G. Ldyard Stbins (Vritio
sxswas long-lastingp
d uolutio i Plts,1950).
By th 1950s Darwin's dram of a matur' profssional sin of volu-
tionary iology was ralizd. orovr, it was gnuinly Darwinian, for al-
though thr had n prtndrs to th ausal thron, notaly Swall For som, lik Dozh:
.lfright's
pross of gnti drift (random hngs in gn frquny in small ought pogrss-prog
popultions du to th vgaris of rding),it was ognizdthat th ky pitur that saw God w
fator in organi natur is its adaptivnss, its manifstation of final aus' divisions that wr ru
and tht natural sltionis a full :rnd satisfying way of xplaining this ph- God's ga nd n rrnfu
nomnon. At th same tilr' prolress-and all th moralizing and philoso- was attratd to th id
phizing that wnt along with it-hd n xplld.No on was going to us ( 1 9 5 5 ) ,w h p r o m o t d
this kind of profssional iology as an us for quasi-rligiousspulations and ayr and Simpson
aout th sttus of humankind and th oligations tht natur lays upon hu- too likd th ida of pr
m a s ( F i g u 2 3 ) ' som kind of dirtion r
Yt, for all this, thr is on mor important fator to th story, and this Baus all ths s
partly xplains why to this day volutionry idas rmin so ontrovrsial to arly twntith ntury,
so many. Although volutionary iology ws upgradd from th lvel of a dripping with mtaphv
popular sinto th lvl of a profssional sin,this did not our sim- surpris if this had not
ply out o{ a disintrstdqust for th truth mn who hd no aims ut th h i m s l f ,t h i s n w l l r d
finding of th workings of natur. Virtually vry on of th nw profssional sional status for thir a<
biologists' from Fishr to Stins,amn volutionistuslr was at- sintists-thn thy wt
tratd to th sujt y thinking that it was mor than just a sintifi thory. pts. Thy ndd to d
 he istor,-oi uolutionr.[,hought 3.]

hangsin gns),natural sltion

.
ag of th arth Was no longr press-
:s of radioativ day showd that
w workings of sltion),iologists
known as popultiotl gtis) t,at
i]atialskltons. In Britain a highly
ras Flnry Huxly's grandson Julian
ous Huxly), who produd a major
io: The odru Snthesis (1,942).
'tant figur was th oford iologist
king studisof sltion in popula-
'und himslfin shool of ..ologi-
hrs likwis intrsted in sltion
.rArthur ain (1954) and Philip
ing pattrns in snails, and Brnard
n industrial mlanism-th ways in
attrnsas thir haitatsar hangd
-rAmria th influntial
figur was Figure22. ThldlsiusDozhansk1''This pitur of Dozhansk frot rrl,
)ozhansky,whos Gtis d tbe
fourtbfrom lft) at a onfrnwith his studntsundrlinshow sussful
ion to a whol gnration of volu- sienmanshvingstudntswho rvill rry on and dvlpyour idas.Th
him wr othrs, notaly th or. HarvrdgntiistRihard Lwlntinis sondfrom th lft in th front rrw;
ornllgntiistBrr"rlWllais riglrt hindhim; an1FrnisrlAyl (whl
or of Sstmtisd th rigi of
g Gaylord Simpson (Tempo d wasa Spanispristforlr .rlan Amrian volutionist)is on tlr far
rightin th front row. larly' Dr,in'svirvson tlr rltivailitislf t
nist G. Ldyard Stins(Vrition
sswas a long-lastingprjudiirl vllutionryirls.

ltur' profssinl sin of volu-

it was gnuinly Darwinian, for al-
th ausal thon, notly Swll
r hangs in gn frquny in small
ing), it was rognizd that th ky
:ss, its manifstationof final aus'
tisfying way of xplaining this ph-
lnd all th moralizing and philoso-
n p l l dN
. o on was goingto us
usfor quasi-rligiousspulations
ligations that lratur lays upon hu-
)ortant fator to th story' and this
ary idas rmain so ontrovrsil to
, was upgradd from tlr lvl of
> n a ls i n t,h i s d i d n o t o
Lthy mn who had no aims ut th
ly vry on of th nw profssional
t an volutinist baus h was at-
'as mor than just a sintifi thory.
For som, lik Dobzhansky, th i1ttrz1tionws pliitly rligious.
roughtpogss-progrssto httmns,that is-inttl a kind of ovrirll world
piturthat saw God working his way through th fosof ntur. Ignoring
divisions that wr ruial to arlir thinkrs, Dozhansky thought that
God,sgraand irn unfrrrlingrtion ould omind. Unsurpisingly, h
was attratdto th idas of th .[suitpalontollgistTilhard d hrdin
Hulv in ngland
(1955),who pronrltdsimitr idas. Oths, lik .[r.rlirr
and ayr and Simpson in Amia, wr slal.i thir thinking, trt thy
too likd th ida of progrss and thought that it showd tht volutitln had
somkind of dirtilnand mning.
Bausall ths sminal volutinists wr popl wh grw up irr th
arlytwntithntr.lry,whn rnulr <lfvolution ws nly a populr sit-t
dripping with mtphsial ilnd moral irplitiol1s,it w>uldhav n a
u r s i m - surprisif this ad not n t lnin motivating ftlr. But' lik Darwin
saw that if thy wr to hav profs-
himslf,this nrv rd of volr"rtionists
sionalstatusfor thir tivitis-so1thingthy rdntly dsird as full-tim
sintists-thnthy would nd ttl purify thir work Of its trasintifias-
pts.Thy nddto dal witlr dlrrinsof prlgrss:rnd with tapoltd
34 h Histor of uolutionr Thought

STUDIS oi\ IltliADIATD POPULAT]ONS oI.. moral hortations. so 1

D no So P I I, M l] I' N oG S1,]tr]l* Progrss' morality, and l
dud as profssional rs
By BRUCE WALLA]]
foundd journal uoluti
Bio|c,ot I,a|ralor,CoIdSqri ll'rbor, N.Y.
authors' dpositd with t
(With Fiv Tet.figures) ing aout his intntions l
ns s possil. On t
(Reciod 28 Jd |95) ..Th fild hs rahed
roulo Also, volutionary rsa
It i well korn tht the ridspead use of ionizing
diations, besuse of tlri gneti from th phylognti ir
effets, poso polem futur genations. The* aditions indue gene
.""g""diog
mutations. Te vast majority valuting th fators of
of muttions haie dletiousfftson individuals rying
them. Und the pssue of ontinued mutatin, on th othr hand, it ha
thse deleteious mutation will
aumulat in popultion. Therefor, an iadiated srh has suffrd in thl
population will, on th avrage, be
lrmd-hav its .fitnes' reduced'-by a ontinual
u*.o." to iradiation. tion'' (lttr to G. G. Fr
The psnt atile ummariz obsrvtions made
on irdiatd po|oi1"". oi lo. Yt, having don thir
so1lhilam|ntr.Som of th.mterial presentedtee
ha n p"ri'a peviously
(Walla, 1950, lg5l; wl]try & King, l95i, 1952). tionists thn wrot othr
This summy,^ho,n".'',u., will intro.
due lrew mtrial in ddition to xtding the oiginai xhorttion as anything 1
obsrvatins.
rrtivs wr fousd on l
lr, AND DtEoDs nintenth. ..Th most s
Th erp'imntnlnop]nlion.s. Tire epimental population
of D' mIn,oslerar kept just this: knowldg' t
in luite and snage.The iginal fli *ee oind
fom an ooniR stin kept' As a first proposition of
lry mass tnsfr for mny year. Fouten letal.
and smi.letl-fee sond homo- volution, it is sumittd
somes w extrted fom thig.stin though the us of a
eriof matings idential to
tJredesribed later (Fig. 2.)'.Flies arying ths sond aquisition of nw truths
homosomes and mitures of
orgon.R and .markd stok' hromosomeothe than th ..""od ly the mutations of th n.
;;;; larental flis
f th popultions.
to othrs and y thir a<
Rief driptions of th pulation a given in Tbl 1.
Th left-hand olumn gives ity)'' (Simpson 1949, 1
th identifying numb fo eah popultion. Th ondolumn
.Stok'indiats indiate tircorgin of th
opulation. populations whooiginal flies aiedluth'l.."u Stlinism was at its pal
*uond
homosomes of oregon-R. deivation. Tlre more rent populations undr thrat aus of t
ae subpopulations
f population 5 nd 6; the deignation in the tabl gives tie^pent,r
p"p"t,l"" and the surprisingly, thrfor, ..
gnation duing whih eggs wI removd to tat t nw
olumn indits the numlr.ofdul in th population
"p'i*;;;.
.,gu'
The tird and undr som urrnt (
.uiu.* to popotu.
tions of aliout ]0,000 indir'iduals,.small'fo ".g"*, ogy whih an mad t<
pultions"qo"oly *]iti u'u. t"n
1000 ildividuals. Th last thee olumns of Tal l giv th typJ of this hr maintaind
ot."*po*u, the
<lose.and the dat th poulatin was statd fo eh population.
. Cholri" rxl)osue
efto ontitrouseposur t radium bms'. No u*oo.uin
th case of poula-
. f.hj rrrk . tlon lldr (]l)ntr&.t
tio. A1'"(30.l).5l-l7,U.S. Atomi n-r.(miirln. Th Past Half ntt
Figur 23. A papr on fruit flis y Bru \/allae.S th aknowldgmnt
at That was all 50 or mor
th^bottomof th pag. By th 1950s' evllutionistswr omi.'g
u.iy rativ
at finding funds for thir work. Doznsky prsuaddthe U.S. synthti thorists in Am
Aiomi nrgy -
ommission that fruit flis w th prfetmodl organism for
studyof the sidrd just as a sin,
effetsof nulr rdiation, a majlr worry at th tim givn th thory dsrvdly is on
o.'goi.,g tstlng
of oms (in.thatmosphr). In nglanj, . B. Frd frsuaddth ur"ld mpiril inquiy. very
Foundation that th study of uttrflisgav insight inio th
sprad of gnsin flourishs as nvr fort
popultionsnd hnewas of gratvalu in studyinghuman gntiallv
used mpirial, hav rahd
ilmnts.
Rosmary and Ptr Gral
paradigmati xampl of
hs n vry imoortant ]
 Tb istor <lfuolutionr Thogbt
JJ
D PoPULAT]olt,s ol. moral hortations.S thy did. Thy took
th tranous thinking aout
LNoGS1,Eht* pro8rss'morality, nd th meaning of
lif out of th sin that thy pro.
rALLcli duds profssionalrsarhrs.st ayr
was th first ditor of th nwly
founddjurnal ullutioll (Figur 24). His
ni llarbor, N'Y. orrspondn with prosptlv
authors,dpsitdwith th Amrian Philosphial
t.figure)
Soity, is highly rval-
ing aouthis intntionstht th new sin
as muh lik stablishdsi-
uJ |9b) nss possil.on th on hand, it had
..Th t mthodologi,tty propr:
fild has rahd point whr quntitativ
,IoN work is dly ndd.
Also, vlutionaryrsarh' as you .."li,.,
nizing radiations, bause of t}rei
hs shiftd almost ompitely
geneti from th phylognti intrst (proving volution)
ntion. Tese ditions indui to an ologial intrst
gene valuating th fatorsof volution'' 1ltirto !. A.
l deltiouffetson individu"l. Goslin, aih 5, |94B).
."..}g
,ttion, ths delteriou mutations on th othr hand, it hd to vlu fr: ..Th
wil.i prstig of volution ar re-
titod pulation will, on the avrge, sahhas suffrdin tlr past ausof
;inual u to idition.
too -oih philosophy and spula-
tion'' (lttrto G. G. Frris, h 28, 1948).
r md-e. on idited populations of Dro-
sentedhelr ben pulished Yt,hvingdon thir professional sin,
pviously thsparadigm-making volu-
952)'Thi summy, howeve,will tioniststhn wrot othr works tht wr s
intro. dripping wl, pog..;, and moral
oiginl obsrvations. xhortationas anything to b found in Hrrt
Spnr, .*..f,, that thir di-
tivswr fousd on th prlms of
th twntith ntury rthr than th
}lEoD8 nitnth...Th mst ssntial matil
ftor in th nw volution se]sto
tal populationot D' ml,nasler juqtthls: knwldg,togthr, nssrily,
a kept witlr its sprad nd inhritan.
fr:T ogon-R
stainkeit As a first proposition of-volutiona.y this
::lb*iid 1n
iial- and smi.lethl-fee drivd r.n- ,p..i."lly human
seond
the use of a seieof mtings identil
"h.on,- volution,it is sumittd tht promotion
t of knowldg is ssntially oth th
resondhromosomesand
mitues of
quisition of nw truths or of ]osrapproimations
to truth (mtaphorial-
hethgn th eondrveethe pantal ly themutationsof th nw volutionl ,n
flies lso its sprad y ommuniation
to othsnd y thi aptan and larning
n in Tabl l. Th left.hand olumn of it (mtapho.i.,l[ it, hrd-
gil,es iry)'' (Simpson 1949,311). Simpson Was
e seondolumn indiates t" writing at midntury, just whn
o.lgi,, #tu
re original flies aied lethal.frer Stalinm was at its pak and in Sovit Russi
seoj iologists ;,' po.,i*l,. *...
oJe recrnt poulationsae ubpopulations undrthratauseof th high position
bIgivesth pntalpopulation of th hrlt,' T. . Lysnk. Un.
and th surprisingly'thfor, ..Dmoray is wrong
.o ..11' tb nw
<lpulations.Th thid
in may of its urrnt aspts
ages:.lg. efers to and undrsom urrnt dfinitions,ut
y::1l dmoay is th only
pula. ogy whih an mad to mr an tlrially olitial idol-
;lopultlons frequentl *ith frwr il,an good soity y th standards
table l giv the type of of thishmaintaind'' (21).
posue, th
r o eah popultion. Choni
eosur
b', No rosue in th ase
of poul-
).J ).ill7,U.S. At,oi Dnrg"r.(.o
m mrl.,. Th Past Half ntury
/alla.Sth
aknowldgmentat
rilnistswr ominguiy ariv ht ws all 50 o mor yars ago. Th
no.Darwinians in Britain and th
,,prsuaddth U.S. Atomi nrgy synthtithoristsin mri uilt good
foundations for thir sin.on-
modl organismfor study of th sidrdjust as a sin,whili it h,. .u..y
right to , modrn voluttonary
...i'.. givnthe ongoingsring thorydsrvdlyis on of th most fo,-n,j-looking
l. f and *itinf ras of
:. . l.ord prsddrh Nuf6ld mpirial inquiry. vry on of th rs
: insightinto th spred of nsin tratd y Drwin in t|te rigin
flourishsas nvr for.Sltion studis
s u d y i n gh u m a ng n t i a l Iy u s d thmslvs,oth thortil and
mpirial' hav ahed a er high
dgre of sophistiation. The work of
Rosmaryand Ptr Grant (1989) on th
ak siz of Gal5pagos finhs is a
paradigmatimpl of llnt
sin.h dvrrtof -o...'ln. biology
has nvry important hr for offring
otl-rnw insights (for instan, th
 signifianof drift at th

IoN
l*{itNl C :EYl}l,ll 1g
nti fingrprinting for d
dtaild in dward o.
Snthsis (1975), hav po
.W

tion and riprolaItrui

t o p l a i n t h i n t i a i sl
from an individualisti p
m p i r i ls t u d i s - f o r i n s t
(|982) on th rd dr of S
a powrful fator in th
sially, th fild (oftn hr
Darwinian fshion looks '
fantiid and sual ha
tion. Plontology has t
major fossil finds-rntl
twn fish and amphiial
mntion th finding in Ind
srs, naturally known as tJ
thniqus hav nald g
vr gtr pision. At
poposd (for instan, th
nd th lt Stphn Jay
othr aras of volutionar
t r n so f t h p s t ( f o r i n s t
| | 9 7 6 | u s d o l o g i a lt h t
pst pttrns of animal
hang, an vnt that o

on lf th iggstprrlms
Figlr 24. Th ovr of th first issu lf ulltir.t.
of thsarly profssionalvolutionistswas of findingpopl,working to thir
standards,who wr nt studyingfruit flisor similar organisrns.Th journal
was launhdwith gntfrom th Amrin PilosophialSoity(flunddy
BenjaminFranklin) in Pildlphia.Rvlingly,t only prsonOn th grxnts
ommittwho opposdth fundingWs th mryologistdwin onklin. Figure25. Puntuatdqu
mryologyWas thn th qunof th illogialsirrs and onklin was not g r a d u a l i s m .. l d d g i r
onvindtht volutionor-rld th asisof ral profssionirlsin.Prhaps l t r n t i v p i r u ( B ) ' i t g l
bausf thsdisiplinarytnsions,volutionistsrsplnddv ignorrng sltationist,whih rathr s
mbryologyand it was not until th 1980s,y wih tim m<llular iology omparativtimsals.F
had ompltlyangdth landsap,that it gin -t a mirjlrlmntin hangan tk pl duri
volutinarystudis. arlygts rorddand a
 '' -*:
"i." h Histor of urllutilttrThought 7
: r ;:-'- i ' ;
"".
:
signifian of diit at th m<llularlvl) irnd nw thniqus(tlrus of g-

UTION
-t
,L oF oANI YoLUlN
nti fingrprintingfor dtrrnininghrdity). Soirlhavior nd instint'
dtailedin dwrd o. !7ilson's magnifint survy Soiobiolog: Th Nc,
Sthsis(1975I,hav powrful nw modls of plnation, lik kin sl-
,".l!tl:.: .,.1,,,t''t,',,,,: tion nd riproalaltruism, <rnfirmingDrwin's htrr-rh that th right wy
to plin th intiaisof soial intrirtions,inluding grotrp hvior, is
Ie +r N r.? from an individualistiprsptiv.Baking th thortial woks ar majo
mpirialstudis_for instan,that of Tim lutton-Brok and his assolats
(1982)on th rd dr of Sltl.lnd-tht slrlwtht volution has indd bn
,.,:;:,l:i powerfulfator in th vrllution of soial bhvior. Somwht ontrov-
.it,l
ril saetion i wirl Pqul* siallv, th fild (oftn hr known as uolutior pshlllgy) lso in vry
.ust , . -.' . . ' . . . . . . . . . ' . . . . . Darwinian fashion loks at humankind, trying to xplain suh things s in-
:,
**"eL .
h {attn8 fntiidand sual havior nd mating pattrns in trms of na rr-rralsl_
ai{*P. 'srcirl ,,,.tt,'.,.'t$'
.. '.':'.,:1, tion' Palontologyhas rr revolutionizd. Nlt tlnly ar tl.leontinud
7pod*y. R,,4sr ,i;
mior fossilfinds-rntly the disovry in th ndian Arti of a link -
.t
twn fish and amphiins (Dashlr, Shuin, and Jnkins 2006), not to
:,t,',u!{t;
mntionth finding in Ind<lnsillof littl hr'rmanlikfiglre (<ltoflorsi-
, ,,:
,.:|i
*8: . :,:lil 51-n. ,a t u r a l l yk n o w n a s t l r . . h o i t ' ' ) ( I } o w n t l . 2 0 0 4 ) - u t m o l u l a
. .,.:l:i*
63 thniqushav nald popl to xplor th past ours of volution with
I i' 'll;* vrgratrprision.At th mor Onptuallvl, nw thoris hav n
. I r.,r'.r1ti
proposd(for instan,th th<lryof puntuatd quiIiria of Nils ldrdg
''"1.
, ,,.''-:|':. and th lat StphnJay Gould 11972|\ (Figur 2.5),and idas drawn fron-l
othrarasof volutionary studis hav bn applid to undrstand th pat-
s9 trnsof th past (tr instan,th way in rvhih th lirtJohn J. Spkoski Jr.
.. . ,tli
[1976lusd ologialthoris aout islanl iogograplryto throw light on
103
past pattrnsof animal divrsity, as wll as on th Grat At-nrianInt-
111 hang,an vnt that ourrd ablut 10 million yrs ago whn Sluth nd

.- :" ir

I F'uolutil. Orrof th iggstprtlln.rs

was of findingpo1rl' working to thir
flisor sinli]arorganisrts. Th journal
lrianPhilosophilSoity(flunddy Morphology |v]orphoIog

valingly,th nly prsol-lon th grants

s th mryologistdrvirlonklin. Figur25. Putuatdquiliriurn.lrr th Drrvininpitur,lld..phlti
iologilsinsanjtlnklin Was not grdualism,'y ldrdgnd Gluld (A), volutiongos smoothly.in thrr
lasisof ral profssionlsin.Prhaps piture(B),it gosy jumps. Gould dnidstronglythat h was a
altentiv
. v o l u t i n i s rt s s t l l l d hdv i g n i n g whih rathrsuggststhat mul-rof th dtwas aout
saltationist,
80s' y whih tim molr-rlr iollg1, mpartiv timsls.For fuit fly gntiist,1,000 1'rsis an ag nd muh
that it aginzrm lnajlrlrntin hngan takpladuring tht priod.For a p[oIltologist, 1,000 yars
arlvgtsorddnd any hngsar ging t()snlinstintanOlls.
]8 he isttlr1'tf uolutirlrlT|,,"r''

Nrlrth Amri ioind and irrrinralsm<lvdnoth to sor.rthand south to noIth

[ a r s l r a l l t l ' 1 9 8 2 ] )( F i g u r 2 6 ) . T h o r i so f g o g r a p h i d i s t i b u t i o nw r
rvolutionizd y th oming of plat ttonis.Until this poir-rt,in a tradition
mrad y Darwin in th rigi, volutitlnists had spnt many happy
hours throrving up hypothtialland ridgs nd finding wys in whih sds
and smll arrinrillsould rtlss lrg odis Of w:lr.Nw th rrrovingof th
ontinnts did ll th rvok for thm. L1'strlsaurus' a marnmal.lik hrivo-
ous rptil found rathr nlor than 200 rillion yars ago) is fat' short' and
squat. It is rtainly not an animl tht would hav roamd far and wid. To.
day it is flund in th sam fossil dposits (Lorvr Triassi) on th ntinnts of
Afria, (Soutlrast) Asia, nd Antarti.This woLrld inxplilwr it not
or th ft tht mor thn 200 rillion ,varsgo all thos olltinntstluhd =[so,t-.".
lvn thl. W part lf Pangaa.Thy lrv sin driftd apart (}.igur27). \
\_=--,!/
,/
Cynognathus
riasic ReptiIe

ryffi
,/r.q-..:--,I}
ffi

Anteaters
Armdi||os
Capybaras
Glyptodonts
Figur 27. Pttrns of foss
Monkeys
y o u a s s u m r h t t h o n t i
Opossums how muh volutioniststo
Porupines w hv ome from th ol
Phorusrhaids
Bears Sloths
Camels ouans
Systmatis has bn tran
Cats oxodonts
th oming of omputrs
Dogs
absorbd, quantifid, al
EIephnts likwis is opn to nw u!
Horses th synthti thorists t
Pecaries oame euolutior det,el
Rbbits ptual bakwatr to th
Raoons volutionary studis (ar
Skunks instn,aout th ways
apirs mans shar th sam un
Wese!s th ways in whih org
among th hottst aras (
Figur 26. lr Grat Amrin Intrhng. By trating North Arrrri and Thr ar' of ours'
Sluth Arrlri s if thy wr t''v islands ing olonizd, on an rvork out th importngg-1hg ygr
th ptl ts li lr.lg anl th numrs tlf invadirrg gr.ouPs ln wluld volutionists' partiularl;
Ii rium.
pt at qr"ri s o m t i m sq u s t i o n s o f .
 Th Histor of ullutior Tbought 39

)vdnorth to south and south to north

rorisof gographidistribution Wr
tonis.Until this point, in a tadition
volutionistshad spnt many happy
lridgsand finding ways in whih sds
dis of watr. Now th moving of th
Lystrosaurus,a mammal-lik hrivo-
00 million ),arsago' is fat, short, and
t would hav oamd far and wid. To.
i s( L o w rT r i a s s i )o n t h ( ) n t i n n os f
l. This would inpliabl wr it not
l yars ago ll ths ontinnts touhd
ly hav sin driftd apart (Figure 27).

Antrcti

Mesosaurus Glossopteris

--ffi
Permin Reptile PInt
*.*

ry#
, . -,.,s *
Anteaters

\.
ng.By trating North Amria and
rds ing olonizd, on an wlrk out
numrs of invding groups on would
fr /w
Armadillos
Cpbaras
Glyptodonts
Figur27. Pattrnsof fossil organisms.Thsdistriutionsmak snsonly if
Monkes
youassumthat th ontinntswr joind. Phnomnalik thsshow jusr
Opossums howmuhvolutioniststoday still work in th Darwinian mod nd yt how far
Porupines we havom from te rigi itslf. (Unitd StatsGologial Survy.)
Phorusrhaids
Sloths
Touans
Systmatis has n trnsformd y nw ladisti thniqus,muh aidd y
oxodonts th oming of omputrs that hv allowd massiv amounts of data to
asord,quantifid,alulatd, and undrstood (Hull 1988). ophology
likwisis opn to nW undrstandings.Aov all' mryology' a sujtthat
th synthetithorists tndd to ignor, has now, spiaily undr th nw
nameuolutilrdeulopmt (evo-dvo), n onvrtd from a nonon.
ptualakwatrto th most xiting ar of rsarhtoday in th fild of
volutionarystudis(arroll 2005). Fantsti nw findings hav mrgd,for
instan,aout th ways in whih organisms s divrs as fruit flis and hu-
mans shar th sam undrlying gneti mhanisms for dvlopmnt, and
th ways in whih organisms ar uilt and vriations ar produd ar
amongth hottst aras of rsarh (Figurs 28 and 29).
Thrar, of ours, ontrovrsisand diffrns.No on wants to deny
th importan-th vry grt importn-of natural sltion.But som
partiularly in aras lik palontology and mryology, whr
volutionists,
somtims qustionsof adaptiv signifianar (as was arlir th as for
40 Tb Histor of uolutilnr Thrlught

popl lik Thomas Hnr.

prhaps othr ausal fat
of th thory of puntuat
many faturs ar not al
that is, nonfuntional by-
L w o n t i n l 9 7 9 ) ( F i g u r s.
..Just
rlying to oftn on
yard Kipling (Figur 32).

l" n Dfd Sr Antp U ad.A Ad-I]

f-trffifI.__l
DRosoPHIL Ox (;N1]s

HAD- x. oo-

Figur 28. Dvlopmnt in th fruit fly. In th ntr W hv th gns that

ontrol dvllpmnt, in th ordr in whih thy Our on th hromosom.
Bllw w hav th lrv and aov w hv th grown fly.

L|vn{d P K o A Y | H o I L L L K I - F Y | \ R Y L l } . t t t V L s o U K I W F o N wKLP\ N v R
i."ii." K s A Y o o V L L K l H F N Y L t ! : t s r r o I K l W l o N R R M K W K K D R$L P N l s
^i'."., PKRsRIAYRoovLtLFKFYNYLRRRVLIAF{ALLsLRoIKIWFoNJRMKWKK LDPl .Nl K | s
" l " . ^ q P K R s R A Y . l R o o V L L L F Y ' \ ] R Y L t v tLs t R o I K I W F o N R M W K K DKL P N K I R s
htkHoB4 P K s A Y o o v L F L K t t ] Y l \ R Y L R R v F I A s L L s t R o l K I W l o N R M K W KKKL DPN KIRs
;;;;;";;; f. fKss A
vYI oooovVLLLLtKKEFFHYYNNR Y YL L R
l RRRRRVVEI A
| A LI L Lf fsLEsRoI oK Il W
K IFW
oNFR oNRM RK
WKK W
KDKKH
L PD\ | * s
i'.],u"'o K L P N KVS
)li"?i'i."* miisvooY!ELKFYIILIRvtnrtllrsqIKIWFoNRRKWK!q KLPNK]

F.igur29. Homologous amino aids.On th lft is olumn tlf th nmsof

gnsin diffrntrganisms.All of thsgnsar ruial,kiking in at similar
stagsof dvlopmnt.Th rows irr th main o irrmad up of lttrs Figure 0. A spandrel fro
orrspondingto amino ids,produdy thsgns,in turn produinga and assistants.Gould's la
vital protin.It n snthat th gnsof frrritflisand humans ar virtually spndrls(tually,th tru
idntial,ldingto th onlusintht dvlopmntis lik uildingmodls sndrlssm dsigndf
with l,go riks.You startwith th sam pisand thn put thm togthrin rsultof othr nds.Hn<
vry diffrnt Ways. p r h p sn a t u r ls | t i o ri
 he Histor li l:,ulltiorThought 41

popllik Thomas nry Huly) not ovrwhlmingly prssing' think that

prhapsothr ausl fators wr sigrrifiant.This Was th undrlying thm
of th thoryof puntuatdquiliri;r,with Gould in partir.rlararguing that
manyfatusr not ptiLllarlyadptiv ut ar mo ..spandrl-lik''-
that is, nonfuntional y-prduts of th volutionry pross (Gould nd
Lwontin1979) (Figurs30 and 31). H ausd Darwinian volutionists of
rlyingtoo oftn on ..Just so'' storis' th fantaulous.lik tals told y Rud-
yard Kipling (Figur 32). Today many vo-dvo nthllsiasts fl muh th

r t LJb d-A Ad-B

Blf-_-l
1o Ct,NA-s

J l I

ABDON

[n th ntr \4,v th gns tht

ih th ()ur On th hromlsom.
tv th gorvn flv.

r ivGoUKIWI oNRMW r.tv

B q f jA F l s L G L R o I K I W F oN R M W D i s R L P N K R s
V |A l - s L L s t o I K I W F o \ R M K W K D H K L P N I K | s
R V EI A | - f f L L s IoK I W lo N R M K W K D H L P N K | s
R V |A H S L L s R o IK I W F o N R R MK W K K D H L P N K I R s
v t n r s o l K l w t o N R R K W o t . t , ' l t s $
R R v t I H L L s R o| | W F o NR R | vK]W K K D H K t P N K V R s
R B Y q l4 ! . s R o I K I W t o N R R K W q ] L .

n th lft is a olumn of th namsof

3 gnsar ttil,kiking in t sirrlill.rr
main o ar mad up of lttrs Figur0. A spandrlfrom th Vill lr:rrrrsin
in Rom, pintd y Raphal
l y thsgrrs,in turn prtldr-ringa ndassistants.Gould's lim is tht rrrnyftursof organismsar lik
rs of fluit flisnd humans r r'irtually spandls (atuzrlly,
th tru thnilnam is pndntiv).Although ths
dvlopmnt is lik uilding mo1ls spandrlssmdsigndfor painting' in truth thy r sirply produd as th
1pisand thn put thm togthrin rsultof othrnds.Hn, on shlr-rldnot lways look for adaptation and
prhaps naturalsltilnis ovrratd.
42 h Histor of uoltior hought

Figu32. Th elephant's
nos am aout aus,
that many storisaout a<

Figur 31. Th Irish lk. Atually, this now-tintast,found in th ogs of

(Gilbrtet a|. 1996, 68,
Ireland,is not a tru lk. lt is rmarkalfor its antlrs.Thy ar so larg it is
hard to imagin that thy ould vr hav n of adaptiv dvantag.A popu. lik ths ar wis and p,
lar hypothsistoday is that thy grWthrough sualsltion'th lks with asts for a nW disiplin
iggr horns dfatingthir rivls and gtting th fmals.It is suggstdthat way' th vigor and xir
th lks mfrtil for thy stoppd growing and prhaps n immtur firmd.
lk would pass on his gns,vnthough latr in lif h and his offspringwould Dos this thn man tl
at a disadvantg. An xplanationlik this shows how arfulon should
tionary thinking has nor
in tlking aout adaptationnd its asn.In on sns,th ntlrsar
many poplas a kind ol
daptiv,but not nssarilyfor survivaland prhapsnot for muh of th
animal'slif. nums of artils and
grams) show o justify i
volution. dward o. W
of books dsignd to sh
sam way. ..Th homologis of pross within morphognti filds provid
flows not iust moral }
som of th st vidnfor volution-just as skltal and organ homolo-
forsts-ut an aItrnati
gis did arlir. Thus, th vidnfor volutilnis ttrthan vr. Th rol
of natural sltion in volution' howvr, is sn to play lss an important But mak no mistak
rol. It is mrly a filtr for unsussful morphologis gnratd y dvlop. nts th human mind v
mnt. Population gntisis dstindto hang if it is not to om as irrl- wys, point fo point i
vant to volution as Nwtonian mhanis is to ontmDorarv ohvsis'' narrativ form is th 1

)w-tintast,found in th ogs of
for its antlrs.Thy ar so larg it is
' nof adaptivadvantag.A popu'
roughsxualsltion,th lks with
)ttingth fmls.It is suggstdthat
d g r o w i n g n d p r h a p s n i m m a t u r
r later in life h and his offspring would
this showshow arfulon should b
r. In on sns'th antlrs r
I and prhapsnot for muh of th
;s within morphognti filds provid
n-just as skltaland organ homolo-
volution is ttr than vr. Th rol
v r ,i s s nt O p l a y l s s a n i m P o r t n t
l morphologisgnratdy dvlop-
o hang if it is not to om as irrl-
:hanisis to ontmporary physis''
b lIistor of t,llutilrhoght 4
Figur32. Th lpant'snos.In his stlris,Kipling laimdtht th lphnt's
nosamaoutaus roldilgradit and pulld. Gould suggstd
thatmanystorisaout adaptivadvantaghav out s muh plausiility.
(Gilrtet a|. |996,368). No dout postrity will tll us whthr sntimnts
lik thsar wis and prsintor mrly th unjustifidffluvia of nthusi-
astsfor a nw disiplin tht is trying to stalish itslf as important. ithr
way' th vigor and itmntof ontmporay volutionry thory ar on-
fimd.
Dos this thn man that th nonsintifi'mor idologial sid to volu-
tionarythinking has now vanishd? Dos volution n longr funtion for
manypoplas a kind lf sular rligion? Not at all. on th ln hand, hug
numrsof artils and olks (not to mntion radio and tlvision pro-
grams)show or justify idologial impliations srid to or drawn from
.Wilson
volution.dward o. is paradigm, for h has pourd out a stram
of ooks dsigndto show that volution is progrssiv nd that from this
.Wilson'
flowsnot just mral hortations-for th prsrvation of th rin
forsts-utn altrnativviw of ration to that of trditiona1rligion:
But mak no mistak aout th power of sintifimatrialism. It prs-
ntsth human mind with an altrnativ mythology that until now has al-
ways,point for point in zons of onflit, dfatd traditionl rligion. Its
narrativform is th pi: th volution of th univrs from th ig ang


11 h istrlrrlf ulutitl1'Tbought

lf fiftn illion \Irsago thrlugh th origin of th lrrrntsand ls- 10,000 yas old and that i
tial odis to th gir.rnings of lif on artlr. Th v<llutionar1'
pi is rgy in just six days' follor
mythology in th sns that th laws it addus h and now ar - vrsal flood' sav only for
livd ut an nvr dfinitivIvprovd to form a aus-and-fft (Rus 2005; Numrs 20(
()lltinuul fronr physis to th slial sirrs,frol this world to all story hr' ut th onstan
othr wrlds in th visil univrs, and akward through tim to th nthusiastsfor this kind of
ginning of th univrs. vry part of istn is onsidrd to thory ut as somthing m.
ldir-rt ttl plrysil larvs quiring no trnI ontrol. Th sirrtist's tionl lif systms that
dr,ltiontl parsimony in plarrtion xluds th divin spirit and hld morl norms. In th'
oth.rtran()Llsagnts' Most importantly, w hav om to th ruial ..It rl
Hnry . orris: is
stag in th istory of iollgywhn rligilnitslf is sujttl th .pla- punishmnt for murdr n
nrtionsof tlr natural sins.As I hav trid to show' slioiologyan mnt for othr sriousril
aount for tlr vry lrigin of mythology y th prinipl of natural s- v o l u t i o n r yd t r m i n i s mt
ltiln ating on th gl-ltiallyvolving mtrial strutur of tl-rhu- is 1989, 148). Apprntl
rrrnbrin. ..ath
Rus,'' wll_known
If this intrprtationis orrt, th final ]isivdg njoyd y si- th morl rot. lvlor rn
ntifi naturalism will lm from its apait to plin traditior-ralrli- has morphd into th morr
gion, its hif ornptition,as a wholly mtril pn<rnrnon. Tholgy gnt Dsign Thory, whih
is nlt likly to surviv irs n indpndntirrtlltualdisiplin. (Wilson q u i r s p i a l r a t i v i n t r
1 9 7 9 ,r 9 2 ) is mor than just a onfli
pting idologis. Th fou
Alrothr sirrtistmuh givn to rhis kind of spulatio-altlrough h dis- Brkl law profssor Ph
likd trditionirl notiot-tslf progrss and wantd to sustituthis own idas l d t r m s :. . T h C h i s t i n
aout th rndonrnsslf lrang-was StprrJay Gould, partiulaly in his an asy targt atrs it ]
popLrlrssays' ..his Vir,v<lfLif,'' pulishd n-rtlnthlt, in |'trtrlistlr.l,. with worldly ids lik hul
Vry arly did Gould fil to draw slmkind ot moral mssagfrlrrrhis writ- whih r atually profoun
ings, 'uvhthraout th raism ndmi in our slityor th nd for onsr. ingly, this humanisti mt:
vtiorr.Not that Gould gv tradititln;rlrligiln muh mo sop than did gay mariag, aortion, ar
Wilsln. Any idas aout lur ing th fvord hiidrr-rof Gld, tlrt w hu-
rnans might th rason for th ati()n'ar simply huris. ..Sin di-
nosaurs Wr n()t nroving toward markdly largr rains, and sin suh a Conlusion
prospt may ii utsiltl-rapailitisof rptilia dsign . . ' w lnust as-
sLlmthat onsiousnssr,vouldnot lravvolvd On our prlantif a osmi - h history of volutiona
tastrtl;lh had rr<ltlirimd th dinlsaurs as r,itirns.In an ntirly litral until 1859, it was littl m.
sns' W oW otlr istn's larg nd rsrningmanl-tills,tl orrr luk idology of progrss, th r
stars'' (Gould l989, 1ti). Hr Go[d antiip1ltdth so-alldnrv athists' It was opposd to traditiol
lik his grat British ountrpat in th rlm tlf popular sin writing' litral truth of th Bil l
Riha]Dawkins. In the T|la God Dlusil(206\Dawkirls i1r]us tlrt hris- that hang oms only t
tiarrity and otlrr rligilrrsar th r:ljlrsours tlf hur-rrrkind's
ills, and tlrirt on th rigin of Spis ,
w nd :r sular philosophy as a sustitut-a sulaphilosophy informd truth of volution on an
y Dirrwiniirn thinkir.rg,whih itslf mks app.lsto traditional rliiorlsnot tion was ignord or down
mrly tltiosut asolr-rtly fls.Phils<lph Danil Dnlrtt'sDt'itz'sDn- Darwin, notaly Thoms
8rorrsld (7995) is rrtlthrmplr of this kind of litratur. volutionary idas as a ki
Hirding th tlthr Way ar th n.ranyritis lf volutionary thinking. In n altrnativ idology to
Amri tlda,stlvy aft srvyonfirms thilt most Arrrriansdrl not - to soil rform. Th thir.
liv in volutiln.Ind]'th grat rnajlritylrlivtht th artlr is lssthan of ndlin gntis. I

tlr origin of th lrrrntsnd ls-
on arth. Th volutilrrarypi is
ws it ddus hr nd ntw ar -
y provd ttl form a aus.irnd-fft
ial sins,from this wold to ll
:, and kward throug tinl to th
lart of istt-l is onsidrd ttl
] I1Otrnal ontr<ll.Tlr sintist's
rtion luds th divin spirit and
ortantly'w hav om to th ruial
r rligion itsIfis sujtto tlr pla-
hv trid to show, slioillogyan
lology b,vth prinipl of lli1turals-
lolving matrial strutur tlf th hu-
h final disivdg njoyd y si-
's :rpaityto xplain trditionl rli-
olly matrialphnomnon.Thology
,ndrltintlltuldisiplin' (Wilsrn
s kind of spulatir-r-zrltl-rough h dis-
rrrlwantd to sustittlris own ids
s SthnJay Gould, partiularly in his
pulishd mlnthly in |JturlHistor.
:rkind of moral mssagfrlm his writ-
Liin our soitylr th n]for onsr-
nal rligilnmuh mor sop than did
Lf:rvordhiidrn of Gol, tht w lru-
ration' ar simply hLrris...Sin di-
rrkedly largr rains, and sin suh a
is of rptilian dsign , We tnust as-
v volvdon our plant if osmi a-
saurs as vitims. In an ntirly litral
nd rastlningmamtnls, to our luky
J antiiptlth so-lldllW thists'
h r l m o f p o p u l : r rs i n w i t i n g .
tsitm(206)Dawkins argLlsthat hris-
t)r soursof humankind's rlls, and that
)stitut-a sular philosophy informd
aks appalsto traditional rligions not
lsophDanil Dnntt's I)ui's D-
rr of tis kind of litratur'
'ny itis of volr.rtiorr:rry thinking. In
rrfimsthat most Amrians do not -
rjoritylivthat th i.rthis lss than
Th istor of uoltior,Thought 4S
10,000yars old and that all organisms wr atdin urst of divin n.
rgyin just si dirys, followd som tim latr y thir dstrution in a ni-
vrsalflood, sav only for thos luky pairs that floatd away in Noah's ark
(Rus2005; Numrs 2006|. Amian vanglial hristianity is not our
storyhr,ut th onstant attks on modn Darwinism show larly that
nthusiasts for this kirrd of rligion do not rgard volution as just sintifi
thoryut as somthing lor' a mtrialisti,sular altrntivto th tradi-
tionallif systlllsthat arris lvithirr it opposition to ll dnt and long-
..ration sintist,''
hldmoral norms. [n th words of th lding th lat
Hnry. Morris: ..It is rathr ovious that th modrn opposition to apital
punishmntfor murdr nd th gnral tndny towrd lniny in punish-
mntfor othr srious rims r dirtly rlatd to th strong mphasis on
volutionarydtrminismthat has haatrizdmuh of this ntury'' (Mor-
..notorious Dirrwinian philosop Mihal
ris 1989, 148). Apparntlv' th
Rus'''a wll-krrown ..athisti hltratrist'''has mad rnirjorontibution to
thmoral rot. Or rntly' th rz.rtlrr ud litrlism of ration sin
hasmorphd into th mor sophistiatdanti-Drwinism ot so-alld Intlli-
gntDsignThory' whih ssasptsof ogani lif as so ompl as to -
quirspial rativ intrvntions y th Dsignr. But still th dat today
is mor than just a onflit of sinvsl'lsrligion and on twn om-
ptingidollgis.Tlr foundr of th lntllignt Dsign movlnnt' formr
BrklylzrwprotsslrPhiilip Jlhrrsorr,is pliit ir-rsing volutin irr th
..Th
oldtrms: hristian philosophy that Was ovthrown in th 1960s was
an asy targt aus it had om idntifid with Amrian ultur nd
with worldly idas lik human prftiilityand th invitaility of progrss,
whihar atually profoundly un-hristian'' (Johnson 1997,106), Unsurpris-
ingly,this humanisti mtaphysisis linkd to moral failings-pornography'
gaymarrig,aotiol-l'nd lik transgssionssuh as soialisrl.
onlusion
Th history of volutionary thory falls into thr stags.From aout 1700
until 1859, it was littl mor than a psudosinriding on th ak of th
idologyof progrss,th notion that humirns an mak ang for th ttr.
Itrvasopposdto tditional rligion not so r-nuhausit wnt against th
it lft no pla for Prrvidn,th lif
litraltruth of rh Bil ut r-rs
thathango-lsonly through Gol's undsrvdgra. harls Darwin's
th origi <lfSpeishangd ll tlris. Darwin stlishdth rasonabl
truthof volution on nd for all. HoWvr, his mhanism of natural sl-
tionwas ignord or downplayd, and th lading volutionistswho followd
Darwin,notaly Thomas Hnry Huxly, Wr muh mor intrstdin using
volutionaryids :rs a kind of popular sin,almost a sulr rIigion' as
an altrntividol<lgyto th lrristin rligilnthy saw [rlokingthir wy
to soialrform. Th third stagstrtd arond 1930 witlr th inorporation
of Mndlian ] ntis. Now thr was th possiility of uilding a nW
46 Tb Histor of uoltiorl, Thought

potssional sin oi volutior.rary lrang. This ourrd, arrd it is this th- Gould, S. J. 1989. Woder
Ory that flourishs today. Yt vllr-rtiorrisrsshould nvr flrgt th past and Nw York: W. !7. Nor
its influ On th prsnt. Alorrg witlr th sin thr is still an ongling Gould, S. J., nd R. C. Lw
dbat about idolog1. nd th tnt to lvhih vrltrtion in som wY ro-
Panglossianpradigm
of th Rol Soit l
vids a sulr altrnatir, to oldr' Ovrtly rligiorrs ways Of viwing th
Grnt, B' R., nd P. R. Gra
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ow. whr. and whn r

ws lutant to addrss r
|, 1871.,to his frind Jr
paragraph that ..it is oft
of a living ing ar now
(and oh what ig if) w
sorts of ammonia nd ph
a Protin ompoulrd W
ompl hangs' at th ]
sord, whih would l
formd'' (Darwin 1887,
Cambridg. It is frn
rprodued in alvin 19t
Darwin's lttr nt onl;
of lif ut also provids t
natur of th asi iolog
sintifi irls. Althugl
alld thm nulin) in 18
in gnti prosss woul
trast' th rols plyd b
firmly ognizd. quall'
jor advans had n m.
wih for a long tim hr
gani ompounds. Th
Alksandr Butlrov, who
laortory synthsis of r
starring matrials, wr r
spratingorganisms frol
 WstminstrRuitl,,67: 244_267 .
til i Plnts, Nw York: olumia

nc humin. Pris: ditions du Stril.

'h
Yers of ltroL,,rs.Phildlphia:

r i i rso d p a r ti n f i n i t I y r o m h Thorigin of Lif

tgslf the Li Soit, Z<lolog :

of NturI Sltio:A Seris rlf sss. leffreL. Bd nd ntonio Lzo

Sthesis,amridg, MA: Blknap
Gnrationsof sintistsmy ).thav to om and go for th qus-
idg,A: Hrvrd Ur-rivrsity Prss. ion of th origin of lif is finl11'sllvd.That it lvill solvd vrttu-
loulations.Gntis |6: 97-| 59 .
ally is s tin s nything an vr amid th unrtaintis that
ding,rossrdingnd sltionin
suroundus.
trntiol lgrss of Gtis I:
ari Alrt Nlson, Tr Is Lif o rs (1956\

How, whr,and whn did lif appar on arth? Although harls Darwin
wasrlutantto addrssthsissusin his ooks, in a lttr snt on Fruary
7, 1877, to his frind Josph Dalton ookr, h wrot in now_famous
paragraphthat ..it is oftn said that ll th onditions for th first podution
of a livirrgbing ar now prsnt,whih ould vr hav n prsnt.But if
(ndoh what a big if) rv ould oniv in som warm littl pond with ll
sortsof ammonia and phosphori salts,-light, hat, ltriityprsnt' that
a protinompound was hmiirlly fornrd, rady to undrgo still mor
omplxhangs,at th psntsuh mattr would instantly dvourd' r
absord,whih wuld not hav n th as for living raturs wr
formd''(Darwirr 1887, 'i, 18). (Th lttr is in th Darwin olltion in
ambridg.It is rfrndas Lttr 747 | in Burkhrt and Smith l994 and is
rprodud in alvin 7969,4_5.)
Dwirrlttr not onlv summrizs in a nutshll his idas on th mrgn
of lif but also provids onsidral insights on th viws aout th molular
naturof th basi biologial prosssthat wr prvalnt at th tim ln mny
sintifiirls.Although Fridrih ishr had disovrd nuli aids (h
alldthm nulin) tn 1869 (Dahm 2005)' th diphing of thir ntral rol
in gntiprossswould rmin unknown for mor than 80 yars. In on.
trast'th rols playd y protirrs in mnifold iologial poSsshad n
firmlyrognizd.qually signifiant, by tlr tim Darwin wrot his lr, ma-
jor advanshad ben mad in th undrstandingof th matrial asis of lif,
whihtr a long tim had bn onsidrd fundarntally diffrnt from ino-
.!76hlr,
ganiompounds.Th primnts of Fridrih Adolph Strekr, and
AlksndrButlrov, who hd dmorrstrtdirrdprrdntlyh fsiility of the
laboratorysynthsis of ura, alnin, and sugars, rsptivly' from simpl
startingmatrials,Wr rognizd as a dmonstration that th hmial gap
spIating organisms fronr th nonliving was not insurrtlr.rntai.

19
 F

50 The rigi of Lif

But how had this gap first n ridgd? Th ida that lif Was an emr- pripittdquartz in th
gnt fatur of natur has n widesprad sin th nintnthntury. Th (Uno t a|' 2006), lthou
major rakthroughs that transformd th origin of lif from pur spula. Collom 2006). Howv
tion into workal and tstabl rsarh modls wr proposals that wr diat iologial sulfat-r
suggstdindpndntly in th 1920s y Alksandr I. oparin and J. B. S. 3.4 billion-yar-old Sout}
Haldan, as wll aS othrs (sBada and Lazano 200). Thy hypothsizd rin nvironmnt inhait
that th first lif forms wr th outom of a slow, multistp pross that nd Low 2004\' Ths r
gan with th aioti synthsisof organi ompounds and th formation of old stromatolits from W
..primordial soup.'' Hr thir proposals divrgd. Whil Haldan argud ida tht th arly Arha
a
that viruss rprsntda primordial stag that had appd prior to lis' origin of lif must hav ta
oprin argud that olloidal, gl.lik systms had formd from th soup, prmit th survival of th
lading to anaroi htrotrophs that ould tak up surrounding organi Th arly Arhan foss
ompounds and us thm dirtly for growth and rprodution. Although quird for th origin and r
th dtails of ths idas hav n suprsdd' th oparin-Haldan hy- ritial fator may hav .
pothsis of hmial volution providd a onptual framwork for th sil s soon as th plan
xprimntal dvlopmnt of th study of th rigin of lif. Laoratory watr. Unfortunatly, tht
xprimnts hav shown how asy it is to produ a varity of organi om- o n d i t i o n so n t h l
pounds, inluding iohmially important monomrs' undr plausibl os- lulr or physil rmni
mi and gohmial onditions. Th roust natur of ths rations has volutionry prosss th
n dmonstratd y th finding that organi ompounds ar uiquitous in ganisms found in the arl
th univrs, as shown by thir prsnin arbon.rih mtorits'omtary ing not only on th o
sptra' and intrstllar lods whr star and plantary formation is taking piod of th origin of lifr
pla. gnral and loal nvlro
impotant for th mrg

A Timsal for th mrgnof Lif

Th Primitiv arth
Although traditionally it had n assumd that th origin and arly volu.
tion of lif involvd svral illion yars (Lazano and Millr 1994), suh Considral progrss ha
viws ar no longr tnal. It is tru that it is not pssil to assign a pr- onditions of th arly
is hronology to th appranof lif, ut in th past fw yars stimatS hmistry may hav or
of th tim within whih this must hav ourrd hav n onsidraly r- thr ar still normous
dud. Dtrmination of th iologial origin of what hav n onsidrd ompounds assoiatdwi
th arlist tras of lif is now a rathr ontntious issu (van Zuiln t al. ing ntitis and how ths
2o0z). This is not surprising. Th gologial rord of th arly Arhan is t u l v i d n o f t h i r i
spars' and thr ar vry fw roks that ar mor than 3.5 illion yars old. hav bgun on arth' w ]
Th roks with ags gratr than 3.5 illion yars that ar prsrvd hav history and undr what
bn xtnsivly altrd y mtamorphi prosss(van Zg||en et a|.2002|, th origin of lif took pl.
and thus any dirt vidn of anint lif has apparntly n largly liquid watr and ogani
olitratd. availabl vidn sugg
Although the iologial natur of th mirostrurursprsnt in th 3.5 lkd protins and dp
illion.yar-old Ap hrts of th Australian !arrawoona formation lotids, ut watr was
(Shopf 1993) has n disputd (Brsir t a|. 2002; Garia-Ruiz t al. vids th mdium for h
2003), thr is vidn that lif mrgd on arth vry arly in its histry. omponnts, as far as w
Th proposd timing of th onst of mirobial mthanognsis,asd on th It is unlikly that Wat
low 1 valus in mthan inlusions, has n found in hydrothrmally ing th Hadan priod tl

dgd?Tlr ida that lit was an mr-
lrad sin tlr nintnth ntury. Th
d th origin of lif from prrr spula-
Lrh modls Wr proposals that wr
s y A l k s a n d rI . O p r i n n d J . B . S .
nd Lzano 2003). Thy lrypothsizd
om of a slow, multistp pross that
4aniomptlundsand th formation of
osals divrgd'Whil Haldan argud
stgthat had appared prior to lls,
3 systms lrd formd frorn th soup,
at ould tak up surrounding organi
r growth and rprodution. Although
suprsdd,tlr Oparin-ldan hy_
idd a onptual framwok for th
'rdy of th origin of lif. Laoratory
s to produ a varity of organi om-
)rtant monotnrs,undr plausil os.
robust natu of ths rations has
organi ompounds ar uiquitous in
i n r o n - r i hm t o r i r s, o m t a r y
star and plantary formation is taking
of Lif
sumd that th origin nd arly volu-
a s ( L a z a n o n d i l l r 1 9 9 4 ) , s u h
. t h a t i t i s n o r p o s s i l r o a s s i g n a p r -
l i f ' u t i n r h p a s t f w . r r s s t i m a t s
v ourrdhav n onsidraly r-
al origin of wlrat hav lronsidrd
1ontntiousissu (varrZuiln t al.
clogial rord of th arly Arhan is
|ratar mor than 3.5 illion yars ld.
i illion yars that ar prsrvd hav
lhi prsss(van Zuiln et a|.2002|,
;int llf has apparrrtly n |arge|
th mirostrutrrrsprsnt in th 3.5
Australian Narrawoon formation
,asirt a\' 20021 Garia-Ruiz t al.
rgd orr ilrth vry arly in its history.
niroilmthanognsis,asd on th
ts, has n found in hydrothrmally
Th rigi of Life
pripitatd qtlartz in th ..Sillion-yar-old Drssr Formation in Australia
(Unot al. 2006), although tlris finding has i.rlstln hallngd (Lollar and
ollom 2006). Howvr, sulfur isotop invstigationsof th sam sit in-
diatiologialsulfat-rduingtivity (Shn t al.2001), and analyssof
3'4 illion-yar_old South Afrian hrts suggstthat thy forrnd ln a ma-
rin nvironmntinaitd v anaroi photosynthti pokayots (Ti
and Low 2004). Ths rsults' omind with rports on .43 illion-yr-
old stromatolitsfrom .JstrnAustralia (Allwood et a|.2007), support th
ideathat th arl,vArhan arth Was tming rvith prokaryots' and that tlr
hrvtakn pla as soon as tlr onditions wr suital to
originof lif mr"rst
prmitth survivl of ths tvps of organisnrs.
h arly Arhan fossil rord spaks for th rlativly short timsal r-
quirdfor th origin and arly volution of lif on arth and suggststhat th
ritialfatorma hav rrt Prsnof liquid watr' whih ampos-
sibls soon as tlr plant'ssurf finlly oold lorv th oiling point of
wtr.Unfortr-rnatiy, thr is no gologial vidn of th nvironmntal
onditionson th arly arth at th tim of tlr origin of lif, nor ar any mo-
luiar or physial rmnants psrvd that provid infornration aout th
volutionaryprosssthat prdd th apparan of th fist llular or-
ganismsfound in th arly fossil relrd.Dirt inforratiorris gnrally lk_
ing not only on th omposition of th trrstrial atmosphr during th
priodof th origin f lif ut also on th trnperatur' oan pH, and othr
gnraland lol nvironmntal onditions that may or may not hav n
impotantfor th mrgnof living systms'
Th Primitiv arth nvironmnt
onsidral progrsshs en lrradin our undrstandingof nvironmntl
onditionsof th arly arth nd how tlr transition frorn ioti to ioti
hmistrymay hav ourrd (for mpl. s Bada 2004). Nvrthlss,
thrar still normous gaps in our dsription of how th simpl Orgnr
ompoundsassoiatdwith lif s w know it atdto gnatth first liv-
igntitisand how tlrsin trrrn volvd into orgnisms tht lft hind a-
tulvidnof thir istnin th rok rrd. o valuat how lif may
havgunon arth' W must asswhat th plant was lik during its aly
historyand undr what onditions th prosssthought to b involvd in
th origin of lif took pla. Lif as w know it dpnds on th prsn of
[iquidwatr and ogani polr,mrs suh as rruli aids nd proteins. Th
vilalvidn suggsts tl-ratduring arr arly stag, iologial systems
lakedprotinsand dpnddlargly on atalyti and rpliativ polyrionu-
leotids'ut watr was likel1.ssntial fronr th vry ginning, as it pro-
vidsth mdium for hmial rations tl tirk pla' Withut ths asi
omponntsJ i1sfar as w know, lif is irrrpossibl.
It is unlikly that watr md its first apparanon artlr as a liquid. Dur-
ing th Hadan priod th volatil omponnts that wr trappd insid th
52 he origi of Lif

arting plant wr rlasd (dgassd)from th intrior of th juvnil on Vnus today, althou

arth to form a sondary atmlsphr.Any primary atmosphr (if on - arly arth.
istd at all) must hav n lost' as vidndy th dpltion of rar gassin Th limat on th
th arth's atmosphompard with osrni aundans (Kasting 1'99I. tors: th luminosity of
As a onsqun of th narly simultanous formation of th arth's or phr. Standard thor
with artion, th mtalli iron was rmovd from th uppr mantl, whih aout 07" lsslumin
would hav allowd th volani gassto rmain rlativly rdud nd pro- th arly arth Wr th
du a vry arly atmosphr tht ontaind spissuh as Ho, NH-., and would hav n frtlz
H,. Sin th tmpratur at th surfa was high nough to prvnt any Wa- ( 1 9 9 a , 1 9 9 3 ) a' o ,
tr from ondnsing, th atmosphrwould hav onsistd mainly of supr- th Hadan and arly
hatd stam along with ths othr gass (Kasting 1993a|' vn though th hous fft that woul
sondary atmosphr may hav n lost svrl tims during larg-impat frzing. Th asi arg
vnts, spially th on that formd th moon, it would hav n rgnr- tory, thr wr proa
atd y furthr outgassing from th intrior, as wll as ing rsupplid from tnsiv siliat Wath
latr impators. loss pross for O' rl
It is gnrally aptd that th imptors during th ltr stags of th a- hav n primarily o
rtion pross originatd from outsid th solar systmand wuld hv n assumptiono sol
similar in omposition to omts. As suggstdin 1961 y th lt Spanish stat atmosphr onta
hmist Joan (John) 016' omtry volatil ompounds, whih appar to ordr to maintin
th most pristin matrials that surviv from th formation of th solar sys- point of Watr.
tm' may hav supplid a sustantial fration of th volatils on th trrs- In summary, th urt
trial plants, prhaps inluding organi ompounds that may hav playd a gst that it onsistd <
rol in th origin of lif on arth (016 7961).It has n suggstdthat th ,o, with lssr amout
watr on th arth was providd ntirly from this sour. Howvr, rnt and N. ar onsidr
masrrrmntsof th dutrium ontnt of watr in omts Hally, Hyaku- izd rgions na vola
tak, and Hal-Bopp indiat that omts dlivrd only a fration of it, During suh arly trr
whil th largest fration Was trappd during th arlir artionary phas and larg ontinnts ha(
and rlasdvia dgassing(Rort 200] ). rations of th arly ar
It is rasonal to assum that th atmosphr that dvlopd on th ling of rustal sdimn
arth ovr th priod 4.4 t'o.8 illion yars ago (prhaps svral tims if it surfa would hav rm
was rodd y larg-impat vnts) was sially a mi of volatils dliv- onat (Slp and Zah
rd y volatil-rih impators suh as omts nd utgassing from th in- nntrations in th
trior of an alrady-diffrntiatdplant. This atmosphr was probaly fomatin of a gloal i
dominatd y watr stam until th surfa tmpratursdroppd to -100" dd th as, th thikn
(dpnding on th prssur),at whih point watr ondnsd to form arly th tlrdr of 300 m, whi
o a n s ( W i l d t a l . 2 0 0 1 ) . T h r d u d s p i s ,w h i h w r m a i n l y s u p - an impato of *100 kr
plid y volani outgassing,ar vry snsitivto ultraviolt (UV) radiation mlting olids has
that pntratd th atmosphr aus of th lak of a prottiv Ozon twn aout .6 and 4
layr. Ths moluls wr probaly dstroyd y photodissoiation, al. yls assoiatd with tl
though thr might hav n a stady-statquilirium twn ths two portd y th impto
prosss that allowd a signifiant amount of ths rdud spis (sp- would hav n wash
ially H,) to prsnt in th atmsphr (for ampl, s Tian t al. n trmd ..ImpatSu
2005). vntually, as H, sapd from th arth into sp, rdud spis H,' and o drivd fro
in th atmosphr would hav n dpltd. Thus, in gnral th ovrall unfrozn oan low t
onsnsus at prsnt is that the arly atmosphr was dominatd y oxi- from UV radiation. Aft,
dizd moluls suh as o,, O' and N,. A similar atmosphr is prsnt plld into th atmos
 Th rigi of Lif t.l

sd) from th intrio of th juvnil o n V n u st o d , a l t h o r r g hi t i s r u h m o r d n s t h a n t h a t m o s p h t t f t h

l. Any primary iltllosphr(if on x- rlvartlr.
lndy th dpltionof rar gass in h limat ()l1th arly artlr t this stg d;lndd mainly otr two f-
osmi abundans(Kasting \99). tors:th luminosity of th sun nd th radiativ proprtis of tlr atlos-
tanous formation of th arth's or phr.Standardtl.lortialsolar-volution modls prdit that th sun W:1s
movdfrom th uppr mantl' whih a o u t3 0 % l s s l u m i n o u s t h n t o . l i r (r G
, i l l i l a n d 1 9 8 9 ) . I f t h e t m o s p h r o f
to rmin rltivlydud and prl- thly arth Wt sam as it is ll()w' th nti stlrfa of th 1rlnt
taind spissuh as Ho, NH.., arrd would hav n fozn. Ho,v, s disr.rssdxtnsivly r. Kstirrg
was high nough tO prvnt any Wa- \ | 9 9 3 a ,1 9 9 3 ) , O , - r i h a t r o s p h r m a y h v n p r s n tr l r r o u g h t l u t
ould hav onsistd mainly of supr- th adan and arly Arhan priods, rsulting in a signifiant grn-
lss (Kasting \99a). vn though th housffttht would hav pvntd th ons on th arly arth from
lost svraltims dr-rringlarg-impat f z i n gT. h s i a r g u m n t i s t l r a t d u r i n g t h i s p i o d o f a r l y a t h h i s -
th moon, it wlr"rldlrav n rgnr- tory' thr ,r prlbly no rrrjlrontinnts, rrd thtls tlrr rr,sno -
.rir'as wll as ing rsupplid from tnsivsiliat wathring. Bar.tsthis pross is nowdays tI-r[ong-trnr
lossprossfor O, rm<lvaland storag,th onlusion is tht o, would
rtors during th ltr stags of th a- havn primarily ontaind in th atmosphr and th oan' With th
: th solar systm and rvould hav en assumptiontlf a solar luminosity of -70. of th. prsnt valu, il stady-
sug'gstd in 1961 y th lt Spnish s t a t a t m o s p r n t a i n i n g - 1 0 r s o f o ' o t r l d l r r , n r q r t i di n
'ltilompounds, whih appar to ordr to mirltin man surf tmpratllr gratr thar-rtlr fzing
from th frmation of th solar sys_ pointof watr.
fration of th voltils on th trrs- In summary, th urrrrtmodls for th ariy trrstrial atmosphr su]-
i ompounds that may hav playd gstthat it onsistd of a wakl rduing mitur of o2, N,, O, and
l 1961|' It has n suggstdthat th H'o, with lssr:rnrountslf H,, So', and H,S. R1tldgass sr'rhas o
.ly from this sor.rr.Howvr, rnt nd NH.. r <lnsidrdto lr:rvn narIy:lsttor prsnt <lrrlvir-rlol-
nt of watr in mts Hally, Hyaku- izdrgionsn:rrvlanos or lrvdrohrmlvrrts.
lmts dlivrd only a fration of it, During suh rly tims whn volani islands may hav n prvalnt,
l during th arlir artionary phas andlargontinntshd not yt formd (Zan|e t al. 2007). Rnt lnsid-
)1). rationsofth arl1,aronyl suggsttht for tnsivttolli ry-
atnrosphrtht dvlopd On th Iingof rustal sdinlntsmol1llon' mrlst trf tlr ron ()n th ilth's
n yars ago (prhapssvral tims if it surfawould lrilv maind trrid in th rtrst nd mantl as illium ar-
vas asially a mi of volatils dliv_ onat(Slpnd Zahnl 2001). hr is thus tlr possiility tlrt th O,
ls omts and outgassing from th in- onntrations irr th arly atmosphrwr not high rroughto prvnt th
nt. This atmosphr was proly formationof gloal i-ovrdoan (Bda t :rl. 1994). If this wr in-
rftmpratursdoppd t -100" ddth as)th thiknssof th gloal i shthas n stimtdto on
L point watr ondnsd to form rly thordr of .]00 m, whih rvr"rld hav n thirr nough to llorv r.r-riting 1'
rd spis'whilr wr mainly sr.rp- n impator lf -100 kr in diirrrrtr.Th frqur-ry of impats of suh i-
snsitivto ultravitllt (UV) radiation mltingolids has n stimtdto on vnt vry 105*10. yars b-
us of th lak of prottivozon twnaout 3.6 and 4.5 illion vars ago' suggstingpriodi thw-frz
y dstroydy photodissoiation, al- ylsassoitdrvith th i-mltingimpats. Th pursor ompotrnds im-
y-sttquilirium trvnths two portdv the inr;lator or svntlrsizdduring th irrrpat' suh as N'
mOunt of ths rdud spis (sp- wouldhav rrwshd into th oan during th thaw priols,rvhih h:rv
..Irrrpat
lsphr (for ampl, s Tian t l. ntrmd Summrs'' (Zahnl t al. 2007). ln additiorr,(jF{",NH1,
n th arth into sp, rdud spis H,, and o drivd from hydrothrmal vnts would hv n stord in th
dpltd.Thus, in gnral th ovall unfroznon |ow th i layr, whih would lrirvprottd ths gass
r atmosphrwas dominatd y oi- fromUV rdition. Aftr larg irr-rpat th trap;ld gass would v n
d N2. A similr rrosphris prsnt plldirrtoth atmosphr'wlrr thy or.rl] lrv prsistdf<lrstlt im
.\4 Th rigi of Lif

for thy wr dstroyd y photohmial rations. During ths Butlrov, who show
pisods highly rduing onditions may hav prvaild. lysts, suh s alil
ars. Th laoratory
to inlud mor or
organi ompounds on th Primitiv arth tnth ntury a lar1
formd (Bada and L
Today organi ompounds ar so prvasiv on th rth's surfa that it is Howvr, ths n
hard to imagin th arth dvoid of organi matrial. Howvr, during th as laoratory simul:
priod immdiatly aftr th arth first fornrd slm 4.5 illion yars ago' to Hookr, ut rath.
thr would hav n no orgni ompounds prsnt on its surfa baus of nitrogn assimila
soon aftr artion th day of rdiotiv lmntshatd th intrior of cano 202a). Th fi
th young rth to th mlting point of roks (\/thrill1990). Volani organi ompounds
ruptions plld moltn rok and hot sorhing gass out of th juvnil Stanly L. ivlillr, w-
arth'sintrior, rating a global infrno. In addition, th arly arth ws also a wk on a mixtur
bing ppprdy mountain-sizdplantsimals,th dris lft ovr aftr th protin amino aids
rtion of th plants. assiv volani onvulsions, oupld with th in- organi moluls (l
tns omardmnt from spa' gnratd surfa tmpraturs so hot that Millr ahivd h
th arth at this point ould vry wll hav had an ..On''of moltn rok simulat th intra
( i . . 'a m a g m a o a n ) . As an nrgy sour
Although tmpraturswould hv slowly drasdas th infali of jts sond-largstnrg
from spa and th irrtnsity of volani ruptions dlind, lvatd tmpr.
turs likly prsistdfor prhaps 100 milliln yars or longr aftr th for.
mation of th arth. During this priod tmpraturswould proaly hav
n too hot for organi omplunds to surviv. !ithout rgani om-
pounds, lif as w know it ould not hav istd. Howvr, on th asis of
data from nint zirons, y approimate| 4 illion years ago (or prhaps
vn arlir) th arth's surf must hav oold to th point that liquid
watr ould ist and gloal oans ould gin tl form (!(zildt al. 2001).
It was during this priod that organi ompounds would hv first startd to
umulat on th arth's surfa,as long as thr wr aioti prosssy
whih thy ould synthsiznd aumulat.!hat Was th natur of ths
prosss?

Synthsisof organi ompounds on th Primitiv arth?

Th hypothsis that th first organisms Wr anaroi htrotrophsis asd

on th assumption that aioti organi ompounds wr a nssary pr-
ursor for th apparan of lif. Th laoratory synthsisof organt om-
-t
pounds from inorgni starting matrial was first hivd in th 1820s whn 500.
Fridrih \Wlhlrdmonstratd that ura ould formd in high yild y Flask
th ration of yanogn and liquid ammoni and y hating ammonium
Figur 1. Th appara
yanat. Although it was not immdiatly rognizd as suh, a nW ra ln was usd to reprsn
hmial rsarh had gun. In 1850 Adolph Strkr ahivd th lora- atmosphr.A spak
tory synthsisof alnin from a mitur of ataldhyd,ammonia' and hy- invntdy Nikola T
drogn yanid. This was followd in 1861 y th primntsof Alksarrdr dishargsin th atm

hmial rations. During ths
hav prvild.
ritivarth
iv on th arth's surfa that it is
rni matrial. Howvr, during th
.ormdsom 4.5 illion yafs
go'
unds prsnton its surfa aus
tiv lmntshatd th intrior of
I roks (.sthrill1990). Volani
srhinggass out of th juvnil
[n addition,th arly arth was also
:simals,th dris lft ovr aftr th
onvulsions,oupld with th in-
.d surfa tmpraturs so hot that
v had an .,oan''of moltn rok
vly drasdas th infall of ojts
ruptions dlind,lvatdtmpr-
illion yars or longr aftr th for-
:mpraturswould proaly hv
o s u r v i v .W i t h o u t o r g a n i
istd.Howvr, on th asis of
ly4 illion yars ago (or prhaps
v oold to th point tht liquid
lgin to form (Wildt al.2001).
-rpoundswould hav first
startd to
; as thrwr aioti prosssy
[at.\ff/hatwas th natur of ths
on th Primitiv arth?
:r anaroihtrotrophsis asd
ompounds Wr a nssary pr-
)oratory synthsis of organi om-
.asfirst ahivdin th 1820s whn
' ould formd in high yild y
n o n i a n d y h a t i n g m m o n i u m
y r o g n i z da s s u h , n w r i n
LolphStrkrahivd th laora-
lf ataldhyd,ammtrnia, and hy-
1 y th primntsof Alksandr
h rigi of I,ife 5.'
Butlrov,who showd that th tratmnt of formaldhyd with alkalin at-
alysts'suh as lium hydroid, lads to th synthsisof a varity of sug-
ars.Th laoratory synthsisof iohmial ompounds was soon xtndd
to inlud mor ompl primntal sttings, and y th nd of th nin.
tnthntury a larg amount of rsarh on organi synthsis had n pr-
formd (Bada nd Lazcano 2003).
Howvr, ths nintnth-ntury organi synthss wr not onivd
..warm littl pond'' that Darwin mntiond
as laoratorysimulations of th
to Hookr, ut rathr as attmpts to undrstand th autotrophi mhanisms
of nitrogn assimilation and O. fiation in grn plants (s Bada andLaz-
ano 2002a).Th first onvining dmonstration of th possibl synthSisof
organiompounds undr prioti onditions Was aomplishd in 1953 by
StanlyL. Millr, who invstigatdth ation of ltri dishargsating for
a wk on a mixtuf of Ha, NH,, LI2' and H,o; rami mixturs of svral
protinamino aids wr produd, as wll as hydroy aids, ura, and othr
o r g a n i m o l u l s( i l l r 1 9 5 3 ' 1 9 5 5 ) .
illr ahivd his rsults y mans of an apparatus in whih h ould
simulatth intration twnan atmosphrand an oan (sFigur 1).
As an nrgy sour Mill hos a spark disharg, onsidrd to b th
sond-largst nrgysour' in th form of lightning and oronal dishargs,
o m -
\r-,0",
Flask
}ona"n,",
.r'
500-
Flask |10ml
Figur1. Th apparatususd in th 1953 illr xprimnt.Th 500 flask
was usdto rprsntth lansand th 5-1itrflask was usd to eprsntth
atmosphr. A spark disharggnratdarossth ltrodswith a Tsla oil,
invntdy Nikola Tsla in 1891' was usdto mimi lightningand orona
dishargsin th atmosphr.
56 Th rigin of Lif

on th arly arth aftr UV rdiation (illr and Ury 1959). Th pparatus

was filld with various miturs of mthan, ammonia, and hydrogn, as H(
wll as watr' whih was thn lrtd during the primnt. A sprk dis-
harg tw th tungstn ltrods,wlrih simulatd lightnirrg atld o-
O /'{ z
rona dishargs irr th arly atmosphr, was rdud using a high-frquny y
Tsl oil with a voltag of 60,000 V. T,h ration tim ws usually a wek
|I + HCN
-\
or so, nd th nrimum prssurwas 1..5rs. With this xprimntlstup'
\
t+NH

illr was al to transfornr alrost 50% oI th originl aron (in th form \ n,n
of mthan) into organi ompourrds.Although most of th syrrtlrsizdor-
gani matrial was an irrsolultarlik solid, h was al to isolt amino
aids and othr simpl organi ompounds from th ration mixtur.
Glyin, th simlst amino id, was produd tn 2Y" yild (asd on th R, R'= H or CnH21*1 :-l
original amount of mthan abon)' whras alanin' th simplst mino Retion 1. h Strker-
aid with a hiral ntr' showd a yild of \". illr was al to dmon- hydrlyaids.
strat that th alanin that was produd Was a ratni mixtur (qual
amounts of D- nd L-alnin). This providd onvining vidn that th
amino ids wr produd irr th xprimnt and wr not iologial orr- (O. + N, + H,o). In gn
taminants somhow introdud into the appratus. ward mor rduing ond
Th othr organi lmpoundsthat Millr was al tl idntify mad it pos- aids, purins, pyrimidirri
sil for him to propos a possil ration pathway for th amino aids nonrdun i g rmosphri
(illr 1957). Th proposd synthti mhanism ws in fat th on d- duing or nutral atmospt
sibd by Adolplr Strkr (18.50)whn h rportd th synthsisof alnin. nt mldl for th aly
his invlvs th ration of hydrogn yanid' amtnonia, arrdaonyl om- r|tdth id of nonr
pounds (aldhydso ktons) to fornr amino nitrils' whih th undrgo numr of primnts W(
hydrolysis to form th amino aids (sRtion 1). Dpnding on th on- modl atmosphers of (
ntration of mmoni in t ration mitur' vr1.ingmounts of hydroy found that not only wr
aids ar produd s wll. This is what illr found, with lgr rlativ glyin was asilly th o
amounts of hydroy ids ing forrd in a ration tnixtu ontining lss A gnral trnd showd t
amronia. mor nutral, th yilds o:
It is important to not that nithr purins nor pyrimidins, th nul- lly, although thy wr :
oass that ar part of DNA and RNA, w lookd fo iI-rth miturs of apprd to spially i
th original Millr_Ury xprinrnt. Howvr' in prirnts arrid out a m i n o i d s .T h m a i n p r .
soon aftr illr's prirnty oro and oworkrs' th formation of ad- o l o g i l I yr l v a n to r g a n i
nin fronr ammonium vanid solutions ws dmonstratd (016 1960; 016 a p p r n tI a k o f i o r m t i o
and Kimll 1961). Latr it was shown that th abioti synthsisf purins th Strkr pathway and
nd othr htroyli ompounds lso tks pla undr th sam ondi- of nuloss (Frris t i
tions as in th original illr-Uy prirnr,ut with muh snrallryilds high onntrations of r(
tharr for th amino aids. In additin' it has brrfourrd that guanin an b tions, and irr ths loaiiz'
produd in a dirt ..on-pot''synthsislrlm th polymrization of quous and ktons may hav
solr-rtionsof amronittm yanid (Yusa t al' 1984). ould hav om involv
AlthoLrgh ths rsults r xtrnrly nourging, th atrnosphri ompo- Whth th prinlitir'
sition that formd th asis of th illr.Ur1' primnt is not onsidrd vant with rspt to th r
today to plausil y mny rsrhrs.It is gnrally grd that fr howv. [t has ntly
oygn was absnt fom th primitiv rth, ut thr is no gnral agr- n i f i n ra m o u n t s o [ m i n ,
mnt on th onrposition of th primitiv tmospr;opinions vry from using O,/N'/liquid wat
Strongly rduing (o + N2, NI_l.r+ H,o, or o, + F{2 + N,) to nutral mino ids found prvr
 r
'8 The rigi of Lif

hydrolyti workup y nitrit and nitrat produd in th rations,as wll as h y d r o t h r m a Iv n t s a r

y th inhiition of synthsisausd y th low pH onditions gnratddur. of ritilorgani mol'
ing th primnt.Addition of alium aronatto th systmto uffr th thir dirt synthsis.
systmnar nutral pH, along with oidation inhiitrs suh as asori aid D s p i t t h p r l
or F2n prior to hydrolysis, rsults in th rovry of svral hundrd tims s y n t h s i st h o y h s b t
mor amino aids than rportd prviously. who hav dsrid h
In th rnt nutral atmosphr primnts, th amounts of fr yanid' b i o h m i s t r ya n d v o l
aldhyds, and ammonia ar low, suggsting that th intrmdiats may b (0.2 ) and high ps
ound as nitrils. Th osrvd amino aids may hav formd y svral mh- a v r i t yo f a m i n o a n d
anisms, inluding the Strkr synthsisor th rlatd Buhrr-Brgspathway. a onrns aout th
Although th amino aids osrvd and thir rlativ aundans ar similar to hmialnvironmn
thos found whn oligomrs formd y th slf-ondnsationof HN in aqu- ath' spially th l.
ous solution ar hydrolyzd (Frris t a|. 1978|, th asn of polymi ma- ( B a d a t a | . 2 0 7 ) .
trial in th primnts ppars to disard this mhanism. primnts With
slightly rduing modl atmosphrs (Shlsingr and illr 1983) suggst
tht th addition of tras of Ho and/or H. in nutral atmosphr simulations Did Prbiotiorg
would likly furthr nhan th prodution of amino aids.
Baus of supposd prolms assoiatdwith th dirt Millr-Ury typ T h d i f f i u I t i ss u p p o
of synthsson th arly rth, diffrnt hypothsis for th aioti synth. thsis of amino aids z
sis of organi ompounds has n proposd. his suggstionrsultd from In th rly l990s h
th disovry of dp-oan hydrlthrmal vnts. A group of rsarhrshav on th rol of omta
argud that th rmarkal proprtis f th hydrothrmal vnt nviron- nd poposd tht rh
mnts, partiularly thir prottion from th harsh onditions ausd by o m t s .a n d i n t r p l n
larg-impatvnts'might hav playd an imprtant rol in th origin of lif. a n t o l i n s d i n gt h
Sin it is thought y som that th last ommon nstraloganism of all x- gin of lif. his on
tant lif on th was a thrmophil' svral rsarhrshav proposd th ompositionof mto
hypothsis that th organi ompounds nssaryfor th origin of lif wr imptsn th arly
atually synthsizdundr vnt onditions (Holm and Andrsson 1995). a. i t d i f f i u l t f o r I i f t o o
jo proponnts of this hypothsis ar vrtt Shok and oworkrs, who hav raw mtrial nssar
alulatd that thrmodynami.asdquiliria favor th formation of om- v i w t h i s h y p o t h s i s
pounds suh as mino aids at hydrothrmal vnt tmpraturs(Shok 1990; l i f i n t h u n i v r s .T
Shok and Shult 1998). Vnt-asdsynthsisis onsidrd to spially for th origin of lif
important in vnts assoiatd with off-ais systms (olm and Charlou plant; rathr' synth
2001; Klly t al.2001). Howvr, at th high tmpraturs(> 350") asso- raks pla on primit
iatd with most vnt dishargs,amino aids and othr iomoluls hav Th possiilityof th
n found to rapidly deompos (Brnhardt t al. 1984; illr and Bada t h s s n t i a lo r g n i t
1988; Bada et a|. 1995). For ampl, amino aids ar dstroydin timsals l fo furthrhm
of minuts at tmpratrrrsgratr thn 300.. Th rat of hydrlysis for a r o n o u s h o r
RNA at pH 7 trapolatd to lvtd tmpraturs givs a half-lif of 2 min- th most primitiv o
uts at 250. for th hydrolysis of vry phosphodistrond; at 350. th omposition. yr th
half-lif is 4 sonds. For DNA, th half-livs for dpurination of ah nu- % wight in son
lotid at pH 7 ar narly th sam as th hydrolysis rats for RNA (illr o r g n i o m p o u n d s i r
and Lazano 1995). lt has n pointd out y Lazano (t997) that ifth ori- r i a ' A u s t r l i )a n d M
gin of lif took pla ovr a suffiintly long priod of tim, all th mpl o r g u i l m t o r i ( l F
organi ompounds in th oan' drivd from whatvr sours' wuld CM a aottt 70" f
dstroyd by passag through th hydrlthrmal vnts. It is thus possil that grind mtril. Bt

produdin tlr rtions,as wll s
th lorv pH lrrditionsgnratddur-
1 aront to th system to uffr tlr
dation inhiitossuh as asori aid
:h rvryof svral hundrd tims
rusly.
rimnts,th amounts of fr yanid,
gstingthat th intrmdiats may
ids may hav fonrly svral mh-
or th rlatdBtrhrr-Brgspathway.
thir rltivaundnsar si-rilarto
th slf.orrdnsation of HN in aqu-
1. 1978|, th asnof polymri ma-
:ardthis mhanism. primnts with
Shlsingrand illr 1983) suggst
r H, in nutral atmosphr simulations
:tion of amino aids.
:iatdwith th dirt illr-Ury typ
rnt hypothsisfor th abioti synthe-
posd.This stiggstionsultd from
nal vnts.A group <lfsarhrshzrv
.s of th hydrthrmal vnt nviron-
rom th harsh onditions ausd y
an imprtant ol in th origin of lif.
ommon anstralorganism of all x-
svralrsarlrrshav proposd th
ls nssaryfor th origin of lif wr
o n s ( o l m n lA n d r s s o n1 9 9 5 ) . a .
vrrrt shok nd or,l.orkrs,whr hav
quilirifvo th ormation of om-
rmal vnt tl-nprturs (Shok 1990;
;ynthsisis onsidrdto spially
lff-axis systn1s(olm and harlou
th high tmprturs(> 350") asso-
no ids nd otl-rriomoluls hav
rnhardt t al. 1984; Millr and Bada
rmino aids ar dstro1.d in timsals
n 300"' Th rt of hydrolysis for
tmratursgivs a hlf-lif of ) mirr-
ry phosphodistond; at 350.C th
Lalf-livsfor dpurination of ah nu-
s th hydrolsis rats for RNA (Millr
l out y Lazcano (1997) that if th ori-
y long priod of tim' all th ompl
vd from whatvr sours' would
othrmalvnts.It is thus possil tht
h ()rigi of L-ii iL)
hydrothrmal vnts r ruh mor fftivirr gulating th onntratiol-t
of ritialorgrrirrrtllulsitr th oans rhirIrirr pl1,irrg signifiantrtll in
thirdirts\,'nthsis.
Dspitth prolms with staility at higli tmpraturs,th vtr-itl]trd
svnthsis thor lrs n hampiond lry Hrrlrr and Wihtshiiusr(2006),
who hav dsridhav primnts limd to ..narrow th gap btrvn
iohmistry and volani gohrnistry.,' Usirrg high onntri-rtions
of HN
(0.2M) and high pssursof Co (7.5ars) in tlr prsnof Ni/F.atlysts,
a varityof amino irrrlhydroyl aids ar prt>lul at 100.. Howvr, thr
ronrnsaor-rt t rlvanof this typ of primntto th natural gtl-
hmialnvironnlntstht w.ould ptd to hlrv istd ln th rlv
arth'spiallyth high onntrti()llsand prssursof strtir-rgrllgnts
( B a d a t a | . 2 0 0 7 ) .
Did Priotiorgani ompounds om from Spa?
Th diffiultisstlpposdly involvd with th ndognous, rth-sdsl.rr-
thsisof mino aids nd nulossld to t dvlopmntof altrnatir's'
In hrly 1990s lr1'airnlSgan (1992| nirlvzdor6's 1961 prlposerl
on th rol of lrntrynuli as sours oi voltils ftlr th prinririv arth
and proposd tht th xognous dliv1,of <lganimattr y strlids,
omts'and intrplntarydust partils (IDPs) ould hav plyd a signifi-
antrol in sdingth arly arth with th lmpoundsnssryfor tlr ori-
gin of lif. This onlusiln was sd on knowldg aout th ogani
ompositionof mtorits.It is important tO llot that if this onpt is virlid,
impatson th aly arth not only ratd dl.llstatingorrditionsthat ma]
itdiffiultfor lif to originat ut also t th sanl tim prhaps dlivrlth
rawmatrialnssill.vto st th stagfor th lrigirlof lif. In n vn .ilr
viwthis lrvpotlrsisould hav prlflr"rrrd irrrpliations|:orth irbtrrr{arr
of
lifin th ttnivrs.Th sour of th ssntialorgani ompoun1srqird
for th origirr of lif is not onstrind r'th onditions on a pz1rtiular
plant;rathr, svnthsisof organi ompottnds is r"rbiquitouspross llat
takspia on primitiv solr-systmodis suh as astroids and otnts.
he possiilityof th origin of lif is thus tlnsidraly inrasd,providd
thssntialrrrgniompounds ar dlivrd intt to a plant tht is suit-
ablfor futhrhnrilvolution. This is t to fully drnonstatd.
aronousllon]rits,a lass ()f st()nvlItOrits,ar rrnsidrdto
th most prir-rritivljtsin tlr solzrr svstm in trms of tlrir lrrrrrtl
omposition,yt thy hav a high urrdanlf lrgani aron. 1()rthan
3% by wiglrt in som ass. Th mtoritslTlosttnsivly analvzd for
organiompounds inlud th Ms urhison (r,vhihfll in 1969 tn Vitl-
ria,Australia)nd Murray (19s0, Kntuk1',Unitd Stats)and th I-lass
orguil mtorit(1864, Frn). ( and I rfr to typs of hondrits.
ar aout 709z"fin-graindmatrial and I ar alrnost lusivly fin-
graindmatrial' Both typs hv prin.ltrtsivaquOus ltrtit.ltl
60 Th rigi of Lif

on th parnt astroid.)Th aron phas is dominatd y an insolul fra. monomrs. Th trm

tion. Th majority (up to 80%) of th solul organi mattr is mad up of nin' rfrs not n
polyyli aromati hydr<larons(PAHs), aliphati hydr<rcarons,aroyli phr whr th aun
aids, fullrns' and amino aids (Bott and Bada 2002). Th purins ad- synthsis may hav ta
nin, guanin' anthin' and hypoxanthin hav also n dttd,as wll mnts' intrtidal zons
s th pyrimidin urail in onntrations of 200 to 500 parts pr illion going wt.and-dry y
(pp) in th M hondrits Murhison and Murray and in th I hondrit ponds.
Orguil (Stoks and Shwartz 1977,1981). In addition' a varity of othr Simpl organi om1
nitrogn-htroyliomplunds, inluding pyridins, quinolins, and iso. is of watr would n
quinolins, wr also idntifid in th urhison mtorit (Stoks and nhan polymizati
,|982),
Shwartz a s w l l a s s u g a r a i d s ( p o l y o l s )( o o p r t a l . 2 0 0 1 ) a n d sorption of moluls r
mmran-forminglipidi ompounds (Damr and Pashly \9s9I. of pomoting polymr:
It has n found that th Cl-typ mtoritssuh as orguil ontain n ortory primnts tl
amino aid omposition distint from that of th Ms (hrnfrund t al. vatd monomrs (Hill
2001). Th simp[ amino aid mitur, onsisting lf just glyin and rami minral-assistdat1
alanin, found in I aronaous hondrits is intrsting aus it has p o l y m r i z a t i o no f a t i
n gnrally thought that a Wid vrity of amino aids wr rquird for 25_50 mr oligonul
th origin f lif. As disussdlatr, howvr' among th andidats for th rd nssaryfo prir
first gntimatrial is pptid nuli aid (PNA), a nuli aid analoguin Baus adsorptior
.Waals
whih th akon dos not Ontin sugar or phosphat moitis (gholm intratins, th
et a|. 19921Nlson t l. 2000). F.or th PNA akon, ahiral mino aids polymrization would
suh as glyin or aminothylyglyin, possily dlivrd y I-typ ar- l998; Sowryt al. .
onaous honditsto th arly rth' my hv n th only mino aids ausas th lngth
ndd for th orisin of lif. tnd to mor firml'
plymrs to invoh
monomrs' thy nd
Prioti PolymrizationProsss th minral' lthough
mrs. A way around
Th organi mtrial on th arly arth for lif istd, rgadlss of its ntratd salt solutio
sour' likly onsistd of a wid array of diffrnt typs lf simpl monomri tidal gions during
ompounds' inluding many that play a major rol in iohmistry today. Th dirt onntl
How ths simpl aioti organi onstituntswr assmldinto polymrs omplishd y vap
and thn into th first living ntitis is prsrrtlyth most hallnging ara of tions, whih, ouplr
rsarh on th origin of lif. adhrn of iohn
Simpl monomrs on th primitiv earth would nd to undrgo polymeriza- al onntrations at
tion, a thrmodynamially unfavoral pross. It is gnrally assumd that rgions and th sut
polymrs omposd of at lst20 to l 00 monomri units (20 to 100 mrs)ar hv n proposd
rquird in ordr to hav any atalyti and rpliation funtions (Joe 202; uls (Nlson t al. j
d Duv 200). Thus arly polymrizationprosssmust hav napalof also n found to 1
produing polymrs of at last this minimum siz. 2000). Ths sam
Thr is no vidn of aiotially produd oligopptids or oligonu- rol in th synthsis
lotids in mtorits, so ondnsation rations larly must hav takn lut solutions of at
pla dirtly on th primitiv arth. Synonymous trms lik ..primordial amounts than in x1
roth'' or ,.Drwin's warm littl pond'' hav ld in som ass to major mis- 25. (Liu and org
undrstandings,inluding th simplisti imag of a worldwid oan, rih in tti frzing is sp
slf-rpliating moluls nd aompanid y all slrts of iohmial lotids (Kanavario

rhasis dominatd y an insolul fra-
r solul organi matt is mad up of
\Hs), lipatihydroarons' aroyli
iotta zrrrdBada 2002). h purins ad-
Lnthirrhav also n dttd,as wll
.ations<lf20
to 500 parts pr illion
on and Murray and in th I hondrit
1 9 8 1 ) .I n d d i t i o n , a v a r i t y o f o t h r
Llirdingpyidirrs,qirrolins,arrd iso-
th lvIurhison mtorit (Stoks arrd
: i d s ( p o l y o l s )( o o p r e a l . 2 0 0 1 ) a n d
l s ( D a m ra n d P a s h l y1 9 8 9 ) .
) mtOritssuh as Orguil ontain an
m thar of tlr s (hrnfrund t al.
'' onsistingof just glvin
nd rami
hondritsis intrsting aus it has
arity of amino aids wr rquird fo
howvr,among th andidats for th
id (PNA)' a nuli ;ridanalogLrin
n sugr or phosphat moitis (glrolm
hPNA kon,ahiral amino aids
n, possily dlivrd y I-typ ar-
'th,may hirvn
th lnly amino aids
sss
rth ftl lif xistd, rgadlss of its
r of diffrnttyps of sirnpl monomri
ry a mjor rol in iohmistry today.
Lstituntswr assmld into polymrs
s prsntlth most hallnging area of
rth would nd to urrdrgopolymriza_
l pross' It is gnrally assumd tlrat
)0 monomriunits (20 o 100 mrs) ar
: and rplition funtions (Joy 2002;
:ion prrsss must hav n apal of
rimum siz.
y produd oligopptids or oligontr-
ion retions larly must hav takn
. Synon1.moustrms lik ..primordial
l . . h v l d i n s o m a s r s t t - tm a j l l r m i s -
-i inr:rgof a worllwil oan, rilr in
nparrid y all sorrs of iohmi:rl
ll rigi o|.Lii 61
monomrs.Th trm rm littl pod, wih lras long n usd for onv-
nin,rfrsn()t nssrilyto th ntir oa ut to parts of th hydros.
phrwhr th umulation rr.lintration of th produts of prioti
synthsismy lrv takrr plzr.lrs inlr"rdoani prtils and sdi-
mnts'intrtidalzons, frsh,trshallow plrrdsand laks, lagoons ndr-
goingwt-and-dryyls, and utti nvironmnts assoitdwith glaial
ponds.
Simplorgani lmpoundsdissolvd in th prirlritivons o[ othr od-
isof watr wlr'rldlrd to otlntratd sotn mhanislrl in ordr to
nhanpolyrnizationprosses.onntr:rtioninvrlvingth sltivad-
sorptionof molulsOnto minral surfashas n suggstdas on mns
of promotingpolvmrization' and this pross as n dmonstratdin l-
oratoryprimntsthat hav usd a varity of simpl ompounds and ati_
vatdmonomrs (ill t l. 1998; Fris 2002). Tlr potntial importan of
minral-assistd atalysis is drnonstratdy rh montmorillonit-promotl
polymrizationof ativatd adnosin and uridin drivativsthat produs
25-50 mr oligonulotids(Frris 2002), th gnral lngth rang onsid.
rdnssayfor primval iohmial funtilns.
Bausadsoption onto surfrsinvolvs wk nonOvalnt van dr
Walsintrations,th minrl-asdonntration pross and susqurrt
polymrizatiorr wluld most ffiintat ool tmpraturs(Liu and orgl
1998;Sowryt al. 2001). This, howvr' prsntssomwhat of a prolm
ausas th lr-rgth of polyms fomd on milral surfsitrrass,thy
tndto b mor fimly lund to tlr minral (orgl 7998)' In ordr for ths
polymrsto involvd in sr.rsqunt intrratilns with ltlrrpolymrs or
monomrs'thy nd to rlasd.This oul1 aomplishd y warming
th minral,although this would also tnd to hydrolyz th adsord poly-
mrs.A way round this prolm would to rlasth polyrnrs y on-
ntratdsalr sllr'rtions (Hill t al' 1998), a pross that ould t:rk pla in
tidalrgionsduring vaporzrtionor frzillg tlf swtr.
Th dirtnntrationof dilut soiutins lf monomrs ould also a-
omplishdy vaporation and y utti frzing of dilut aquous solu.
tions'whih, oupld with othr physiohmial mhanisms suh as th
adhrn of iolrmiallonotrrsto ativ sf;rs, ould lrv risd lo.
l onntr:rtionsnd protrrotd polyrrizzltion.h vllporation of tilal
rgiona s n d t h s u s q u n t o n n t r e t i o n< l f t h i r o r g a n i O n s t i t u n r s
havn proposd in th synthsis of a varity of simpl ogani mol-
u l s( N l s o n t a l . 2 0 0 0 ) . u t t i f r z i n go f d i l u t r a g n t s o l u t i o n s h a s
also n found to promot tlr synthsis of k,v iomoluls (Lvy t al.
2 0 0 0 ) .T h s s a m O n l l t i l t i ( ) n p r o s s s l s o o u l . li r v p l a y d k y
rolin th synthsis of polyrrrrs.It hs n shown that tlr frzing of di-
lut solutionsof ativatd mino aids at _20.C yilds pptids at highr
amountsthan in primnts with highly onntratd solutions at 0o and
25" (Liu and orgl 7997)' ln alditiln,rDtstudis hav slrown that u_
ttifrzingis spiallyfftivin th nor-rrrzymati syllthsisof oiignu-
lotids ( K a r r v r i o ti t a l . 2 0 0 1 ) .
62 Th rigi of I,if

Salt rins mv also hav rrimportnt irr tlr formtiorrof ppids:rnd dns monolayr of ioni'
prhaps othr important iopolymsas ,ll.Aording t<lRod and owork- of pyrit. ln addition,
rs (Rod 1999),..sit-indu] Pptid formirti<lnration'' (SIPF)provi1sa shusr amino aid_ppt
pathway for th ffiintsynthsisof pptids and possilv protins dirtly likly to hiv format
for simpl anrino aids in lnntratdNlll solutilnsotrtaining u(II). fftiv autoatalysis pr
Yilds of di- and tri;lptidsin tl-rrang of 0.4 to 47o l. n rpotdus- ory" (Ross 2008).
ing starting arrrinoaid onntrationsin th 40_50 m N,I ang. lay rl-riner. Non of ths xperill
als suh as montmorillonit aPParntly promot th rtion. Again, th aids ar th volutionar
vaporation of ti1allyflushd ltrgoonsr th frzingof th primitiv oans th rsults ar also oml
ould hav produd th onlltrirtdsirlt-rilrrins nd1 t()promot this primordial souP in whi'
salt-indud pol,vmrizatilt-l
ptlss. sour of ltrons for tI
bl that undr rtain g(
hav rdud not only
Did Pyrit Ply a Rol in th origins f Lif on arth? dp-savnts, and ld l
stan. ould hav takr
.tO?ihtrshlusr
Ft>rsom tim, one of th most srius rivals o th ,.primordial soup'' ht- and 19
rotrophi thory was drivd frm th ids of $7;ihtrshziusr (1988).A- hargs via a N4illr-typ
orling to his ..pionrmtaoli thory,'' lif gan with th apparanf ss did not rmain ol
n aLltoatalytitwo-dimnsiotr:rlhmlithorrophi mtalli systm that surrounding aquous n
took pla on th surfaof th highly insolul minral pvrit in th r,iiniry ioti soup, not of a tw(
of hydrothrmal vnts.Rpliation followd th apparanof this nonorgan. Th ssntial qustior
isml iron sulfid*asdtwo-dimnsi<lnal..lif,''in whih hmoautotrophi not whthr pyrit-mdi
aron fixation ttllk pla y a rdutiv iri aid yl, or rvrsrlrs CO, rdution nd syntl
vl' li rh typ origirrallvdsrid{or th photosnthtigrrrsirlfirrlra. pothtial two.dimnsi<
tIitlm hlorobium limiril' Molular pvlognti trs show that this Proof of Wihtrshius
mod of aron fixatilr-r and its modifiations (suh as th dutivatyl- tight oupling of th r
oA or th rdutiv malonyl.oA pathwa,vs)ar flund in anaroi zlr- t i o n v i a r d u t i v i t r i
ha irnd th mtlst dply divrgrrtutri:r'whih has n intrrtd of homologotls yls th
s vidn o[ its primitiv iratr(aln 1995). But is th rvrsKrs wll as rpliation (a
yl truly primodiirl? imntal sults ahivd
Th rtion FS + H,S = F.Sl+ H, is vry nrgtialli,favoral"It has and Co ar onsistnt !
an irrvrsil,highlv rgoni haratrwitlr a stndard fnrgyharrg th rol of slfur-rih
of (;., = -9.2 ka|lrol, whih orsptrdsto rdtrtion potntial ^ = mulation of organi or
-62m ). Thus th S/H,S omination is a strtlng rduing agnt nd has
shown to provid an ffiint sour tlf ltrons for th rdutiln of or-
gani ompounds undr mild onditions. Pyit formation an produ mo- How Did Rpliati
l t r l a rh y d r o g n : r r r dr d u i r r a t t o a m m < l n i aa, t y l n < l t h y l n ,a n d
thioti i.lttl ati ail, .rs wll as L)olrrotingmor ompl syDthss Th pimordial roth r
(adn 1995), inluding pptld lnds tht sult fronr th ativatio of drland in whih a wid
amino aids with aron monid and nikl/iron sulphids (Hur and sizd' dstroyd, or in
\/dlrtrshiusr1998). Although pyrit-mdiirtdO, rdLrtionto orgtri Rgardlss of th orp
ompour-rdshas not n ahivl,th fiatil undr plzrusilepi<lti ould not hav volvd
onditions of aron monoi]into ativat]ati aid l.lva mitur f o. guarant th maintnl
.Wihtrsh;ir-lsr
pripitatd NiS/FS/S has n rportd (u and Ponnts undr th at
1 9 9 8 ) . H o w v r , i r r t h s p r i m n t st h r i o n so t l r i t l n e q u o u s n . molular ntitis a]
viltlnrntto whih powdrd pr,rit has rrddd; thr, do not form a would hv markd tl

:
5

otntin th formation lf pptids and
rs wll. Aording to Rod and owork-
formatin rtion'' (SIPF)provides a
pptidsand possily protins dirtly
ltd Nal solutions ontaining u(II).
g of 0.4 to 4o ae n rportd us-
; in th 40_50m rang. lay minr-
rtly promot th ration. Again, th
o r t h f z i n go f t h p i m i t i v o
salt.rihrinsnddto promot this
)riginsof Lif n arth?
rs rivals to th ..primordial soup'' ht-
. i d a so f W i h t r s h i u s r ( . l 9 8 8 ) .A -
lry,'' lif gan,ith th appran of
molithotrophi mtoli systm that
insolulminral pyrit in th viinit
owd th apparanof this nonorgan-
nal ..lif,''in whih hmoutotrophi
:tivitri aid yl, or rvrs Krs
Or th photosynthti grn sulfr.rra-
lar phyiogntitrs show that this
fiations(suh s th rdutiv atyl-
)athways)r found in anroi ar-
:utria'whih has n intrprtd
Madn 1995\. But is th rvrs Krs
, is vry nrgtiallyfavorl. It has
]trWith stndard fr nrgy hang
sponds to a rdution potntial ., =
tion is a strongrduing agnt and has
:of ltronsfor th rdution of or-
ns. Pyrit formtion n produ mo-
l ammonia, atylnto thyln, and
ts promoting mor ompl synthss
rds that rsult from th ativation of
rnd nikl/iron sulphids (Hur and
t-mdiatdo, rdution to organi
h fiation und plausil prioti
:tivatdarti aid y a mitur of o-
lortd (Hur and $7ihtrsh:iusr
th rationslur in aquOus n-
has n ddd: thy d<l not f()rm a
h rigi of Life o.)
dnsmonolayr of ionially ound moluls or tak pl on th surf
of pyrit.In addition' a arful aminatiorr of th Hur and !zihtr-
shiusramino aid-pptid synthsisshm hs found that it is highly un.
likly to ahiv formation of oligomrs that had th aility to promot
..pionrmtaoli th-
fftiv utoatalysisprosssassoiatdwith th
o r y " ( R o s s2 0 0 8 ) .
Non of thsxprimnts provs y itslf that oth nzyms and nuli
aidsar th volutionary outom of surfa-ounddmtaolism. In fat,
a n 5 th rsults ar also omptibl with a mor gnral' modifid modl of th
primodial soup in whih pyit formation is rognizd as an important
sourof ltronsfor th rdution of organi ompounds. It is thus possi-
ltht undr rtain gologial onditions th FS/H,S ombination ould
havrdud not only o ut also O. rlasd from moltn mgn in
dp-savnts, and ld to iohmial monomfs. Pptid synthsis' for in-
stan,ould hav takn pla in an iron and nikl sulfid systm (Hur
and Wihtrsh:iusr 1998) involving amino ids formd y ltri dis-
hargsvia a Millr-typ synthsis.If th ompounds synthsizdy this pro.
ssdid not rmain ound to th pyrit surfa ut driftd away into th
surroundingaquous nvironmnt, thn thy would om part of th pr-
iotisoup, not of a two-dimnsional autoatalyti ration shm.
Th ssntial qustion in diding twn ths two diffrnt thoris is
not whthr pyrit-mdiatd organi synthsis an oLlr' ut whthr dirt
o, rdutionand synthsisof organi ompounds n ahivd by a hy-
pothtial two-dimnsional living systm that laks gnti information.
Proof of !7zihtrshiusr's hyplthsisrquirs dmonstration not only of th
tightoupling of th rations nSsryto driv autoatalyti O, assimila-
tionvia a rdutivitri aid yl ut also of th intrwavingof a ntwork
of homologousyls that, it is assumd,ld to all th anaoli pthways' as
wll as rpliation (adn 1995). This has not n ahivd.ln fat, pr-
imntalrsults ahivd so far with th FS/H,S omination in duing N.
and o ar onsistnt with a htrotrophi origin of lif tht aknowldgs
throl of sulfur-rih minrals nd othr atalysts in th synthsisand au-
mulationof organi ompounds.
How Did Rpliation First originat?
Th primordial roth must hav n a wildring organi hmial won-
drlandin whih a wid array of diffrnt moluls Wr onstantly synth-
sizd,dstroyd' or inorporatd into yls of hmial transformations.
Rgardlssof th omplity of th prioti nvironmnt' lif as w know it
ouldnot hav vlvd in th asnof a gnti rpliating mhanism to
guarantth maintnan,staility' and divrsifiation of lif's asi om-
ponntsundr th ation of natural sltion. Th apparan of th first
molulr ntitis pal of rpliation, atalysis' and multipliation
.What
would hav markd th orisin of oth lif and volution. wr th
64 The (.)rigiof Lif

furrdamntalhratristisof ths first mlIr-rlar living ntitisthat distin- th PNA-RNA and P]

guishd thr fr<lmnonlivirrg hmistry? paling as possil ar
Th lap from iohmil monlmrs nd small oligomrs to livirrg nti- titis, both PNA and
tis is an norlolls lap indd. How th uiquitlusgntisystm of tant ngativ fators ar t
lif asd on nuli aid originatd is ot-lof th major unsolvd prllms that would prdu t
in
ontmporay ioltlgy. Th disovry of atalytialIytiv RNA moluls onditions and th la
gav onsidralrdiility to prior suggstionsthat th first living PNA is attrativ l
ntities
wr largly asdon atalyti RNA mlluls(riozyms)'in ir hypothtial whih liminats th r
stag in th arly volution tlf lif alld th RNA wold (Gilrt 1986; origin of lif. Its orn
Joy
2002). This possiility is now widly aptd,ut th lirrritdstaility hav n produd ur
of th
omponnts of RNA implis that this molul was proaly not a dirt Howvr, PNA is sus
podut of priltivolutitln. rarrangd PNA. Thil
Th rilsomponent tlf RNA is vry trnstal,whih maks lts prsnr th N-trminal positi
i n t h p r i o t i m i l i u u n l i k l y ( L a r r a l d t a l . 1 9 9 5 ) . B o t h l a d h y d r o i d on th asis of an
( Z u a 1 9 9 8 ) a n d o r a t m i n r a l sa p p a r n t l ys t a i l i z r i o s ( R i a r d o
t al. Pars to suprior t.
2004), and ynamid is known to at with rios to form a stal iyli to its as-pairing attr
addr-rt(Springstnand.|oy2oo4). Howvr, in ordr t involvd in Shiningt al. 2000).
th
polymrizatitln rations that lad to RNA, rios would likly hav during th ration a
to
Prsnt in solution, whr it would pon to dmposition. Morovr, dsrid y Butlrov
th hug nurr of possil random ominations of drivativs lf nul- ould hav bn rad
trass,stt]rs'and phosphat that may hav n prsnt in th prioti (sRation 2), whi
soup mks it unlikly that an RNA molul apal of atalyzing its own tion of formaldhyd.
slf-rpliationros spontnOusiy(J.ly2,002).Ths diffiultishav ld rdus this prolm
to roposals of pr-RNA wlrlds, in whih inf<lrmationalmaromoluls oligonulotids om1
with akons diffrnt from thos of tant nuli aids may also hav sizd undr prioti r
n n1owdwith atalti ativity, that is, with phnotyp and gnotyp It is possil that P]
also rsiding in th sam rrrllul.Thus a simplr slf-rpliatormust hav to th fist TNA mol
om first, rrd svral possil tlntndrshv n suggstd.Th RN omponnt of TNA w
prursor would hv had th apaity to talyz rati()nsnd to stor in oprtion. org1 aI
in-
formation, although th onrponntnr-rloss and th akn that hld hiral sugar dinulot
th polymr togthrwr not nssariIyth sam as thos in modrn urrd simply y han
RNA
and DNA. Th natur of th grlri. poly,-,-'.,,nd th irtalyti agnts PNA.indud oligom.
that
may hav prdd RNA is prsntly unknown. providd mans of
Thr ar n<lwsvral known anrplsof rolular gnti systmsthat hirality into polymr
hv n studid in th laoratory, and ths pr<lviiampls of th tntially solvs an onf
ryps
of molular systmsthat ould hav givn ris to arly slf-rpliatingnti- for th origin of lil
tis (s Bda 2004). Possll andidats inlul nuli aid anaigus
( g h o i m t a l . 1 9 9 2 ; i l l r 1 9 9 7 ; s h n m o s r1 9 9 9 , 2 ( ) 0 4 ;
Nlson t al.
2000) suh as pptid nuli aid (PNA), whr th akon onsists
of
linkd amino aid drivativssuh as N.(2-aminothyl)glyinor (AG) (th o
..--..---.-.---
I{o f
nuloass r attahd y an ti aid linkag to th mino group
of
glvin), nd thrtlsrruliaid (TNA), whr th akon is mad Formaldehyde G.
up of
L-thosonntd y 3', 2' phsphodrstronds. Both of thsandidts
Retion 2. h forma
form doul hlil strlltursthrough !tson-rik as pairing with
om. formaldehydeand gly
plmntary strands of thmslvs,RNA, and DNA. '$7ithPNA-PNA
hlis of rations'and oth
oth lft. and right-handd strutursar prdud in qual mounts, susquntations.
whil

irst molulr living ntitis that distin-
ry?
rrs nd smali oligomrs to living nti-
th uiquit<lus gntisystm of tant
; on of th major unsolvd prolms in
. of atalytialiytiv RNA moluls
suggstionsthat th first living ntitis
noluls(riozynrs),in a hypothtial
d t h R N A w o r l d ( G i l r t1 9 8 6 ;J o y
aptd,ut th limitd staility of th
is molul was proaly not a dirt
.ry unstal,whih maks its prsen
l d t a l . 1 9 9 5 ) .B o t h l a d h y d r o i d
)parntlystailizrios (Riardo t al.
at with riostl fom a stal iyli
Howvr, in ordr to involvd in th
l RNA, rios would likly hav to
) pn t domposition. orovr,
r ombinations of drivativs of nul-
ny hav n prsnt in th prioti
molulapl of atalyzing its own
Joy 2002). Ths diffiultis hav ld
whih informationl mromoluls
of tant nuli aids may also hav
, that is, with phnotyp and gnotyp
hus a simplr slf-rpliatrmust hav
ndrshav n suggstd.Th RNA
:y to atalvzrations and to stor in-
ruloass nd th akon that hld
rrily th sam as thos in modrn RNA
polymrs and th atlyti :1gntstht
unknown.
rpls of molular gnti systms that
nd thsprovid ampls of th typs
givn ris to arly slf-rpliatingnti.
idats inlud nuli aid analogus
s h n m o s r1 9 9 9 , 2 0 4 ; N l s o n t a l .
]NA), whr th akon onsists of
N-(2-aminlthyl)glyin o (AG) (th
id linkag to th amino group of
IA), whr th akon is mad up of
,distrorrds.Bth of thsandidats
l !atson.rikas pairing with om-
jA, and DNA. With PNA-PNA hlis
ar produd in qual amounts' whil
The rigi of Life
th PNA-RNA and PNA-DNA hlis ar both right-handd. Although ap-
palingas possilandidatsfor th first slf-rpliatingmolular living n-
titis,oth PNA and TNA hav positiv and ngativ aspts. Th main
ngativfators ar th lak of ny dmonstratd oligomrization pross
that would produ ths nuli aid analgus undr plausil prioti
onditionsand th lak of any dmonstratd atalyti proprtis.
PNA is attrativ aus its akon is ahiral (lking handdnss),
whih liminatsth nd for th sltion of hirality for th tim of th
origin of lif. Its omponnts, AG and nuloass linkd to ati aid'
havbnprodud ndr simulatd prioti onditions (Nlson t al. 2000).
Howvr'PNA is stlsptilto an N-ayl migration ration that produs a
rarrangdPNA. This prolm ould minimizd, howvr, y loking
thN-trminal position y tylation' for xampl.
on th asis of an tnsivstudy of sugar-asdnuli aids' TNA ap-
pafsto b suprior to othr possibl sugar-asd nuli aids with rspt
to its as-pairingattriuts,spiallywith RNA (shnmosr1999, 20041
Shningt al. 2000). Th ttrossugar in TNA ould hav n synthsizd
duringth ration asad that taks pla during th formos ration first
dsridy Butlrov in 1861. Th 4-aron sugars thros and rythros
ould hav n radily synthsizd y th dimrization of glyolaldhyd
(sRation2), whih in turn ould hav n produd from th dimriza-
tion f formaldhyd. vn though th prsn of a 4-aron sugr in TNA
rdusthis prolm to 2 sugars and 4 stroisomrs' to dmonstrat how
oligonulotidsomposd only of L-thros ould prfrntially synth-
sizdundr prbioti nditions is a formidal hallng.
It is possil that PNA prdd TNA and in fat assistd in th transition
to th first TNA moluls. As statd arlir, th sltion of th hirl sugar
omponntof TNA would hav rquird som sort of sltion proess to
in opration. orgl and oworkrs hav suggstd that th inorporation of
hiral sugar dinulotids at th nd of a PNA hain' whih ould hav o-
urdsimply y han, an indu hirality into a nuli aid produd y
PNA.indud oligomrization (Kozlov t al. 2000). PNA ould thus hav
providda mans of onvying th ritially imprtant iologial proprty f
hiralityinto polymrs nar th tim of th origin of lif. This possiility po-
tntiallysolvs an ongoing dilmma whthr th origin of hiality ourrd
forth origin of lif r during th volution of arly living ntitis.
HCo'o?-Ho 9Sr91o Io2-FIoH.Co-CH2oH
tlo
Formaldehyde Glyolaldhyd
Reation2. Th formation of ttrossugars from th dimrization of
formaldhyd and glyolaldhyd.This is nly part of the ovrll formos sris
and
of reations, ot hxossnd pntosswould hav n produd y
subsquent rations.
h ()rigitt f 1,if

Rgardlssof th typ of analogu simiIar to nuli aid or othr typ of th asi mahinry
rpliator sstm that ws usd y tl-rfirst self-rpliatingntitis, polvmr .
partmntalizd into
staility and survival would hav n of ritial importan. Nuli aidsin Dspit th susl
..with
grrralhv vry shrtsurvival tims at lvatdtllpraturs. RNA is r,r lms with this
unstabl ausof th prsnof th phlsphldist ond involvirtgth 2'- ordr to maintain a s
ydroyl group of ris. Th staiIiryof TNA is unkrrorvnat prsnt' ut it ating systm insid
is proaly somwht similar to th staility of DNA. It is possiltlrat otlrr layr barrir. This
ttros sugar-asdnuli aids wr lss stal thirn TNA usof thir as amino aids. As a
tndny ttl for lss stal hlial stfrrtul]s(snmtlsr|L)99|rrd tlrt sid th vsil'' (Gr
th sltion of TNA Was th rsult of this fatlr,althotlgh this is also not surfs ould hav
known. Tlr stility of PNA lrs n prtly invstigatd,and providd th ules that usd th z
N-ayl migration ration an rninimizd, th mid linkag in PNA volvd in polymriz
rvould xptdto hav a stailit at nutral pH similar to that of pptid omplxity, synrgis
honds in pr<ltins(.Wng1998). This suggststht in nr,ir<lnmnts rvith tm- of th omponnts il
pratursof around 25", its survival tim wotrld in th rang of 104-105 portd into th vsi
,vars.Salry'rins trray lrav playd a rol in ariy nuli aid suvival - formd a singl om
aus high salt onntrations hav n found to nhn nuli aid st- rsults showing that
bility. For mpl, th staiiitis lf svral transfr RNAs (tRNAs) wr om polymrizd
signifiantl.vinrasdin |-2 Nl soluti<lnin omplrison rvith staility pidi arri may no
in pur water (Thit al. 2002). Th staility of DNA also inrasswith in-
rasil-lgsalt onntration (fo ampl, sBda 2004).If thsrsultsr,ver
ppliale to othr nr'rliaid analogus suh as PNA nd TNA, thrrsalt Th Transition l
solutions ould hav providd a prortiv nviront-trnt that ould hav n-
h:rndth survival oi arl1'slf-plitingmolular ntitis. Th volution of th
-h
$rmmrns rquird for th mt:gnf rpliating systms? tirly on RNA (i..
mrgnof lif m-v st irndrstoodin trt-t-ts of th dynaIlis and vo- volution toward m<
lution of sts of hmial rpliirtingntitis.\7hthr suh ntitis Wrn- on molul that
..t
llsdwithin mmrns is not yt lar, ut givn rh priori availallity Laoatory-basd
of amphiphili ompounds, this rn,vhav wll n th s. Mmrn- atalyti RNA mol
forming lipidi ompounds may hav n providd to th primitiv rth rang of important
from xtrtrstrials()Llrs(Dmr and Pashl 1989). This sotlr ould RNA fragmnts (lig
potntially supply a wid varity of aliphati and aromati hydroarons,al- 1999|. Th list of dt
lhols, and r;rnhd arrd straight fattv aids, inluding som that ar that th RNA world
nrman-formingompunds. Prioti lipidi moluls ma1,lso hav r. uls, prhaps fun
sultd frorlr aioti synthsis, ars shtlwn y th formatiln rrf nornral ftty 2002). Th omplx
ids, glyrol. glyl phosphat, and tlrhr tipids (iherg t a|. t977\. ti transfr, nd vaI
Also, primntshav shown that simpl miturs of fatty aids and glyrol dmonstratd in th
form fro.l miturs <lfmt>no.,di-' nd triglridsur-rdrild lnditions immatuity of tlris
(Hargravst l. 1977)' It is vry atrrativtO assul tht orpartmntal- Ch 2002; Joe2|
ization within liposot-llsformd y amphiphili molulsof prioti origin Th invntion of
\vs ssntiillfor th mrgnof lif. RNA nr<lltrls lsordorlt las hiny ndd for l
suh as montmorillonit' whih an atalyZ th formation <lf RNA Four of th bsi r
oligoms, an e napsulatdinto fattl, aid vsilsrvhos forlltion in ozyms, and it has
turn is alratd y th lay. By inorporating additiorral fatty aid rni- tions is not likly a
lls' ths vsils an gow and divid whil still rtaining a portiln o{ oigin, most pra
tlrir ontnts ndd o sLlpprt RNA rplition. In this mannr' som of th as' thn th r

similar to nuli ail or othr typ of
6rst slf-pliringntitis' p<llyrrrr
lfritial imp<lrtarr. Nuli aids in
at lvatdtmpratus.RNA is vy
: phosphodistrond involving th 2.-
g of TNA is unknowtl t prsnt, ut it
ailitv of DNA. It is possil that t>thr
lss staltharrNA ausof rhir
rllturs(shnmosr1999) and tlrat
lf this fator, altough this is also not
l ptly invstigatd,and providd th
'inirnizd' th amid lirrkag in PNA
tt nutrlpH sinrilar to that of pptid
uggststhat in nvionmntswith tm-
til would in th rang of 101_105
rol in arly nuli aid survival -
n found to nhan nuli aid st-
svrltransfr RNAs (tRNAs) wr
solution in ompzrris<ln with sterility
;tirbiIityof DNA also inrasswith in-
I 's B d 2 0 4 l . l ft h s r s u i sw r
gussuh as PNA and TNA, thn salt
tiv nvironmnttl]t ould hav err-
rting molulrnritis.
:mrgnof rpliating systms? Th
lod in trms of th dynamis and vo-
n t i t i s .W h t h r s u l l n t i t i sW r n -
:ar, ut givn th prioti vailaility
hv wll n th ils. mran- atlytiRNA moluls
n providd to th primitiv arth
n d P a s h l y1 9 8 9 ) .T h i s s o u r o u l d
phati and romati hydroaons, l-
[amy aids' inludig som that ar
>tlliidi molr-rls mv als<lave r-
vrr y th fornl:rtitlt-tof ntlrrrlal fatty
L d o t h r l i p i d s ( i h r ge t a l . 1 9 7 7 ) .
pl miturs of fatty ids and glyrol
rd triglyridsundr mild tlnditions
ativ to assumthat ompartmntal-
p h i p h i l i m o l u l so f p i o t i o r i g i n
RNA molulsdsod onto lilys
rr atalyze th f<lr:mationof RNA
ltty aid vsilswhos formation in
otporating additional fatty aid mi-
lid whil still rtaining a portion of
. rpliation.In tliis mannr, sonr of
Th rigin of Life 67
thasimahinndd for RNA slf-rpliatilnould hv Irom-
prtmntalizd irrto poto-typils (rizy t :rl.200; hn 2006).
Dspitth sussof th RNA napsulatioIrprimnts,thr ar pro-
..rvithin
lmswith this th vsil'' snario (Griffiths 2007). For mpl, in
ordrto maintain a stdy supply of moluls ndd for survival of a rpli-
atingsystminsid a vsil, moluls would nd to ross a lipid-lik i-
layrarrir'This rrlay hav n mirjor ostal fo hargd spis suh
..out-
as aminoaids.As a rsult' prlrirpsth first rpliating systm volvd
sid th vsil'' (Griffiths 2007). In this as, vsils attahd on minral
surfasould hv n in los proimity to sirnpl slf-rpliating mol-
ulsthat usd th atalyti proprtisof th minrl to nhan rations in-
volvdin polymrizationand rplition. As thstlr two systmsvolvd in
omplity,synrgistiintrirtiotrsould irav dvlod thar lllwd som
of thomponntsinvolvd in th rpliating systm to om frly trans-
portdinto th r,sil.At this point th two systms ould hav rrrrgdand
formda singl omartmntalizd, slf-rpliation systm. Howvr, rnt
rsultsshowing tht ativatdnulotidesan diffrrsinto liposoms and -
ompolymrizdwithin thm suggsttht undr rtain onditios tl-rli-
pidibarrirmy not hav n irnpntral(nsy t al. 2008).
Th Transition toward a DNA/RNA/Protin World
Th voiutionof th first livirrg rrrolularlir,ing rrtitisinto ons basd n-
tirlyon RNA (i..,th RNA rvrld) would lrav n th nt stp in th
volutiontoward modrn ihrnistry.RNA has n found to an all-in-
onmolulthat an not only stor information ut also atalyz rations.
..tst-tu-volution''primtrtshav dmonstatdthat
Laoratory-asd
(riozyrns) hav th pitv to ary Out a wid
rangof irportant iohnrial ations,irrluding th joining togthr of
RNA fragmnts (ligation) and pptid-ond formation (Bartl and Unrau
1999).Telist of dmonstratdatalyti rations is tnsiv ad suggsts
thatth RNA world ould hav had a lrg rprtoirof atalyti RNA mol.
uls,prhps fr"rtining in onrt with on an()thr(Doudna irnd Ch
2002).Th otrrplsrisof rtionsnddto prr-r.rit rnultiplition'gn-
ti transfr,and variation rqr.lirdin th RNA wrld has so filr not n
dmonstratd in th laoratlry,ut optimism rlrrainsbausof th rlativ
immaturityof tis ara of rsrh (Bartl nd Unrau 1999; Doudna and
h2002;Joy 2002).
Th invntion lf protin synthsisand th npsulation of rtioll ma-
hinryndd for rpliation n1y have takn p[ during th RNA world.
Fourofth asi ratitlnsinvolvd in protin ilsynthsisar atlyzdy ri-
ozyms,and it has n notd that th omplmntary natur of ths ra-
tions is not liklv aidntal ut rathr suggstiv that thy had a ommon
origin,most proably in th RNA wlrld (Kumar atr]Yarus 2001)' If this was
thas,thrrtlr primitiv nltloasod usd tor protin iosyntlrsislrad
Th rigi lf l,if

its origin in th RNA worll, lthough th b:rssusd in th early od ould Prthrmophili organt
hv en diffrrrtforn tll ons usJtody (Kol t l. 1994). modrn iology was a
It is possil tht y th tim RNA-asd lif appad<lnarth, th sup- DNA/protin-asd lift
plis of simpl ioti orgalli omportndsdrivd from th soursdisussd At first glan, oth
arlir would hav n grtly lirinishd.nr. of th lrponntsof th PPr to support a }
primordial soup had likly n tnsivl1.onvrtd into poiymrs, inlud- arly. Larg-sal anal
irrgthos ssoitdr,r,ithliving rrtitis,nd thtrsth rrv nl:ltrialsnddt of primitiv arth was
sustain lif had om largly hr.rstd. This inlplis tht simpl mtoli- rrmaind moltn for
lik pthwys must hav rrin plir in ordr t() nsur a supply of th in- (Nfthrill 1990), but n
grdintsnddto sustain th istnof th primitiv living ntitis.In this hydrosphr implis th
as som mtrllipathways ndd to produ sslltial omPonnts Howvr, thr is th
rquird y primitiv living ntitisWr prhaps origirrallynonnzynllttior wnt lat rtion im.
sminzymatiautoatalyti prosssthat ltr brfin tund as iozy- may hav oild th o
mati nd protir.r-sd nzmti poessing rltlmi rrnt. Could modrn bioh
ore than with rlier living rnolulrntitis,th main limittion in th ditions? Th proposals
RNA rvrld would hv n th xtrm instllilitv of RNA. This i turn lms, inluding th h
implis that RNA m<llulsmust hav n vry effiint in arrying out ompounds suh as ar
slf-lpliatiOnationsin ord tl t-nintainarr adquat invntory of mol- moluls, whos ha]
uls ndd ftrr survival. Th instiIity of RNA otlld hv n th pri- 250_350. ar at th n
mary rson ftl th transition to th DNA/prtltin world, tvhr, usof It is tru, of ours,
rh inrsdstility of th gntimoluls,srrrvivalwould hr,nlss fstr. Primitiv nzyI
dpndnton polym staility.Aording rlJoy (2002)' it is possilthat nd thus th rat nha:
irr th RNA rvorlliozyms ros tht ould talyz th pllyrnriztionof on way to ovrome
DNA, nd in this mannr information stord in RNA ould tansfrdto rizd lswhr (Islasi
th mor stal DNA. Anothr rsotl for DNA takovr ould hav n
1. rdud onntr
that aus of inrsd staility muh longr oligomrs ould hav au-
and NH.;
r-rruiatd.his rvould hav providd n nlrandstor:rgapaity fo infor-
2. lowr Stady.stat
mation that ould passd on to th nt gnraion lf living niris.
whih at tmpra
Befor long, RNA, whih had on playd tlr singular rol tlf rpliatitrnand
frmamid and tl
atalysis,ws rplad y th mlffiintnd rousr DNA/prtin .orld,
3. instability of ra
rvhrinRNA wrrsdrnltdto a rol of mssngr/transrir of DNA-stord (RFIo(Nt{,)
irrformtion ndd for rtltin iosvnthsis.
o f a m i n o a i d s ;a
4. loss of organi
s t a i l i t yo f g n t
Did Lif Aris in a High-Tmpraturnvironmnt?
Th rognition thz
vn tlrouglrDNA is nror st:rlthan RNA, it is still ridly dgrddat l- ar oupid y hyp
r,atd tmpraturs.In addition, Protin nzyms dnatur rapidly at lvatd itslf onlusiv pro
tmpraturs.This rtrsthr,at lst irtitillylimitd th nr,ironmntswh lif (Brohir and Phi
DNA/protin-asd lif ould surviv for anr. signifizrntprid of tim, nd as sriptions of th vo
was th as for ll othr arli living ntitissd tln nuli aid, sr.lrvivl biohmia1ompoul
would hav n most likly und ool onditiorrs(Bda and [,azno2002). bly niv to attmpt
Nvrthlss, sevral rsarhrslrav advoatd high t1praturs,spill living systms from
thos zrssoiatdrvith [rydrlthrnralvrrt systnrs,as th nr,ironmnt whr rl gntransfrof.
DNA/protin_asd ntitis first ros. Prlponntsfor high.tmprturOrt- misd th gnti r
gin it th f:tttht th trnivrsltr lf tI1tlif irpprst<l[.l<ltdin hv- onlusions aout th.

r th assusd in th arly od oLlld
ltody (Kolt aI.1994).
-sdlif irppardOn art' th sup-
rnds drivdfrom t sours disussd
ishd.:rrrylf th omporrnts of th
sivly onvrtdintrl polymrs, inlud-
s, and thus th rw mtrials nedd to
td.This irrrplisthat simpl mtoli-
in ordr tO nsur a supply of th in-
oi th prinlitivlir.ingntitis.In this
id to produ ssntial tlnrponnts
- ' r 1 . l r l i pt lqr i g i n l I l r n o n n z v m a t i o r
rlratiatr amfin urrds riozy-
ossingamdominnt.
ul ntities,th main limittin irr th
rrninstaility of RNA. Tlris in tr'rrn
r v n v r y f i i n ri n r r y i n g < l u t
rintainan lquatitlvntorr.of mol-
'ility of RNA ould hav n th pi-
)NA/potirr world, whr, aus of
oluls,sul,ir'aIlr'luldh:rl'lln lss
dirrgto.|oy(2002), it is p<lssiltlrat
lt ould t|yzth polvmization of
stord in RNA lr"rld trzlstrrdto
n for DNA takovr ould hav n
rh long oligomrs ould v tt-
n nhan{ storagapaitr,for infor_
h nt gnration of living ntitis.
yd th singulirrrllof rplition nd
intand roust DNA/prorin world,
{ mssngr/transritrr
rhsis.
rturnvironmnt?
RNA, it is still rapidl1'dgraldat l-
n nzymsdnturapidly t lvatd
ritially limitd th nvironmntswhr
l any signifintpriod of til, and s
ntitisasdon nr.rliaid, survival
onditions(Badaand I,zno20O2).
dvotdlriglr tnlpraturs,spi:rlly
nt systms'as th nvironmnt whr
P r o p n n t s o rl h i g , h - t m p
tnt lit appars to ro<rtdin lry-
Tbe rigi of Li|. 69
prthrmophiliorganisms. Thus if th lst om]on anstor (LA) of all
modrnilogy was a hyprthrmphil, thn it is onluded that tlr first
DNA/protin_asdlif nrust hav arisn in a similar typ of nvironmnt.
At first glan' oth th molular and th palontologial fossil rords
apparto support a lryprthrrophiliorigin of lif. Lit on arth aros
arly.Larg-sal analysis suggsts that son aftr its formation th surfa
of primitivrth was xtrml hot. Th plant is gnrlly thouglrt to hav
rmaindmoltn for som tim aftr its formation 4.6 illion yars ago
(Wthrill 1990|, ut minralogial vidneof a 4'- to 4'4-illion- yar-old
hydrosphr lmplis that its surfa apidly oold down (\r'ild t al. 2001,).
Howvr,thr is thortial and mpirial vidn that th plant und-
Wntlat artion impts (Byrly et a|.202; Sl-ronrg t al. 2002) that
mayhavoild th oans as lat as 3.8 illion yas ago (Slp t aL.|989).
ould modrn ihmistry hav arisn undr ths high tmprtur on-
ditions?Th proposls of a high-tmprtur origin of lif fa major pro-
lms'inluding tlr hnrial domposition of prsumd ssntial iohmial
ompoundssuh as amino aids, nuloass, RNA, and othr thrmllail
moluls,whos half-livs for domposition at tmPraturs twn
250-350"ar at th most a fw minuts (!(/hit1984;Millr and Bada 1988).
It is tru' of ours' tht high tmpratursllow hmial rations to go
fasrr.Pimitiv nzyms' on tlry appared, ould hv n inffiint
ndthus th rat nhanmnt ssoiatd with highr tmpraturs wouid b
onWay to ovrom this limitation (Harvy 1924), Howvr, s summa-
rizdlswhr(Islaset a|.207)' high-tmpraturrgil-tls would lad to:
1.rdudonntratiOnsof volatil intrmdiats'suh as HN, H2O,
and NH,;
2. lowr stady-statOnntrationsof prbiti prursors lik HN'
whih at tmpratursa littl aov 100. undrgoshydrolysis to first
forrnamidnd thn formi id arrd ammoni;
3. instabilityof rativ hmial intrnrdiatslik amino nitrils
of DNA-stord
(RHo(NH')N), whih play a ntal rol in th Strkr syr-rthsis
of amino aids; and
4. loss of rgani ompounds y thrmal domposition and diminishd
stailityof gnti polymrs.
Th rognitionthat th dpstranhs in rootd univrsal phylognis
aroupidy lryprthrnophilsis ontrovrsil and dos not provid y
itslfonlusivproof of a higlr.tmpraturorigin of DNA/protin-asd
lif(Brohirand hilipp 2002). Givn th hug gap xisting in urrnt d-
sriptionsof th voltrtionarytransition twn th prioti syr-rthsis of
iohmialorrrpounds and th LA lf ll tarrt living ings, it is pro-
ly naiv to attrmpt to dsri th origin of lif nd th natur of th first
livingsvstemsfronr nrolularplrylognis(Islaset aI. 2007).In addition, lt-
ralgntransfr of thrmoadaptativtrits has apparntly grtly ompro-
t urrlr i - misdth gnti rord prsnt in modrn organisms, r,vhih maks any
olusions aot th nvironnrntwhr th DNA/protin world origirrtd
70 hc rigilt of I'if

q t l s t i o n a l ( D o o l i t t l | 9 9 9 , 2 0 1B r a t a l . 2 0 0 7 ) . A r r a n a l y s i so f extant forms of lif ut

protin sqtlIlshs f<lr-rnd onl,vorrnzvn1'rvrsgYras' tlrar is spifi Ponnts of ths om;
to hyprthI.mophils; ()thr protins ar pparntly not anstalto thsor- hangs in thm ar vt
ganisms and ar liklv sirnpiy hatd-adaptdvrsions of thos prsntin that no anint inipir
<lllr-tmprtur ognisnrs(Fortrr2002). vn if rh LCA was hypr. ular struturs hav b
thrnropil (Gaulrr t al. 2008; Gouy and haussidon 2008), thr ar al- an ddud, nd th
trnativ planations for thir asal distriution' inluding th possiilitv fshiond and ontm1
that thy ar (1) a rli fr<lmarl1,Arhn lrigh-tmpratur rgimsthat may For instan, th fat
hv srrltdfrom lr svrimpat rgim (Slpt al, 1989;Gogartn-Bokls forming all th ration
t al. 1995); (2) adapttion of atria tO trm nvrronlnntsy lateral rlir, that protin i
trnsf of rvrsgyr.1s(Fortrt al. 2000) and othr thrrnoadaptativ 1998), that is' that th 1
traits fronr hirt-lovir-rg Arha; nd (3) Otltomptionof oIdr rnsophils RNA. This possibiliry
v hyprthrmophilsoiginallr' daptd tl strss-induingonditions othr tht th ribosom's atl
than high tmpraturs(Millr and Lazano 1995). omposd of RNA (Bar
ganization of primitiv
gns, whih ar squr
From th Origin of Protin Synthsist th Last pliadon vnt. Fo ins1
ommon Anstor f Lif gous gns that no
triphosphat)-dpndr
Orr DNA/potin_lrsdlif, nd proaly RNA-sd lif as wll, ar vidn of th istn
dorinant and volvd in stlphistiatitln,th way aron was produd nd synthsis that took pl
s1ustrd rr th rth,ssurf was fovr hangd' spiall aftr pho- mntal vidn of in '
tosynthsisepparl.Th amor.tntof organi aron produd y al1 lif trations laking oth
via the utotrophi fiation of ),' ,,, and simpl organi ompounds (Gvrilova et a|.1976;
suh as fornri and :rtiids rvuld hav far ddth anr<luntsof or- anstral protin syntht
girrrilmpt-lundsr.til-lirg. frln irhr l-rom-
lr still irlg syntlrsiz-d, Bioinformati analy
!]r()Wn prosss or tratrrstrial sours undr tl.rst ondititlns. h domains an b usd t
srvoir of rgani nrtrial prsl1ton th arth thn shiftd from on ini- squns to hartr
tialll. hrtrizdy rlmpounls<lfaioti <lriginto on mad Llp ntirh,' rsulting st is domin
lf iltlgiaIlydriv1omponnts. volvd in gn xprss
It is likiy tht only in th DNA/protin world that iohrnialmahinry whih unwind RNA r
olnpartml.ltlizdy ll-lik lrrmranstrutursl omparirl to thos onsrvd as many tra
usd in lltllrn illogy,ilrwidspad.This is tus tlr first tim tha volution of a ontrol
an dirt r,idnof lif's istl-t might hirvnprsrvdin th form of viding additional suPP
phvsial fossils in th rk rorl.rlir stagswould hav lft hind only prominnt rol during
molulr nll1nts'ilnd ths h11vnt sr.lrvivdth rvlrgsf ghmil biosynthti pathway
us ovr illions of vars of gologi tin-r.Although tras of hydroron rophosphat synthasr
iomarkrs hav n dttdin 2.7 illion-yar-oldsdimntryroks, th taolism. Although t
lldst unmigutlus nlllulr flssils flurrdto dat (Broks t a|. |999), orrsponds only to
diagntiProsss'itllogial ssimilatiotl,and post.dposititlnalmtamOr. potd hr are likly
phism hav long sin radiatdth rlir reod of aioti organi hm. protin domains oml
istrv and th ompontrtsassoidwith primitiv lif. Th high lvls of
All knolvn organisnls shrrth sam ssntilfatursof gnlmrplia- imply not only that r
titln, gn prssion, si antroli rtions, and rnmran-ssoiatd th omplx Bnom
-Pas-rndiatd nrgv prodution. h molular dtails of thsunivrsl dupliation vnts r.
prosssn()t ()nlv prlr,i1 dirt r,idnoi th r-rrlrrophvlti lrigin of ll tion is usd t uild

B r r a t l . 2 0 0 7 ) . A n a n a l y s i s o f
: nzm'[vrsg},rs'that is sifi
]r apprntlynot nstralto ths or-
J-adaptdvrsions of thos prsnt in
r 2 0 0 2 | . v n i f t l r L A w a s a h y p r -
,uyand haussilorr2008), thr ar ill-
l distriution,inluding th possiility
hanhigh-trnpratur rgimsthat may
l (Slpt al. 1989; Gogrrl-r-Bokels
ia to trm nr,ionmntsy latrl
al. 2000) and otr thrmoadptativ
(3) outornptionof oldr nrsophils
)tdto strss-indr-ring onditi<lnsothr
a z a n o! 9 9 5 \ .
nthsisto th Last
oaly RNA-asd lif as wll, anr
tln, th way aron was produd and
s forvrhangd,spilly rftrpho-
organi aron prdud y arly lif
Ho, and simpl organi tlmpounds
] hav f ddth amoutts of or-
irrgsynthsizl'fom ithr horn.
ours undr th st onditions. Th
on th rth thn shiftd frorn on ini.
aioti origin to t>nmad tlp ntirly
ltin world that iohmial mhinry
ran strttturs'omparal to thos
sprd.This ls tlrus th first tim that
righthav nprsrvdin th form of
lir stagswould hv lft .hind only
ot sttrvir.dth rar':rgsof gohmil
: tim. Although tras of hydroaron
billion-r.ar-oldsdimntaryroks, th
s found tl dt (Broks et tl|. 1999),
lation,nd post-dpositionalmtamor.
rlir rord of ioti organi hm.
.ithprimitivlif.
l sslltialfattlrsof gn<lmrpli.
) rati()ns'nd mmrn-assiatd
h molui:rrdtails of ths univrsl
lnof th mlnlph1.lti origin of all
h origi of Lf 71
tantforms of lif ut also imply that th sts of gns that nod th om-
ponntsof ths omplx traits wr frozen a long tim ago (i.e., major
hangsin thm r vry stongly sltdagainst and ar lthal). It is tru
that no anint inipint Stagsor volutionary intrmdiats of ths mol-
ular struturshav n dttd, ut th istn of gradd intrmdiats
an b ddud'and thrfor th suprnatural origin advoatd y oth old-
fashiondand ntmporary rtionists is rndrd unnssary.
Fo instan,th fat tht RNA moluls y thmslvsar apl of pr-
formingll th rtions involvd in pptid_ondformtiorr suggsts's statd
arlir,that protin biosynthsis volvd in an RNA world (Zhang and h
1998|,that' is, that th first iosom lakd protins and ws produd only by
RNA' This possiility is supportd y rystllogphi data that hav shown
thatth ribosom'satalyti sit, wlrr pptid-bond formation taks pla, is
omposdof RNA (Ban t al. 2000; Nissn t al. 2000). lursto th gnti or-
ganizationof primitiv forms of translation ar also providd y paralogous
gns,whih ar squnsthat divrg, not through spition, ut ftr a du-
pliationvnt.For instan' th prsnin all known lls of pairs of homolo-
gous gns that nod two longation fatrs, whih r GTP (guanosin
triphosphat)-dpndnt nzyms that assist in protin iosynthsis, provids
vidnof th xistnof a mo primitiv, lss rgulatd vrsion of protin
synthsisthat took pla with only on longation fator. In fat, th xpri-
mntalvidnof in vitro tanslation systms with modifid ationi onn-
trations lking both longation fators and othr protin omponnts
(Gavrilovaet aI' I976; Spirin 1986) strongly supports th possiility of an oldr
nstralprtin synthsisapparatus for th mrgnof longation fators.
Bioinformati nlysis of sqund llular gnoms from th thr major
domainsan usd to dfin th st of th most onsrvd protin.noding
squnes to haratriz th gn omplmnt of th LCA of xtant lif. Th
rsultingst is dominatd y diffrnt putativ ATPass' and y moluls in.
volvdin gnprssion and RNA nrtaolisr.Th so-alld DAD gyrass'
whih unwind RNA moluls and lav thm rady for dgradation, ar s
onsrvds many transription and translation gns.This suggststh arly
volutionof a ontrol rnhanism for gn xprssion at th RNA lvl, pro-
vidingdditional support to th hypothsis that RNA moluls playd a mor
prominntrl during arly llular volution. onsrvd squnslatd to
iosynthtipathwys inlud thos noding putativ phosphoriosyl py_
rophosphatsynthas and thiordoin, whilr partiipat in nulotid m-
taolism. Although th information ontaind in th availal dataass
orrspondsonly to minor portion of biologial divrsity, th squnsre-
portdhr ar likly to part lf an ssntil and highly onsrvd pool of
protindomains ommon to all orgarrisms (Brra et a|' 2007a).
Th high lvls of gnti rdundany dttdin all sqund]noms
implyot only tht dupliation has plyd a major rll in th artion of
th ompl gnoms fond in tant lls, ut also tlrat for th arly
dupliationvnts rvald y th larg protin familis, aus th duplia'
tion is usd to uild mor mplity, simplr living systms xistd that
72 Th rigi of Lif

lakd th larg sts of nzvms and th slplristiatdgltoryailitisof

thi mintnan nd r
n r m O r i ] 1 ' g n i s m s . h r ' a r i l t i t ' n ' . ' i t r l i t s o l r ln l o l l t t l t an t s p i s
sythsizd y pioti
an asily plaind as th outm of divrgntprossesfrom n ans-
sums that th raw mat
trl lif form that istd for th sparation of th thr major iologial
tiv hmial systms w
domains, that is, th last ommon anstor (LA) or nanstor.No pal-
quird t driv th h.
ontologil rmnnts likly rmain that :rrtstimony of its istn'so th
may hv bn provid
sarh for::r fossil of th nilrrstoris ound to prov fuitlss.
high.ngyompolrrrds
Analvsis of an inasirrglylarg nulrrlrof ompltlv squndllular
Th si tnet of thr
gnomshas rvaldmajrlr disrpanisin th topology of riosomal RNA
maintnan and rpro.
(rRNA) trs. Vry oftn ths diffrns hav n intrprtd as vidn
on priotially synth
of horizontal gn trnsfr (HGT) vnts twn diffrnt spis,asting
hs n no shortag o
dout uporr th fasiility of th ronstrtttior and prlpurrlrstanding f
dil soup took pla.Br
aly iologil histov (Doolittle 1999, 2000). h is lar vidrrtlrt
It is vr unlikly that z
gnoms hav a mosai-lik natur whs omponnts otll from a wid v-
of organi ompourlds
rity of sours (Ohman t al. 2000). Diffrnt advoatshav dsrida
and that th onsquer
wid sptrum of mix-and-math romination prosss'ranging from th
ndognous synthss
latral transfr of a fw gns via onjugarion, trnsdutiorr'r transform-
synthsis in dp_sav
tion to |l ftrsionvnts irrr,lvingorganistrrsfrom diffrnt domains.
orits, and intrplrrlt
Unil,rsl gn-basdphylognisultinatlyrah a singl univrsal ntitr,,
th rlativ signifian
ut th atrial-lik LA (Gogartn t al. 1989) that w favor Was not
simply ognizs th
alon. It would hav n in th ompany f its silings, a population of n-
pounds' th raw matri
titis similar to it that istd throughlut th sam priod. Thy may hav
As disussd hr, th
not survivd, ut som lf tlrir gnsdid if rl-rvam intgrtdvia ltrl
biohmialmonomrs
transfr il-ltotlr l-A gnonl.Th nanstoris thtrson t>ft last volu-
tions is wll stalish
tionary outoms of a srisof anstralvnts' inluding latral gn trans-
ffiint, ut th wid r
fr, gn losss, and paalogous dupliatilns, that took pla for th
ompounds an syn
sparation of Btria, Arhaa, nd uarya (I,azano t al. 1992; Glans-
uilding bloks of lif ;
dorff 2000; astrsana 200 1; Dlay t l. 2004; Brra t l. 2007a).
to thm do not tk pl
ation onditions bul
Th synthsisof h
onlusions and Prsptivs zymati prosssund
thy wr ith ssnt
Although tl.rrhav n onsidral advans in th undrstnding lf hm-
ronmnt. Nonthlss,l
ial prosssthat may hav tzrknpla bf<lrth mrgnof th first living
stituntsof living orgzr
ntitis, lif's ginnings i-tfstill slroudd in mysty' Lik vgtaiol-li a
too striking to fotui
mangrov swamp' th ots of univrsal plrylognti trs r sumrgedin
portd y th ourn
th muddy Watrs of th prioti roth, and how th transition from th non-
onous mtorlts (
living to tlr living took pla is still unknown. Givn th hug gap that ists
oms plusil, br-
in urrn1sriptions of th volutionarytrnsitiontwnrh priorisyn-
pimitiv arth. Fo all
thsis of illrmialmpon1s and th last otnmon anstorlf all tant
a P P r st o u s h a t t h [
living ir.rgs, it is proah' naiv to attlllpt tl ompltly ]srith oigin
stalishdvnts tht
of lif ar-rdth natur of tlr first living systmsfrom molul:rrphylognis.
Mainstram sintif
Our urnt undrstanding f gntis, iohmistry, ll iology, nd th
vlopd within th fr
asi molular prosssof living organismshas hallngdtrr.rnyorigina|as-
walth of xprimnt
sumptions lf th htrltroplrithory. Th r,irvdvoatd hr ssums tht
arrtiv tht links m:
vn if tl-rfirst living rrtitiswr ndtlrvdwith nrinir-nirl svnthtiailitis'
issus.It is tru tht th

:h sophistizltd rgultory ailitis f
l n s o f t r a i r s o t n m o n o
of divergntprosssfrom an anrs-
parationof th tlrrma jor biologil
stor ([-A) or nansto.No pi.rl-
Ltartstimonvof its xistn,so th
s orrndto pov fruitlss.
mr of orrrpltlysqr.rr.rd
is in th topollgyof riosomal RNA
ns hav n intrprtdas vidn
n t s t w nd i t f r n ts p
tstrllti'nand propr un{rstandinglf
}9,2000). Thr is lar vidn tht
I o s o m p ( ) n n ts( ) m f r t ' n l w i d v -
. Diffrrrt dvats hav dsrid
mintion prosss'ranging from rh
jugation,transdution,or transforma-
anisms from diftrntdomins.
tirnatlyralra singl ur-rivrsal
r t al. 1989) that w fr,or ws not
' a n y o f i t s s i l i n g s , p o p u l a t i o n o f n -
hout th sam priod. Th,v nray hav
tid ii thy atrrintgrtdvi latral
nnstoris thus on <lfth last volu-
]al vnts,inluding latrl gn trans-
llitions, tirt took pla tr th
urya (Lazano et a\. 19921Glans-
: t a | ' 2 0 4 ;B r r ae t a \ . 2 0 7 a \ .
rdvI-ts irr h urrdrsrirrrdir.rg
: trth mrgnof th first livirrg
" l d di n m y s t r ) ' L. i k v g t e i < linn
al phvlogntitsar surgd in
t, :rndhow tlr transition frm th nor-r-
known. Givn th hug gap that ists
ry transition tu,nth prioti syn-
h lst Ommon nst()rof all tant
mpt to ompltlydsrith oigin
ystmsfrom molular phylognis.
s, iohmistry.ll iollg,v,and th
nismshas hirllngdmarly liginalas-
Th viw advoatd hr assums that
owd with minimal synthti ilitis,
Th rigi of Lif /)
theirmintnan and rpliation dpnddprimily on organi o-tpounds
t a n t s p i s
synthsizdy prioti prosss. An updatd htrotrophi hypothsis as-
sumsthat th raw matrial for ssmling th first slf-maintaining, rplia-
dv hmia]svst1s\t,as th utom of ilti syrrthsis,rvhil th nrg),
quirdto driv th hmil rtions involvd in growth irnd rprodr.rtion
may hav n prlvidd ,v yanamid, thiostrs' glyin nitril, or othr
high-nrgy omporrnds(d Dtrv 199S:Lazano and Millr 7996).
e[lulzrr
Th asitnet of th lrtrotroplrithory of th origirr of lif is that h
maintnn anl rprodution of th first living ntitis dpndd primarily
on prbiotiallysvnthsizdorgani moluls. As summarizd hr, thr
i s . s t i n 8
hs nno shortgof disussitlnaottt how t formtion of th primor-
dial soup took pl. But lr:rvtoo mal1y ooks spoild th slup?Not rally.
It is vry unlikli,that any singl mhanism n aount for th wid rang
of organiompounds thr rnay hv aumulatd on th primitiv a]rth'
nd that th itnsquntprioti sollp Was formd y orrtiutionsfrlm
ndognoussynthssin a rduing tmosphr, mtal sulphid-mdiatd
synthsisin dp-savnts' and ognous sours suh as omts' llt-
orits,nd intplirntarvdust. This lti viw d<lesnot g th issu of
ntit,
th rlativsigrrifianof th diffrnt sours of organi ompounds-it
simply rognizsth wid varity of potntial sours f organr om-
pounds,th raw nratrialrqrrirdflr th mrgnof lif.
As disussdlrr,th istnof diftrrttioti mhanisms v whih
biohmialmonomrs arr synthsizdundr plausil prioti ondi-
tions is wll stalishd.of ours' not all prioti pathr'vaysar qually
ffiint, ut th lvid rang of pri-rr-rtal
lnditions und.rr,hihoreni
olpundsarr synthsizddm()llstratsth21tprbioti synthssof th
buildingloks of lif ar roust, that is, tht th aioti rations that lad
to thm do not tk pla undr a nrrow rang dfind highly sltiv
rationondirins bur athr uirdr wid varity of primrrtlstings.
h synthsisof hmial Onstiturntsof ontr-rrporary orgnisms y nonn-
zymtiprosssundr laoratory onditilns dos not nssarily implv that
thyrvithssntialfr tlr origin of lif or avirilirlin tlr primitiv nvi-
of hrn-
ronmnt.Nor-rthlss, th rrnarkal oinidrrtwt-t rh molr-rlzrr
on-
stituntsof living organisms arrd thos synthsizd in prioti xprimnts is
too strikingto fortuitlus,nd th roLlstnssof this typ of hmistry is sup-
portd1.th ourn t>fnrt>stof ths iohnril ompounds in th ar-
onaous mtoritsof 4..5illion y:lrsago (lrrnfrundt al. 2001). So it
bomspiausil, ut ntlt provl1' that similar synthsis took pla on th
primitivarth.}-orall th unrtaintissurrunding th mrgnof lif' it
aprsto us th21tth forrrrti<lnof tlr pioti slup is orr ot th m<lstfirmly
stablishd vntsthat took pla on th primitiv arth.
ainstramsintifihypothsson th origin of lif, whih hav n d.
vlopdwithin th framwok lf an volutionav nalysis, lrav Id ttl a
wlthof primnt*llrsults and th dvloprrrntof a ohrnt historial
narrativtht links many diffrnt disiplinsand raiss major philoslphil
issus'It is tru tht thr ar lirrggps in th urrnt dsri;ltilnsof th vo-
74 Th rigi of Lif

lutionary transitiorr twn th prioti synthsis of iohmial om- Burkhadt' F., and S. Smi
rlas Dri, 182
pounds and th last ommon anstorof ll tirntliving ings' ut attmpts
Univsitl' Prss.
to rdu it hav llowd mor pris dsription of th ginning of lif,
Bvrlv' . R.' ). R. Lorr,
Nowadays ntral issu in origin-f-litrsarh is to undrstnd th abioti l:r}'fnl th Pilr
synthsisof an nestrlgntipolymr ndowd with atalyti ativity and its C a l r , i n ,M . l 9 6 9 . h m i ,
furthr volution to an RNA world and ultimatly to th DNA/RNA/protin LiL,ig Sstms o tb
world haratristi of all lif on arth. ! fa major unsolvd prolms, but C s t r s a n a ,J . 2 0 0 1 . o l
Bilpbsi t 150
thy a not ompletly shroudd in mystry, unsolval, or unknowal.
hn, I' A. 2006. Th .
15.58-15.t9.
hy, .. and . Sagan
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W.ahtrshliusr, (i. 1988. Bfor nz1.rs and tmplats; of surfa
mtallrsm.irrlbitllogil Ruitus 52 452-484.
.Wang,
. S. 1998. Stility f gnti infrmatinal moluls urrdrgologil
ondititlns.PhD thsis,Sripps Institution of Oarrlgrlrphy, Univrsit of
lifrni t Srl l)igl.
Wthrill' G. w. I990. HlrmtrIrot tlr eart, ttul Rt,uiu,,rlf rtb Pltr
S i 1 8 : 2 0 . 5 - 2 . 5 6 .
Whit, R. . 1984. Hydolyti stilit.vof iomllulsIrt high tmpratursand its
implitior-r for lif t 2.50 .. Ntur 10 430_42.
r l r i t Wild, S. A.' J. !0. \/aliy,W. H. Pk, nd . M. Gahm' 2001. vidn to-r
dtritlzirons {or th istnlf tltltinntirlrrrstnl ranson th t]r4'4
G i . r g . N t t t 4 9 | 7 5 _ 1 7 8 .
Yr'lirsa, S., D. Florv, B. Basil' nd J. Orl. 1984. Airi sr.nthsisf purit-tsirnl
othr ht()vliompounds v th ation of ltritrldishargs.Jtlurl of
llulr tl lut itl 21': 7 6_80'
Zanle,K., N. Arndt, . okll' A. ttallidy' . Nist, F. Slsis' and N. . Slp.
2007. mrgl.llf a hitl plnt. Spae Si Ruitus129l.35-78.
Zhang, B., nd T. R. h. i998. Ppti1ond flrmtilnv in vitro sltd
ribozyms. Ntt'tr.]90:96_l00.
Zuay, G. l9!)8. Studis On th llr]-taiyzl syntlrsislf aldlprrtoss. ()rigis of
l-if d t'llutil of tb l3iosp|lr 28 1.2_26.
 Narrative

There are many ways to re(

nren)'fvpical examples,Bent
o f l 0 s t a g e sf r o n r t h e o r i g i n
Paleontologyand the History l.The origin of life.The tt
of Life fossil recorcl,some 3.6-
c e l l sw e r e p r o b a b l y l i k e
but they lived in the irbs
Michael Benton 2. Eukarycttesttnd the oriS
nucleus,are reporteclfrr
ar.rdhencepresum:rblya
And out of the ground the Lord God formed every beast of the field, and o f s e x u a lr e p r o d u c t i o n
every fowl of the air; and brought them unto Adam to seewhat he would eukar,votes, possibly no
call them: and whatsoeverAdam calledeveryliving creature,that was the
mrrterialancl reconrbin
name thereof.
3. Multicellularity. The hr
G e n e s i s2 : 1 9
billion-950 nrillionyea
m o l e c u l a re v i d e n c et h a t
s o m e 1 . 2 b i l l i o n y e a r si 1
d i r e r s i f yt h e f u n c t i o r rrs
4 . S k c l e t o n sN. l r n y l n r m a
Peoplehave always been astoundedby the diversity of life, although perhaps
skeletonsabout 545 rni
in different ways. In prescientifictimes farmers saw how their crops and live-
Phanerozclic,during thc
stock were merely part of a much larger richnessof life, ar-rdpeople have al-
are known from many I
wrrys striven to understandthe complexity and arrangementof living things.
d i v e r s i t yo f p l r v l a r c p r e
From Aristotle to Linnaeus,scienristsatremptedto catalog life and to under-
skeletonsu,ereaccluire
stand where it had come from. During the eighteenth century it becameclear
skeletonsclearly ofierec
to all savantsthar the earth had beenpopulirtedformerly by strangeand mar-
g r o u p s t o e n t e rn e w l i f c
velous creaturesthat had since becomeertinct. Bv 1820 some rough picture
5. Pretlation Perhapslink
of the successionof floras and faunas through geological time was beginning
protective outer shells,r
to emerge.Charles Darwirr, during the voyage of HMS Beagle in the early
predato rs, tnacrosco1't ic
1830s, becameincreasinglyconvincedthat life was more diversethan he had
strategiesto feed on the
imagined-every island he visited sported a new crop of plants and animals.
prey becamea st:rndar
He saw the lateral (geographic)and vertical (historic) links between species
6. Bictlogical reefs. Reeis l
and realizedby 1837 that speciesrvereall linked by a great tree.The tree con-
been made from a broa
cept made it clear why speciesthat in his time were geographicallyclose
polychaeteu,orms in th
should also be genealogicallyclose. Further, the tree concpt made it clear
l.ate Cambrian and alg
why the fossil mammals he found in Argentina should be similar to the living
tabulate corals in the O
mammals of the region.
groups of algae,sponge
This essayaddressesfour concepts:what we know about the sequenceof
through time, but reefs
the history of life, how life has diversified through time, how speciation oc-
scleractinian c o r a l s .R e
curs, and how good (or bad) the fossil record is as a sourceof data on the his-
rnrrjor physical geograp
tory of life. These were all issuesthat concernedDarwin (1859), and they
habitats for life.
concern us still today.
7 . Terrestrialization. The t
huge arra,vclf new life z

80

d th History
rnrd vry ast of th fild, and
unto Adam to s wht h would
vy living ratur' tat ws th
G n s i 2s : 1 9
divsityof lif, although prhaps
mtssW how thir rops and liv-
rihnssof lif, and popl hv al-
y and arrangmntf living things.
lmptd to atalog lif and to undr-
ightnthntury it amlar
:ltd formrly strngand mar-
tint.By 1820 sonr rough pitur
ugh gologial tirn was ginning
,yag of F{S Begle in th arly
rt lif was mor divrs than h had
l a nw rop of plants and animals.
i l ( h i s t t l i l)i n k s
linkd y a greattre. Th tr on-
ris tim wr gographially los
hr, th tr onpt mad it lar
ntina should similirr to th livins
tt w know out th squnof
l through tim' how spiation o-
rd is as a sourof data on th his-
lnernd Darwin (1859), and thy
Plotolrg1'
d th istlr of Lif 8,l
Narativ
.hr
r many ways tO r()untth strrytlf th history of lif. As <lnof
manytypialapls,Bnton anl Harpr (|997,2008) psntd squn
of 10 stagsfrol th lrigin of lif to th origin of rldrnlrulnns:
|.T origin o| lii' Th apparirtrof th first vilt-ts of lif irl tll
fossil rlrd,s<lm.6-3..5illi<lnyars ago. Ths first, pr<lkirryori
lls wr prbaly lik modrn yanoatria (..lu-grnirlga''),
ut thy livd in th asnof oygn.
2, ukrotes d th lrigi of s.Th first ukirryotlls, wit a
nulus,ar ptlrtdfrom rlks latdalut 1'.]-1 illion as lld
rrdhrrprsurnlyllpplrd som tilll [rttlrt. h inititi<llr
of sxul prldution,possily at th sam tim as th lrigirrof
ukaryots,plssilynot, opnd up th pilssiility of miing rlf gnti
matrial and rominatin.
3. ltillulritl,'Te first multilld flssils,r1alga, are |'26
illion_950 millio yars rlld, tl]this rng of 1trts lrrspon]s to
molulirrr,idnthat points t() th origitl lf rnultilldlrgirnisnrs
s o m 1 . 2 i l l i o n y a r s a g o . o r g a n i s m s t l n s i s t i n go f m a n y l l s a n
divrsifyth fr"rntions of thos lls, and thy :rn orlarg.
4.Skltls.Mlrny aninral groups apparntly quird hard, minr:rlizd
skltonsalrrt545 millin s ago' at th ginning of th
Phanrozoi'durir-rgtlr sl-lld(,irrrinpllsi<lrr. Anirri:il tilssils
ar known frlm m:rny lo[itisin th pr1ing diaararr,llut th
divrsityof phyla rprsntdy fossils parrddhugly wl.rrl
skltonswr quird. Th rasons for this .rr unknown, ttt
skltlnslrlv offrd pottion nd sr'rpplt rrd prrr-rittd rtarn
groups to ntrrw lif Zons.
5. Prdtiltt.Prhaps linkl with th spralof skltlns,t1.piall
prottiv()utrshlls, ray lrav n th ris lf nw kinds of
prdators,larosoi animals sh s triloits tht mplovd nw
stratgisto fd on thir larg pry. Arms ras twnprdl1torsand
t w ns i s pry lrmlr stilndard fttrlf anim:rl volutitln thrafrr'
6. tsiologilrfs. Rfs lrr,istd sirrirnlbrin tirrrs.Tlry l-rv
nrad from road r'rrrgof aninrals;lminnshift1frlrn
polyhatworms in th arly amrian tl rhaoyathansin th
Lat amrian and lga, rr,ozoans'strolatoporoids!and r.rgos and
taulattlrirlsin th ordoviiirn, Siluin, n1Dvonian. Diffnt
groups of alg;r,sPongs' rt,ozoilns,nd orlrls m and wt.tt
thouglr tirrr,ut rfs rtlr.rindand ar l1dlmirratl'
slratiniarrorals. Rfs r uilt from orgnisms, ut thy [orr
major physial gographi fatursand pr<lvida plthora lf nw
haitatsfrlr lif.
7.rrestrilizittio.Th "llvlf lif onttl land similrly opnd up a
hug rrir\,of nw lif zorrs.Soils ar knorvrrfr<lmslnllat
82 Plotrlllgd th Histor of Lif

Pramrian sussions,and soils imply lif on lalrd. Burowing 2. lndpedt rpli

animls ar known from ()rdoviin soils, whil mny smll vasular or gs' r linkd
plants and artlrropods ar known from th Silurin. Land plants lrromosom pr
am larger z.rndmor divrs in th Dvonin, nd tlr divsit of wlrn on gn is r
land-dwlling,animals pandd to inlud Worms' mlllusks,many Pvnts olllPtitio
mor arthropldgroups, and vrtrats. ()()pratiOnon thI
8. res nd flrsts.Just as rfs in th szrar gographially 3. RN s g|1d ei
sustantial iologial struturs,so too ar forsts.Th first trs il-t laor btwn two (
th Devonin Wr largly isllatd,btlt massiv forstsdvlopd in that stornd trans
th aronifrous, with trs up to 20 m tall. Not only did varilus rations and form
land-plant groups divrsify s trs' ut othr plants and animals musl, trrdn,hai
that ploitd th nw habit:rtsratd th trs ls divrsifid. otlr funtions. Th
9. Flight. A ftlrthr pansion of osp;rwas markd y th origin of :rnd potin rquirr
flight. Instsaros in th arly Dvonian, ut th first tru flyrs squndtrmin
wr Cronifrous in ag. Insetsdominatd th skis from thn on. 4. ukrlts d org
Th first flying (gliding) vrteratsaros in th Prmiiln and h:rva singl irtll
Triassi, whil powrd flight in vrtratsaros at last thr a n l a t i a( l u
tims' in ptrosaursin th Lat Triassi, in irds in th Lat Jurirssi, ontains rod-shap
nd in ats in th Trtiary. struturs alld or
10. lnscilusnss. Th origin of nsioust-tss in humrrsis muh lr ukaryos inlr
datd.Is this a fatr'rroi Homl stpisalon, and hen psnt lld Amoir and
only in th past 200,000 vars or so, or did arlir spisof Hlmo, 5. Saul rproLlutior
or vn pursor gnra suh as ustrlopithcs afld Ardipithus' i n d i v i d u a l sa r i s a s .
possssonsiousnssas mlth as 2 t<l4 mi|lion yars ago? two. In nrost kar
(]orrsilusnss ll<rwdhuns to atthings nd t rrrodifvtlrir ll division may
nvironmnts and is th basis of th prlfound impt humans rvhih nw individ
having on th volution of lif. gmts' produd l
6. D i ffret it d e lls.
Ths 10 stagsar linkd losly to major iologial innoviltions (lif' s,
lls lf onl OI1or
rnultillularity' skltons, pdatit>n,onsitlusnss)and major pIlsitrns
allong anirrrrls, pla
in haitats oupid y lif (rfs,trrstrialization,trs, flight). All 10 ar
. kinds of lls,suh .
doumntd to a gtrr lssrtnt in th fossil rord.
h individul,th
In a mor iologilly orintd prsntation' .|ohn aynard Smith and
inforration (trvo in
rs Szathmar (995) idntifid ight major stps from th >riginof lif to
pr<llm,of ous,
human soietiswith languag:
informtion, thy al
7. Rplitig molules. Th first ojts with th proprtis of furrtion.
multipliation, vriation, nd hrditywr probaly rpliating 7. olonil liuig. ol
Inoluls,sirilr to RNA ut prhaps simplr and apal of t h i r s p i s u t n o t
pliation ut not informationl atrsthy did nOt spify othr s,wasps, nd tr
struturs.A popula viw is tht RNA am for DNA ausit irrdividuals rpodu
n at oth as il gn and an nzym' virvsnapsulatdin th trm suprorgism' aoa1
RN orld' If evrllutiorrwer to prod futh,it rvas nssary u'orkrs ar nalrg
that diffrntkinds of rpliatingmoluIshould ooprat,ah produing individ
poduing ffts that hlpd th rplition of lthrs.For this to olonis is importn
hppn, populations of molulshad t<ll.lnlosdwithin som iomassof th Ama
kind of mmrztn,or omrt|t|l1t' orrsplndingto e simpl ll. mr,rhth sm is pr

; imply lif on land. Burrowing
ln soils,rvhil mani, small vsulr
from th Silurian. Lnd plarrts
th Dvonin, and th divrsity of
l inludwoms' rr-rollusks, mny
rats.
h s r gographially
c too ar forsts.Tlr first tres in
l, ut massivforstsdvlopd in
o 20 m tll' Not lnly did vaious
s' ut othr plants and animals
:atd th trsalso divrsifid.
spaWas mrkd y th origin of
]vonian, ut th first tru flyrs
;dorninatdth skis from thn on.
s aosin th Prnrin and
]rtratsaros at least thr
iassi,in irds in th Lat Jur;rssi,
iousnssin humans is muh
spislon, nd lrnprsnt
;o' r did rlir spisof lmo,
\ustrlop itus ad rdip ith es,
2 ttl 4 million yars ago?
ratthings and to modify thir
r pofound impat umans ir
or iologial innovtions (lif, s,
r s i o u s n s s)n d r n , jro r
rialization,trs' flight). All 10 r
n th fossil ord.
lttion' John Mayrrard Smith and
ajor stpsfrom th origin of lif to
jtswith th proprtis of
J i t w r p r l yr p l i a r i n g
raps simplrand pal of
|atlsthy did not spify othr
lNA m forDNA ausit
m' viws napsuitdin th trm
rod furthr, it was nssary
rolui should l<lprt,
lpliatilnof lthrs.For this to
l d r o n l o s dw i t h i n s m
l/, <rrrsplr.rding to simpl Il.
l,lottllog,ndth Histor1,of LiIa 8l
2. Idpedtreplittlrs.In isting organisms,rpliating moluls,
or gs' ar lirrkd togthr 'nd tO nd to forl hromoso|7's ( singl
hromosompr ll in rlstsimpl rganisnrs).This has th fft tht
whn on gn is rpliatd,ll ar. This oordinatd pliation
prvntsonrptiti()tlbtwngns within a omprtmnt and fors
ooprationln thm, sin if ln fails, thy all fail.
.RN s gn d ezl,m.in modrn organisnrsthr is a division of
ior twntwo lasssof mlul..lei', ids (DNA and RNA)
that stor and transmit information ad protis tht atalyz hmial
rationsanl form rnuh rlf tlr strutur of th ody (for an-r;ll,
musl,tndln,hair). Prhps originirlly RN molr.rls prflrmd
oth funtions. Th tansition frlm an RNA rvlrld to a wlrld of DNA
and prrlrinrquird th vllutionof th gntild,r,vhyas
sqtlnd r m i n sp r t i ns t u t u r .
4. uktlrltsd orglllls.Prlkrr;tslak nulus nd usualry
hv singl irulr hrtrrosom.hy irrlLrdth atri nd
yarroatria(lu-grnalga).ttkrots hav nulus that
ontainsrod.shpd lrromoslmsan]usuall othr iltrallulr
strtttllrsa||dlrgtle//es' inludirrgmitohlndria .rrdhlroplasts.
Th uk:rryotsinlud all othr lluIar organisnrs,fronr th singl-
lldAmt>it,.lnd hlm1,dtlrlsttp to hul.nans'
5. Sul rprodtiot;.In prokaryots nd in som ukyots'lrw
individuals aris as s|tllos y th divisiln of a singl ll into
trvl.I most kar\'ots,in ()iltrastithis prlssof multipliti<lrry
ll divisilnmay intrruptdy a pross <lfsul rprodutio tn
whilr a nw individLtirlariss y th ftrsion of twl s lls, lr
gamts'produd y diflrntindir.iduls.
6.Difirtitd clls. Protists ist ithr as singl lis or as olonis of
lls of onlv ol.tor a vry frv kinds, rvhrsmultilld orgarrisms
p i r n s i o n s
among aninrls,plarlts, and fungi ar omposd of rrrnydiffrnt
kinds of lls, suh s musl lls, nrv lls, and pithlil lls.
alr individul, thrfor' itrris nOt ()nopy of rh gnti
i n f o r m a t i o n( t w o i n a d i p l o i d ) u t m a n y m i l l i o n s o f o p i s .T h
prolm, lf tlurs, is tht lthough all th lls ontain th sm
inf<lrnratior-r, thy irr vr 1iffrntirr shap' omp<lsition,trrd
funtion.
7. okil /iuzg.ost orgaisr-rrs ar solitary, il1tratingwith othrs of
thir spisbut nor dpn{nton rlrr.Oth anirn:rls,notzrlyants,
s,wasps, and trmits,liv in llois ir-rwhih ouly a fw
i n d i v i d l sr ; l r o d u S
. u h a o l o n y h s nl i k n dt o a
s|lrorgiszl'an:rlog<lus t<la multillul:rroganisr.Th stril
workrs ar nl<rgolts to th ody lls of arr individual, and th
ah rprtrduingindividuls to th lls lf th grr-rr lin. Tlr origin of srrh
ollnisis irnportant;it has lrrstimatdtlrt on-third f th animal
ionrssof t Amazon rain ftlrstot.lsistsof ants nd trmits,and
nruh th szrtlris proaly tru of ltlrrhaitts.
lotoklg d th Histor of Life

8. Primt socitis, bum soitis, d th origi of lguge. The yars' thn living sp
disiv stp in th trnsition from ap to human soity was proly th past 600 million
th origin tlf languag.In many wys human languag is lik th gn- 2. Th sond argumn
ti od; information is stord and transmittd,with modifiatron, past 450 million ya
down th gnrations.ommuniation holds soitistosthr and 1 0 m i l l i o n y a r s ,a n d
allows hunrans to sapvolution. trrstrial spis tha
aynard Smith and Szthmiry (1995) argu that all ut two of ths ight 3 . h t h i d r g u m n ri
transitions wr uniqu, ourring just on in a singl linag.Th twl - s p i so f o s s i l m r i
ptions ar th oigir-rsof multillular organisms, whih happnd thr numr of known m;
tims, and of olonial animals with stril asts'whih has happnd many Allowing for nonprs
losss,this ould als
tims. Had any of th othr si trnsitions not happnd, and tht inluds
th origin of lif itslf (numr 1), thn w would not hr. Tht only two spis (Spkoski 19!
of th ight hav dmonstraly happnd mor than on spks against
In ths disussions it has
Simon onway Morris's (2003) thsis that ()nvrgntn natur is so pr-
valnt that humanity and mny othr tnt styls of lif r virtually tion fo mrosopi o
invitl. ros ar ludd u
and vry littl is known of
vr istd ar aliv tod
Divrsifiation to 750 million spis rn t

NURs
oDLs
Lif is astonishingly divers today, with stimtsrnging from 5 milliln to
Biodivrsity, thn, has p
100 million spis, ut prhaps' 1Orsorly, homing in on a 6gur -
twn 10 million and 15 million spis(sth min ssay ..Th Pattrn and tody. But how? Thr ar
Pross of Spiation'' y argart B. Ptk and Shala J. Hankison in this ths an prssdin t
vllum).Of ths,fwr than 2 million hv n rognizd and dsrid a straight lin, an ponI
ruptd inras (Figur 1;
so far.
i m p o s d ( F i g u 1 B ) .
It is ommonly assumd that iif today is mor divrs than it has v
n.This smsin som Ways ovilus,ut it ould aIso onstrud as x- The lir modl rprs
traordinary vanity, somhow akin ttl th viw that all volution was plannd r of nw spis in h
o t h r s i s a n t i n r a s ,i .
to lad to human iggs, nd that somhow this instant in th vast span of
tim is th most importnt of ll. Howvr, it is vidnt that all living organ. volutionay ,rnhing m
isms and ali organisms known as fossils driv fom singl ommon ans- tim spiation ats hav
larly at a rat suffiint to
tor (on th asis of th vidn of shrd ompl haratrs,sr-rhas th
DNA-RNA systm of inhritan,homoo gns,and th lik). That om. in rh t of volution in
mon anstor, th singl spis that gv ris tl all of lif, istd som t o t l n u m r o f s p i si s
.5_.8 illion yars ago. But how many spishav vr istd? t h o r d m a i n s f i x d ;h
Biologists nd palontologistshav trid rrumrof lins of rsonlng to o r p r o a i l i yo f s p l i t t i n g
stimat th totl numr of spis tht hav vr istd. Prhaps living ally n rjtd as impr
iodivrsity is only 24Y" of th total that hs vr istd' as is suggstdy The epoetil model i
thr lins of rasoning (Spkoski 19921a 1994): tion. If spiation and xt
will rgular douling <
1. Th first argumnt is asd on an assumd pattrn f spisdivrsity of volution at th lvl of
inras nd th known avrag duratiln of a spis for it oms nlltial rat of inas ol
xtint. If spisdivrsity inrasdroughly linarly (additiv modl) modl has n applid to
through th Phanrozoi' nd vagspisduration is 5_10 million th divsifiation of lif ir

d th origi of llgug. Te
p to human soitywas proaly
; human lnguagis lik th gn-
a n s m i t t dw
' i t h m < l d i f i t i o n ,
cn holds soitistogthr nd
guthat all ut two of ths ight
l in a singl linag.Th tW()x-
orgnisms,whih happnd thr
] asts'whih has happnd many
ls not happnd,and that inluds
would not hr. That only two
:d mor than on spks against
e t o n V r g n in n t u r i s s o p r -
tant styls o{ lif ar virtually
)Stimats ranging from 5 million to
iorly, homing in on figur -
th main ssay..Th Pattrn and
ak and Shala J. ankison in this
tavn rognizdand dsrid
'y is mor divrs than it has vr
rut it ould also onstrud as x-
r,iwthat all volution ws plannd
tow this instant in th vast span of
:r, it is vidntthat all living orgn-
|rivfom a singl ommon ans-
:d omplx haratrs'suh as th
bo gns,and th lik). That om-
v ris to ll of lif, istd som
spishav vr xistd?
d a n u m o f l i n so f r e s n i n gt o
t hav vr istd. Prhaps living
' has vristd,as is suggstdy
tla |994):
;umdpattrnof spisdivrsity
rtion of a spisfor it oms
roughly linarly (additivmodl)
;spisduration is 5_10 million
Plotolog d th Histor of Lif 8.'
yrs,thn living spisrprsnt2-4" of thos that istd duing
th past 600 millin yars.
2. Th sond agumnt is asd on th divrsifiationof insts ovr th
past 450 million yars. If th avragduration of an inst spisis
10 million yars, and th group has divrsifidlinaly, thn 5% of all
trrstrialspisthat vr xistd ar aliv today.
3. Th third argumnt is asd on prsrvailityof fossils. Som 250,000
spisof fossil marin animals hav n namd, similar to th total
numr of known marin animal spisaliv today (200'000).
Allowing for nonprsrvationof soft-odidorganisms and othr
losss'this ould also rprsntonl 2_4Y" of th total numr of fossil
spis(Spkoski 1992).
In thsdisussionsit has only n possil to xamin pattrns of divrsifi-
ation for mrosopi organisms, that is, typial plants and animals. Mi-
rosr ludd ausit has n hard to stimat thir urrnt divrsity,
and vry littl is known of thir arly fossil rord. If 24% of all spisthat
vrxistdar aliv today (10-1 5 million), thr must hv n som 250
to 750 million spisin th past.
MoDLs
Biodivrsity,thn' has pandd from on spisto many millions of spis
today.But how? hr ar many Ways to go from n spisto many' and
thsan pssdin trms of thr mathmatial modls, rprsntdy
a straightlin, an xponntial urv' and a logisti urv' first as n unlntr-
ruptdinras(Figur 1A), and sond with som mass tintions supr-
imposd(Figur 1B).
e liner modl rprsnts additiv inras' th addition of a fid num-
rof nw spisin h unit of tim. (Th inrasin this ampl and th
othrsis a nt inras,i.., tru ins minus xtintions.)In trnrs of an
volutionryranhing modl, additiv inras would man that though
tim spiationrats hav dlind, or tintion rats hav inrasdrgu-
larly at a rat suffiintto mop up th ssspiations.Th implid dlin
in th rat of volution in th linar modl oms aout simply usth
total numr of spisis inrasingrgularly, yt th rt of inrasaross
th oard rmains fid; hn,for any individual volutionary lin, th rat
or proabilityof splitting (spiatin)must dlin. Suh a modl has gnr-
ally nrjtdas improal.
The pontilmodl is mor onsistntwith a ranhing mod of volu-
tion. If spiatilnand tintion rats rmain roughly onstant' thn thr
will rgular douling of divrsity within fid units of tim. A stady rat
of volutiont th lvl of individul volutionary lins salsup to n xpo-
nntialrat of inras ovrall sin total divrsity is vr inrasing. This
modlhas n applid to th divrsifiationrats of individual lads and to
thdivrsifiationof lif in gnral (Bnton 1995; Hwzulla t a|. 1999|.
86 Plrltllog1'
d th Histor of Lif

LAND AND sEA (

Iogisti Thr ar major diffn
and in th sa, and th his
.= frnt. Today aout 85%
.
land, and th main grouPs
.z p r s n t8 r t d i v r s i t y i n t
o ^ddr*"rr////
/
n volving in th sa s
I
rord is dominatd y m.
exponential
srid forms. This domir
rlll by th fats that virtually
600 500 400 300 200 '100 0 arly alozoi (600-450
A Geologia|time (My) history of lif on land ap
Also, fossils in rtain ma
thn thos in many ontin
d y i s a p p r n t I yf i v t l s i
of th insts' ould an
votd y systmatists to l
this diffrn is vn prt
o
divrsifition on land tha
.z
o In studis of th divrsif
is vidn for a short pl.
yars) and a longr on fr
lion yars). This is follow
ponntial inrs in di
600 500 400 00 200 100 0 rising lmnt of a third lo
Geologial time (My) th last 25 million yars o
a
ahivd 125 million yars
Figur l. Thortialmtldlsflr th divrsifitionof lif plottdfrlr th last arin invrtrat divt
600 million yars:(A) in th asnof major prturtionand (B) with two th sussion of thr m:
"rass
xtintionssuprimplsd. In h asth uppr urv is th logistior lgs of diffrnt phyla .
quiliriummtldl' th middl urv is th :rdditivor linr modl,nd th
wr thn rlad. Th (
]owrurv is th ponntiaIrnodl.
and thn divrsifiation slo
proahd. Th ponnti
in th rly ordoviian, r
largly supplanting th al
Th logisti mil involvs on or mor lssi S-shapdurvs' ah n- th nd of th Prmian' wJ
sisting of an initial priod lf sllw divrsity inrs,a rapid ris, a slowing of fauna dmatially, th M
th rat of inrs as a rsult of divrsity-dpndntdamping fators' and trm ris in divrsity.
thn a pltau orrsponding to a limiting or quiliriurrrvalu. Th llgisti In studis of th divrsif
modl has rrusd to xplain pattrns of divrsifiatilnof marin organ- rium intrprtation has ls<
i s r n s( S p k o s k i1 9 8 4 ) n d o f p l a n t s ( N i k l a s t a l . 1 9 8 3 ) . ssion of major Bplti
Thr is larly n() onsnsuson whih modl st plins th divrsifia- vasular plants in th Dv<
tion lf miOr storsof lif thrugh tim, or on whthr all pattrnsof divr- ronifrous to th Prm
sifiation adhr to th sam modl of inras. Th hoi of modt is and angiosprms from th
imptlrtant sir-r h maks prlfourrdlydiffrnt laims out voluion. ing spiation rats and inr
 Plotoklg d tbe Histor of Life 87

LAND AND sA oPARD

Thr ar major diffrns twn th pattrns of divrsifiation on lnd

and in th sa, and th history of lif in ah ralm may hav bn rathr dif-
frnt.Tody aout 85% of dsrid spisof plants and animals liv on
land' and th main groups (plants, arthropods, vrtrats)hav rahd thir
prsntgrat divrsity in th past 450 million yars. Plants and animals hav
bnvolving in th sa sin at lst 600 million yars ago' nd th fossil
rordis dominatd y marin spis,whih mak up som 95o%of all d-
sridforms. This dominan of marin organisms is partly aountd for
y th fats that virtually th only organisms known from th Vndian and
arly Palozoi (600_450 million yars ago) ar marin, and that th arly
historyof lif on land appars to hav ourrd at rlativly low divrsitis.
Also, fossils in rtain marin nvironmnts ar mor likly to prsrvd
than thlsin many ontinnta1sttings.Th osrvation that lif on land to-
day is apparntlyfiv to si tims as divrs s lif in th sa, largly aus
of th insts,ould an artifat that rfltsth gratramount of tim d-
votd y systmatists to ontinntal than to marin organisms. Howvr, if
this diffrnis vn patly orrt' thn it would imply a muh mor rapid
divrsifiationon land than in th sa.
In studisof th divrsifiationof marin animal familis (Figur 2A), thr
is vidnfor a short platau in th amrian (lsting aout 40 million
yrs)and a longr on from th Ordoviian to th Prmian (out 250 mil-
lion yars).This is followd y a long phas (250 million yrs) of nar-
ponntialinras in divrsity through th sozoi and Cnozoi, th
200 ,100 0 rising lmnt of a third logisti urv, whih shows a hint of a slowdown in
time(My) the last 25 million yars or so, suggstingthat a third platau lvl may b
a ahivd125 million yars in th futur (Spkoski 7984).
i r i o no f I i f p I o t t d o t h l n s t arin invrtratdivrsifiationhas n plaind (Spkoski 1984) as
or prturbationand (B) with two th sussionof thr major phass of volution, in whih road assm-
' t h u p p r u r v i s h l o g , i s t io
lags of diffrnt ph|a (uolutionr fus) dorninatd th oans and
'dditivor linarmodl,nd th wr thn rplad. Th amrian fauna divrsifid xponntially at first'
and thn divrsifiationslowd as th quilirium lvl of 85 familis was ap-
proahd.Th xponntial divrsifiationof th Plozoi fauna thn bgan
in th arly Ordoviian, rahing an quilirium divrsity of 50 familis and
largly supplanting th mrian fauna. Finally, aftr th mss xtintion t
lssiS-shapdurvs' ah on. th nd of th Prmian, whih rdud th global divrsity of th Palozoi
g inras,a rapid ris, a slowing of fauna dramatially, th odn fauna ontinud and latdits long-
y-dpndntdamping fators, and trmris in divrsity.
3 or quilirium valu. Th llgisti In studisof th divrsifitionof vasular plnts (Figur 2B), an quili-
of divsi6tion o f m i n o r g e n - rium intrprtationhas also n givn (Niklas t l. 1983). Thr was a su-
tstal. 1983). ssionof major Bpliie (ground plans or arhtyps)of plant typs:arly
modl stxplains th divrsifia- vasularplants in th Dvonian; lyopods' frns' onifrs, and othrs in th
or on whthrall pattrnsof divr- aronifrous to th Prmiarr; gymnosprms in th Triassi to th Jurassi;
inrs.Th hoi of modl is and angisprmsfom th rtaous onwrd. Thr is vidn fo dlin-
iffrntlaims aout volution. ing spiation rats and inrasing spis durations during ah of th first
88 Plotoklg nd th Histlr of Lif

Ibrith o dorul amphiiar

Mrine invertebrates Vas<ularland plants
1000 Palozoi; arhosaurs (dir
,9 and lissamphiians (frogs
s i r d s ,a n d m a m m a l s f r o m
: 500 assoiations rpla ah r
milial divrsity lvls,ut

z
iations.
Th divrsifiation of ir
D rb P r Jul spially in h Msozo
400 200 200 1,99). This suggsts that
time(My0
GeoIogia| t i|m e( M y r )
Geo|ogia
xpansion that prhaps sl
a t t h t i n s t d i v s i t yi
Plots of th divrsifia
Non-marinetetrapods
lif (Figur 3) by Bnton
ation. Th ontinntal
and land plants' is pon
lozoi platau lvl and
ward th Rnt, whih l
approahing an quiliri
ontinntal familis (Fig
D Crb P Tr Jur Cret
ting xponntial urv (F
400 200 0 400 200
Geologialtime(Myr) D time(lvly0
Geo|ogia| flting th ontriurio
annot ignord.
Figur 2. Pattrnslf divrsifiationof fnrilisof marin irrvrtrts (A),
v s u l a rl a n d p l a n t s( B ) ,n o n - m a r i n
ttrapods ( ) , :rnd i n s t (
s D ) .
Stratigraphiarviations:, arnrian1r, aronifrous;(irt, xPLANAloNs
rtaous;D, Dvonian;Jur, Jurassi;O, Ordoviin;P, Prrnian;S, Silurian;
Trt, rtiry;Tr, Triassi;V, Vndin. (Basdln Spkoski 1984; Nikls t al. In omparing logisti anr
1 9 8 3 ;B n t o n1 9 8 5 ;a n d l , a n d i r a n d S p k o s k i19 9 . ) th ky distintion is tv
modls. Th formr imp\
xpansion modls assum
th radiations as th nw st of lads partially rplad th old. ah nw last that suh a iling h
radiation ld to an inrasin.total gloal divrsity,whil th divrsity of th quilibrium modIs fol
pdingfloras dlind.Angiosprms apparntly ontinud tl divrsify t a an influntial ody of r
high rat. It is hrd to idntify platatrsin land-plant spisdivrsifiation, data on divrsifiation as
and it is hard to find vidn for lgisti modls of divrsifiation.qually, (2) intrations twn<
th total urv of spisdivrsitisthrlugh tim is not oviously ponn- is show that lds ma;
til, and, if anything, th pattrn appars tO suggstlinr inras in divr- rat of divrsifiation slov
sity through tim. phnomna, suh as om
Th divrsifiationof ontinntal ttrapldfmilis (Figur 2) apprs to dution of spis rangs
OrrspondtO an ponntial rodl of irs(Bnton 1985). Divrsiry lv- lows xpliitly from las
ls rmaind low, at som 0 to 40 familis, during th lat Palozoi and of on population supor
muh of th sozoi. Thy thn ros to aot l00 farilisat th nd of th sour. An initial po
rtaous and, after rovry from th nd-rtaous tinti()n vnt' f:1- lowd y a platau whI
milial divrsity inrasdrapidly toward 330 fmilis and shows rro sign of a (usually food); this orr
slowdown. Th pattrn of divrsifiationmay disstdinto susstvra- Th pattrn of divrsif
diations of thr gloal lad ssoiations:sl ttrapods (formrly tmd prtd (Spkoski 1984,
 Plontolog d the istor of Life

Ibrinthodot amphtians) and synapsids (mammal-lik rptils) in th lat

Vasularland pIants
Palozoi;arhosaurs (dinosaurs,ptrosaurs' roodilians) in th Msozoi;
and lissamphiians (frogs and salamandrs), lpidosaurs (lizards and snaks),
birds' and mmmals from th lat rtaous to th prsnt day. Ths lad
assoiationsrpla ah othr and ar assiatd with vr highr gloal fa-
milial divrsity lvls, ut it is diffiult to fit logisti ufvs to any of th asso-
iations.
Th divrsifiation of insts (Figur 2D) was also apparntly xponntial,
r Jur ret ert spiallyin th sozoi potion of th urv (Labandira and Spkoski
1993).This suggststhat insts hav had a long and ontinuous pattrn of
GeoIogiaItime (My0
pansionthat prhaps slowd somwhat during th Trtiary. This may indi-
atthat inst divrsity is approahing an quilirium lvl now.
..all''
Plots of th divrsifiation of fmilis of marin. ontinntal. and
lif (Figur 3) y Bnton (1995) onfirm ths varying modls of divrsifi-
ation.Th ontinntal uv (Figur 3B), dominatd y ttrapods, insts,
and land plants, is xponntial. Th marin urv (Figur 3) rtains a Pa-
lozoiplatau lvl and apprs to show a slowdown in divrsifiation to-
ward th Rnt' whih may indiat that marin divrsity lvls today ar
approahingan quilirium lvl. Th urv that omins all marin and
ontinntalfamilis (Figur 3A) ould intrprtd as a singl poorly fit-
s D Crb P r JUr Cret ert
ting ponential urv (Hwzulla et a|. 1'999),ut th alozoi platau, r-
) Geo|ogial time (y0 flting th ontriution of marin invrtrats (ompar Figur 3),
annot ignord.
s of mrininvrtrts (A),
d s ( ) ,a n d i n s t s( D ) .
r, arlnifrous; rt, XPLANAloNs oF PARNs oF DlVRsIF|A|oN
)rdlviian; P, Prmian;S, Silurin;
d o n S p k o s k 1i 9 t l 4 ;N i k l a s t l . In omparing logisti and ponntial modls for th divrsifiation of lif,
r k o s k i1 9 9 3 . ) th ky distintion is twn quilirium and nonquilibrium (or xpansion)
modls.Th formr imly th istnof gloal quiliria in divrsity,whil
pansion modls assum that thr is no iling to th divrsity of lif, or at
atiallyrpladth lld. ah rrw lastthat suh a iling has yt to rahd.
divrsity,whil th divrsity of th quilibrium models forth pansion of th divrsity of lif wr asd on
parntlyontinudtl divrsify at an influntial ody of ologial thory. Logisti modling of gloal-sl
r land-plantspisdivrsifiation' data on divrsifiationassums (1) intrationsamong spiswithin lads,
modls of divrsifiation.qully, (2)intrationstwnlads,and (3) glol quilibrium levls.Many stud-
gh tim is nlt oviously xponn- is show that lads may radiat initially at ponntial rats' ut that th
tO suggstlinar inrasin divr- ratof divrsifiation slows at a tain point as a result of divrsity-dpndnt
phnomna,suh as omptitiv xlusion, inrasd spis paking' and r-
od familis (}.igur2) apprs to dutionof spisrangs (Spkoski 1,984,1'996).This styl of rasoning fol-
r a s ( B n t o n1 9 8 5 ) .D i v r s i t yl v . lows xpliitly from lassial xprimnts in omptition whr th inras
[is,duing th lat Palozoi nd of on popultion supprsss anothr that dpnds on th sam limiting r-
out 100 familisat th nd of th sour. An initial ponntial inras of th sussful population is fol-
l d - t o us t i n t i n V n t 'f i r - lowed y a platau whn th spis gins to dplt th limiting rsour
30 familisand shows no sign of a (usuallyfood); this orrspondsto th loal arrying apaity.
r a y b d i s s t di n r o s u s s i v - Th pattrn of divrsifiation of marin familis (Figur 2A) has n intr-
: asalttrpods(formrly trmd prtd (Spkoski 1984' 1996) in trms of a thr-phas logisti modl that
90 Plontollgnd th istor of Liie

500 400 00 200 100 rprsnts th bhavior <

nd odrn. Th rpla
aility to pnetrat vr
2000
sitis.Thr is som vid
=
'= imlsou|durow d
't500
mor ingniorrs ways. T
o tms of lg-salonr
1000 pting rahiopods, nrar
popular appal of suh s

z 500 ,^:-../;r:
'
-
.\ .a=
/\- 9-
tition runrl whn th
Bnton 7987). Spkoski
0
of lads t diffus or
spis in on lad ar
A
Howvr, thr ar four
G e o I o g i a It i m e
1.Thr is no indpr
600 500 400 00 200 100 ing apaitis on th
1600+ d i v r s i t i si m p l y t h
-t
1400i Continent|organisms is filld. If a nw s
1200 owvr, osrvat
* faunas om into <
1000
(g
insinuat (ntr nr
800
o 2. ultipl logisti m<
600 twn mmbrs
group will gnrall1
z 400
200 nrajor ioti rpl
l i k l y t o d i s a p p a rl
0
as a rsult of intra

B PIaozoi

G e o | o g i |t i m e

600 500 400 00 200 100

1200-.lr Figur 3. Pattrnsof th
M a rine Organisms in numbrs of familisx
1000 (A)' ontinntalorganism
.9 mimumand minimur
800 and haitat-prfrn tn
(o
familisorddas dfin
= 600
U / d f i n i t l ys p n n i n gt h t s
,},--+:, th min or ontinnta
-

400 ',*\' //
f
doubrful strtigrpitt
z 2oo haitat dsignations.Th
organismsis qual to th
m i m u m m s u r sh, o w
t ausfamilis wit]
our in oth marin nd
ontinntal.Stratigrphr
G e o I o g i a It i m e
Centlzi;P' Prmbt
 Plontolog nd tbe Histor of l-if 9]

rprsnts th haviorof th tlrrvolutionaryfaunas:ambian, Plozoi,

nd odrn. Th rplaing furunasar said to hv n hratrizd y th
ailityto pntratvr wid sts of nihs and hIrtl ahiv highr divr-
sitis.Thr is som vidnin favor of this id; for ampl, ltr maritr an.
imalsould urrow dpr,flrm mor ornplx rfs,and aptur pr in vr
mor ingnious Ways. Th quilirium rnodls ould intrptd sirnply in
trmsof larg-salomptition twn major lads' with ivalvs outom-
ptingrhiopods,rammlsoutomptingdinosurs,and sl on. Dspit th
ppular:rppalof suh suggstions,mosr supposdssof larg-slomp-
titionrumlwhn th vidnis xamind (.g.,Gould and alloway 1980;
Bnton1987). Spkoski (1996) attriutdth pattnsof waing and waning
of lads to diffrrs Omptition twn thm at th spis lvl, wh
spisin on lad ar gnrally omptitivly sr"rpriorto thos in anothr.
Howvr,thr ar four aras of onrn with quilirium modls:
Itime
i. h is no indpndr-rt vidn for quiliria, that is' for fid rr.-
200 100 ing apaitison th arth today. In volutionary trms quilirium
divrsitisimply that all availal rsoursar in us and all ospa
i s f i l l d .I f a n w s p i s r i g i n t s ,i t m u s t d i s p l a a p i s t i n go n .
Howvr, osrvtionsof ass whr prviously isoltd floras and
faunasom into ontat suggstthat spisar just as likly to
insinrrat(ntrnw nihs) nd not aus xtintion of othr taa'
2. ultipl logisti md[s imply prditl outoms of intratiorrs
t,nmmrsof th diffrnt faunas. that is. that nrmrsof on
.!her
goup will gnrallysudwhr thos of rrothrwill fail.
majr ioti rplmntshv rrinvstigirtd,on group is mo
likly to disappar ausof an tintion vnt (Bnton 1987) than
s r s u l to f i n t r a t i o n s .

r lt i m e

200
'r00
Figur . Pattrns of th divrsifiation oi lif thrlugh tim in trms of hangs
in numrs of fmilis tant r strtigraphi stag, plttd for all organisms
(A), ontinntal orgnisms (B), nd marin organisms (). In ah grph a
mximunr nd minimum ar slrown, asd on a ot-r-rination of strtigraPhi
and haitt.prfrninforn-ratin.Th minimum masur inluds or-rl
familisrordd as dfinitly prsnt within ah stratigrphi stag or as
definitlyspnning tat stag, and only fmilis dsignatd as rstritd solly to
th marin or ontiDlltl rlm. Th miInr.rmmasllr inluds trlso all
doutfLrlstratigraphi attriutions of familis and ll quivoal nd shard
haitatdsignations.Th sum of minimum masurs fr ontinntal nd rnain
organismsis qual to th minimllm masll for all taxa togthr. Th sum of
mimur Ilasllrs'h<lw.vr, dtls not qul th mimum m-asufor ll
Jur ret txa usfamilis with qtrivol nvirnmnta[ ssignmnts and thos that
our in oth marin and ntinntl sttings ar ountd as oth marin and
ontinntl.Stratigrerphiarviations as in Figur 2 with th ddition of n,
nozti;P' PrirmriaIr.(Bsd lr.rBrrton l99.5.)
e2 lotrllog1'd th L{istorof Lif

3. Th dir.rsifiationof th Mtrdrn fun snrsmor prol()ngdand

Q U l L | B R l U l " 1o F
slorvrthan prditd y a ltlgisti mtlr']l.Th rising phirsof th
logisti uv has last]for 2.50 million vars,with som vidnof a It is hard to slt btwi
slowdown toward th psnt.If th is no rrntplatau,thn it til rntly, quilirium
would Smthat in th sond half of th Phanrozoi,th st-known gists, just as thy hv
part ()f t[rfossil rord, th logisti ruls lr:rvn forglttn. ologists ar now qust
4 . T h l s s i l o g i s t i u r v s i S p k o s k i( 1 9 8 4 , l 9 9 6 ) m a v a r r i f a t so f ptdin the 1960s and
th lvl of analysis.lr urvs ar plt>ttdat th lvl of fanrilis. versifiation might hav
\ihnths ar translatdto th gnri lvl, th logisti pttrns
Th quilirium mod,
gin to rak down, and at spislvl th pttrn oul1 aommodat only rt;
ponlltiai (Bnton 1997). Th shap of th urvs swiths aus paity is rahd, nt
ah gr-rr.rs ontains man spis,and ah fmily many gnra.In an no sh assumtion n
voltrtilnrytr th spisar th tir-rlrl twigs, whil gnra tnd with no ultimt limit i
dpinto th tr, nd fmilis dpr still. So, althouglr th numr Paloiologists hav <
of familis might dfind t a onstant lvl, th nr-rmrof spis rt. rhaps all lif has
in eh ould pandirrgponntiall},. xpansion modl (Stanl
fid in diffrnt wys. I
Th ltrrrltiv to quiliiurrris p|t1sio. Ar tlr ill]gletPattrns,oI lvl of stailit, and pz
at last stlll-llf tlrm (FigLrs2 and 3), t rsult of unotrstraindpan- whil lif on land lnay
siorr? rtailrly, som lds (suh as insts,angiosprms,ids, and mm- and arthropods rpt :
nrals) sm tl ontinu rdiating linarly or ponntially for many tns or quilirium and xpans
h u n d r d s t l f r n i l l i o r r so f y a r s . S u h v r - x p a n d i n gp a t t r n s i m p l y t h a t p l o i o l o g i s t s , u t o r
t h s g r < l u p sa r h i g h l y s u s s f L ral n d a d p t : r l
. h o v r l l p : 1 t t r n so f stat of gloal iodivrs
divrsifiation (Figur ) ilrporatth n[tllr()usonstitllnt lads, sot
panding' tltrs diminishirlg, nd othrs rrnainingat onstilnt divrsity t
any Partiul:rrtim. From an pensionist viwpoint, thr is no prdition Spiation
of how th individual lads irfft ah othr. Nw glo1 divrsity lvls
my ahir,dy ominati<lrrsof nw adapt:rtions,habit:rth:lngs,and h ffort to undrstnd
xtinti()nvllts.Irr th past,50 nlillion }'arsth divrsifitirlnof lif as is at th uppr nd of th
n dmintd y th sptlrularradiations tlf rtain lds both in th sa nd th important ross<
(dapods, gstropods, tlost fishs) and ln land (insts' arahnids, an- studis is spitio (see
giosprms, irds, mammals). Thr is litl vidn that ths major lads tion'' y rgart B. Pt
lrav run otrt <lfstm and nothing to indit tlrat thy will nlt ontlnu to mation of nw spis b'
pand into tlrv()sps. Ploiologists lrl
ponntil inras oLlllimply that ]ivrsifitionwtrttld last forvr' annot tst whthr any
Prsumal thr is a limit tt>th numrs of familis or othr taa that an thy sk to apply th r
inhait th llrth at any tim: suh a limit rvould ausd not last y th distinguishd y diffrr
mount lf srndirrg room on th ark. If a limit of living spl1Wr aP- a wak sustitut,ut th
prlahd'vr smallr tlrganisnrswluld prsuralv fvord v sltion. ultional thinking, sk t
quallr., s irs happnd llrlrny tinls lrrringvtllr'rtion,trrgelnismswould tistil studis of lrg
tak unptdmasursto surviv, for irnrpl,y oupying th ir' ur- invoIvs morphologil
rlwing into sdimnts,and, ill th as of sonr atria,living dp within tion. In pati' this is
th arth's rllst. With siz rdution, th ultimat linrit to th divrsifiation atists of modn ogn
of lif might thn <lrn th vilaility of th hmial omp()nntsof lif, from morphology, and I
priniplly rlt"t. lss stud. Systmatistsr

la smsm prolongd and
dl. Th rising phas of th
n Yars'with som vidnof t.t
is no urrntplatatr,hn it
th Phanroztli' th st-known
ls hav rrforgltn'
( | 9 8 4 , 1 9 9 6 )n r a y a r t i f a t so f
ttdat th ivl of fnrilis.
i lvl, th logisti pitttrns
vl th pttrnoul]
of th urvsswithsaus
l ah familv many gnra.In arr
rl twigs,whil gnrtnd
jr still. So, although th numr
ant lvel' th nurrrlf spis
r Il1'.
o. r th aggrgatpattrns' or
l s u l to f u o n s t r a i n d p a n -
rs, angiosprms,irds, and nr:rm_
)r xponntiallyfor many tns or
-panding pttrns imply tht
laptl.Th ovrll pttrrrsof
umrousonstituntlads, som
rrnaining at onstant livrsity t
viwpoint, thr is rro prdition
lthr. Nw gloal divrsity lvls
rdaptations,haitat hangs,arrd
's th dil,rsifiatilnof lif has
rrsf rtair.r lds oth in th sa
on land (insts, ar;rhnids,an-
vidntht ths rnlrjlrlads
: ; l t t h a t t h v w i I I n o t n t i n U t o
livsifiatioI.r
wotrld lst forvr.
of famiiisor othr ta that n
would ausd not last y th
a limit of living spar Wr ap-
sumaly favord y slti<ln.
ring volution, <lrganismswould
:ampl,y oupying th air, ur-
som tria,iivirrg dp rvithi
ltimtlimit to th divrsifition
I th hmialomponnts of lif,
Plontolog d th istor of Lif 9
Q U I L l B R I U o R P A N S | o N ?
It is hard to slttwnth two modls for t divrsifiationof lif. Un-
til rntly,quilibrium modls hav dominatd th thoughts of paloiolo-
gists'just as thy hav dominatd th minds of ologists.Howvr, just as
ologistsr now qustioningth ovrsimplistiquiliritlrmodls thy a-
ptdirr th 1960s and 1970s, so paloiologistsr ronsidrirrghow di-
vrsifiationmight hav happnd in th longr trm.
Th qrrilibriummodl ssums that spifi nrajor ologial realms n
aommodatorrly rtain numrs of spis,nd th:rt whn th rrying
apaityis rahd, nt divrsifiation ass.Th pansion modl maks
no suh assumptio and allows for ontinuing, if pisodi, divrsifiation
with no ultimat lirnit in sight.
Palobiologistshav dtd arrd ontinue to dat whih modl is or-
rt.Prhaps all lif has divrsifid aording tl ithr an quilirir.rmor an
pansionmodl (Stnlv 2007). Or prlrapsdiftrntstors of lif divrsi-
fid in diffrnt Ways. volution in th sa my hav rsultd in a gratr
lvlof staility, and pattrns of inras may hav gnrally n logisti,
whil lif on land may hav divrsifid ponrrtially sin th first plants
and arthopods rpt autiously out of th watr. Th impliations of th
quiliriumand pansion modls ar profoundly diffrnt, not mrly for
paloiologists,ut for vryon onrnd aout th presnt and futur
s t a t o f g l o a | i o d i v r s i 1 1 ' .
Spiation
Th ffort to undstand how lif diversrfid fron-lits oigir-rto th prsnt day
is at th uppr rrd of th spetur of studis ilt lcrouolutill.At th lowr
nd th important rossovr twnpaloiology and nrodrn volutionary
studisis spcitirln(sth main ssay ..Th Pattrn and Pross of Spi-
tion'' y lvlargrtB. Ptk nd Shl J. ankison tn this volum), th for-
mation of nw spisy th splitting of liegs(volutionarylins).
Paloiologistslarly annot us th biologil speisoept sin thy
annottstwhthr ny two fossil spimnsar apal of intrding,so
thy sk to apply te mlrphologil spis oept, in whih spls ar
distinguishdy diffrnsin thir phnotyps'This may at first sm to
a wak sustitut,ut thr :rrshad assumptiOns:palobiologistsus pop-
ulationalthinking, sk to hratrizspisboundaris on th sis of sta-
tistial studis of larg sampls of spimns' and assum that spiation
invlvs morphologial diffrntition assoiatd with rprodutiv isol-
tion.In pati,this is not vry diffrntfrom th approah of many systm-
atistsof modrn organisms-spis of mollusks are gnrally dtrmined
from mopholog, and fossil and tant matril may form parts of a sam.
lssstLrdy.Systrntistsrarly rry out itltrrdingtrils.
Pleotolog d th Histor1,of Lif

Until 1970 most volutionists assumd tlrt fossils <luld s,v r,y littl Th puntuatd-q
original about spiation' Tn ldrdg and Gould (197z) hallngd th 1970s and 1980s was
onsnsuswith thir thory of volution y puntuatd qtriliria.ldrdg tagonists kpt shifting
and Gould trmd th standard viwpoint phlti grdulism. It assumd a s do n t h i d a s i
that volving linags Wr hanging at vail rats (Figur 4A) ut wr rl supports of pun.
hanging mor o lss ontinuos| (gsls/.Spiationr'vassn as a y- ida of speies sltio
podut of this pross lf hang: sointi,slinagsha.lom so diffr- and' in som modls,
nt from thir starting points that thy ha] volvd into ir nw spis (th ordr haratrs. Sp<
hroosp:iesonpt). Spiation y splitting (ldogesi.s/ould lstl hp- and vn in oppositio
pn, ut th pross Was no slowr or fastr than normal rats of linag ida of an ..pansior
volutin. (Gould 1990), with h
Th opposing viwpoint' volution puttttdquilibritl (Figr-rr48), Darwinian viw. Suh
proposd that rapid morphologial harrgt ladognsis,rathr than alr- most nthusiasts, and
gnsis,Was th most important pross.Tlr normal stat Of a linag was natural sltion duri
s/asls (no hang),nd fronr tim to tim spitionwould hpprr,ut suh ally onsidrd to t,
spiation vnts wr rapid and rvolutionarY.Th puntutd-quilirium to rsult in signifiant
modl maks two laims: tht rts of hang ;rlonga lingr'plrntuatd' tion of individuals. H
nd that rapid hang is orrlatd witlr s;liationvnts.Th lattr lairn dn pointd to its ir
was th most ontrovrsial,ut it has n rgardd as th ky, dfining l- to xplain it (Land 19
mnt of th puntuatd-qullrbriummodl L.'yits suppotrs. A plthora of as
ldrdg and Gould asd thir nw modl on two osrvations: |99 that purpotd
1. Stsis is ommon in th fossil rord. Fossils an min onstant ln quiliriumviws..ar
not provide nough v
apParanthrtlr'rghmny mtrs of siimnt(i..'thotrsndsor
millior-rsof yars),and thn vrythingsmsto hang' of migrtionsof tx i
2. If most spiation happns ording to Mayr's lloptri (gographi- wht was a gradul i
stppd, puntuationa
splitting) modl, thn it would appar as a rapid vnt in rh fossil
rord. Th dtail of th gadual divrgrrof two is<llatl 1981; Shldon 1987) t
populations
intns fous on fin-s
would not sen.
thy wr still sujt
triloits showd a gr
ags just did not sp
mntd a numr of pu
intrprtd as mor lik
nonBnti hang in sl
h fossil rord dr
and Ansty (1995) sur
tns in th fossil r.
angd from radiolri
stratigraphi ags rang
iority onntrating in
of th arth. of th .5
|\,4ophoIo9y with anagnsis (15
6 3 % ) . I t s m s l a , t
F-igur4. ntrastingxptatiorrs of spis.lvl
r.llutionill th [lsi and it had not n pr
plrylti-grlr'talisn-r
modl (A) and tl-rpuntutd-quiliiu modl 1B). Wht thn of th pr
Dgrof nltlrphologiallrngis inditlon th -is, tit-t.l th 1,-1y1,.
t.lt.t at? As vr. th Dr

] thirt fossils ould say vry littl
n d G o u i d \ \ 9 7 2 ) h a l l n g dt h
)y puntuatdquiliria. ldrdg
ft p1,ltigrtdulisnt.It assumd
arialrats (Figur 4A) ut wr
est/.SpiationWas sn as a y-
ns li1s had om so diffr-
rd volvd into a nw spis (th
.ting (ldogezesls/ould aisl hap-
lstr than nomal rats of linag
plttutdquilibri (Figur 48),
at ladognsis,rilthr th:rn na-
Th normal stat Of a linag was
;pi:rtion.tluldhappn. ut suh
'nary. Th puntuatd-quilibrium
rg along a linag punttlatd'
;pitionvnts.lr latrr laim
r rgadds th k' dfinir.rg l-
t y its supportrs.
ldl on tr,oosrvatios:
F o s s i l s l n n l i n t r t l 5 1 1n','
dir-t-rr-rt (i.., tlrotlslrrrdslr
g smsto hang.
t ayr's lloptri (gograhi-
. a s a r a p i d v n ti n
th fossil
rgnof two isolatdpopulations
l\,4orphology
-lvlvolutionirrth lssi
u t d - q u i l i r i rmrmo d l( B 1 .
t l r lt h - i s .t i m t r nt h y - . r l s .
Pleotolognd th Histor of Lif 95
Th puntuatd-quiliium vrsus pylti-gdualism dat of th
|970s and 1980s was hug fun for all onrnd' nt last austh pro-
tagonistskpt shifting ground and rvising thir laims. Th first phas ws
asdon th idas in lddg and Gould's ssays' From 1975 onward sv-
ralsupportrsof puntuatdquiliria xtnddth modl to spousa nw
ida of spissltion, a highr_ordr pross in whih spis wr sortd
and' in som modls, ould evn undrgo sltion of ..emrgnt'' highr-
ordr haratrs. Spis sltion ould, in thory, our indpndnt of,
and vn in opposition to' natural sltion (Stanly 1975). This ld to th
ida of an ..pnsion'' of volr.rtionarythory to many hirarhial lvls
(Gould \990), with hints of an apparnt rjtin of aspts of th no-
Darwinian viw. Suh xtrm positions w andond fairly rapidly y
most nthusiasts,and paloiologistsontinud thir staunh adhnto
naturalsltionduring th lat 1980s and 1990s. Spissltion is gnr-
ally onsidrdto thotiallpossil ut trmly fl and unlikly
to rsult in signifiant adaptations of th typ that our in th natural sl-
don of individuals. owvr, th notion of stasis stuk: too muh fossil vi-
dnpointd to its imp<lrtn,nd qllantitativ gnti modis wr found
to plain it (Land 1986).
A plthora of s studis ws ssmld and pulishd from 1975 to
|99 that purportd to tst th phyleti-gradualism vsus puntuatd_
quilirium viws. arly fforts wr oftn inadquatly doumntd and did
not provid nough vidnaout th auray of dating and th possiility
of migrationsof taa in and out of th study ara. It soon am lar that
what was a gradual angnti pattn to on prson was n oviously
.Williamson
stppd,puntuationl pattrn tO anot. Som studis (.g.,
1981;Shldo |987) involvd hundrds of thousnds of spimns and an
intnsfous on fin-sal dating and statistial analysis of hug sampls' ut
thy wr still suit to itiisnr. For mpl, Shldn (1987) study of
triloits showd a grat dl of anagnsis and stasis' ut his sampld lin-
a g sj u s t d i d n o t s p i a t . . W i l l i a m s o n( i 9 8 1 ) l i v dt h a t h h a d d o u -
mntda numr of puntuational spiation vnts)ut thy wr gnrlly
interprtdas mor likly to xampls of ecopheotpic hang, that is,
nongntihang in shp rsulting from tmporary nvironmntal strsss.
Th fossil rord dmonstrats th widsprad ourrn of stasis. rwin
and Ansty (1995) summarizd th rsults of 58 studis of spiation pat-
trns in th fossil ord pulishd twn 1972 and 1995. orgnisms
rangd from radiolaria and foraminifra to ammonits and mammals' and
stratigraphi ags rngd from th ambrian to th Nogn, with th ma-
jrity onntratingin th Nogn' th past 25 million ys of th history
of th arth. of th 58 studis, 41 (71%) showed stasis, assoiatd ithr
with anagnsis(15 ase 7o) or with puntuatd pattrns (26 ass;
63%),|t smslar, thn, that stasis is ommon in speis_lvlvolution,
and it had not bn prditd from modrn gnti studis.
What thn of th puntuatd-quiliriumvrsus phylti-gradualism d-
bat? As vr' th potagonists on ithr sid sought to dmonstrt th
Pleotolog nd the Ilistor of Lif

uiquity of thir modl, and yt ommon snsSuggststhat thr ar lar Rhizosolenia

iologial rasons that ah modl prvails undr prtiula irumstans. (p|anktonidiatom)
Bnton and Parson (2001) notd that spiation is a onsqunofrpro-
dutiv isolation, and hn th frquny of spiation in a grlup is likly to
b rlatd to th as with whih rpodutiv arrirs ppr. At on - 5.0

trm thr ar organisms suh as planktoni protists that liv in hug plpu-
4.0
lations that sldom nountr arrirs to disprsal, and that do not possss
omplx bhaviors assoiatedwith rprodution. For ths,gnti isolation 3.0
of populations is a rar vnt, and spiation,whih prhaps oLlrsrarly' is
2.O
probaly gnrally long trm and gradual, lasting prhaps half million
yars. At th othr xtrm ar organisms suh as frshwatr fishs that liv
in spatially struturd and oftn-transintnvironmnts. r spiation might
b so ommon that vry lak and rivr has its own rprodutivly isolatd
population of a partiular typ <lffish, ut thspopultions might nithr 2.8

vry distintivnor last vry long. In twnmight lie th mjority of invr-

trat and vrtrat groups' gnrally hibiting stasis ut from tim to
tim spiating in a puntuational way as a rsult of a major prturation in fr
th nvironmnt. | "GAUss
A fin xampl of spiation in a marin protist is sn tn Rhizosolei' a
t\
planktoni ditom that ours today in hug aundan in th quatoril Pa-
ifi (Sorhannust al. 1998)' Th siliousvalvs of this gnus hav aumu. Figur 5. Grdual spi
latd for millions of yars on th sad,and thy hav n sampld ak to morph<llogialharat
3.4 million yars ago from ors. Hug sampls of th valvs an takn two living spisR. ber
s a n l p l i n gs t t i o n so n t h (
vry fw millimtrs through th sdimntpil' and thsan datd au-
divrgntwnth
rtly. Today thr ar two spisof Rhizlsolei|lving sid y sid in th
populationsslowly diff
oans' R. bergoii and R. prbergoii, nd ths an trkd ak for an datd y fossils
som 2.6 million yars. Thn, from 2'6 to .1 million yars ago, th mor- m g n r o h r o ns | a t
phologis of the two spisonvrg and fus, and thr is apparntly only
a singl lineag for tht tim (Figur.5). This splitting vllt is rported
from ight diffrnt sad ors' and th morphologial divrgn ours
in svral morphologial haratrs,so it was vidntly not a loal vnt ut ount th gntis z
ourrd throughout th quatorial Paifi. Diatoms gnrally rprodu rlatd xtant spis
asually, ut thy oasionally produ sual offspring. Prhaps th m- p h o l o g i | d i f f r n t i a
ination of a gnrally sxual rprodutivmod and th arrirlssPaifi dols sms o hav
Oan xplains wh Rbizosolei speciatd phyltially and ovr a span of was spially apid fr<
som 400,000 to 500,000 yars. paring in that tim,
It sms proal that sually rproduing animals tht liv in varid ag. owvr, th int
haitats mor oftn show puntuational spiation. trrbdotos ls an as- y information from ]
ophoran hilostom ryozoan that is rprsntdtoday in th arin thr ar qustions ov
by thr spis.oastal roks of th Dominian Rpuli and lswhrin vl. th oigins of th
th arian doumnt th past 10 million yrs of sdimntationin shal- ar more onfidntly d
low sas, and thy yild aundant fossils of this ryozoan. Th fossils show Th pluality of v
tat trrbdotos radiatd dramatially from 8 to 4 million yrs 8o' ism plus stasis, puntu
splitting int som 12 spis'most of whih had did out y th Quatrnary vironmntal ontrols l
(Figur 6). Studis y Chtham and Jakson (1995) hav stblishda vari- ( | 9 9 6 | p o p o s dt h ;
ty of protools for distinguishing spis within trrbdolos' taking into n v i o n m n r sa n d h a

snssuggststhat thr a lar
s undr patiular irumstns.
:iation is a onsqunof rpro-
lf spiationin a grup is likly to
'tiv barrirs appar. At on -
ri protiststhat liv in hug popu-
iisprsal,and that do not possss
rtion.For ths,gnti isolation
'n, whih prhapsours rarly, is
,l' lasting prhaps half a million
suh as frshwatr fishs that liv
vironmnts.Hr spiation might
ls its own rprodutivlyisolatd
hesepopulationsmight nithr
n might li th majority of invr-
h i i r i n gs t a s i sb u t f r o m t i m t r l
' rsultof a major prturation in
protist is sn in Rbizosolei, a
;aundanin th quatorial Pa-
valvsof this gnushav aumu.
d thy hv nsampld ak to
mpls of th valvs an takn
pil, and thsan datd au-
osoleniliving sid y sid in th
nd thsan trakd ak for
l 3.1 million yars ago' th mor-
fus,and thr is apparntly only
). This splitting vnt is rported
morphologial divrgnours
,s vidntlynot a loal vnt ut
i. Diatoms gnrlly rprodu
lual offspring. Prhaps th om-
mod and th arrirlssPaifi
d phyltially and lvr a span of
ring animls that liv in varid
i t i o n . l r r b d o t o si s a n a s -
lrsntdtoday in th arin
inian Rpuli and lswhrin
)n yars of sdimntationin shal-
f this ryozoan.h fossils show
from 8 to 4 million yars a]o'
h had did out y th Quatrnary
ln (1995)hav stalishda vari-
vtthin tr rbdotls, taking into
Plotolog d the Histor of Life 97
Rhizosolenia
(p|anktoni
diatom)
g R. bergonii
l R. prebergonii
Hyalinearea
#
"GAUss" l "MAUYA|\A,
Magnetohron
Figur.5.Grdual spiationin th diatom Rhizosolni.On of svral
morphologialhratrs,th ightof th hyalinra' that diffrntiatth
two living spisR. bergoii and R. prbrgonii.This plot' from on of ight
saplingstationson th floor of th Paifi Oan, shows a long-trm
divrgn twnth two spisfrom 3.1 to 2.6 million yars ago, as the
populationsslowly diffrntiatd. Th sdimntaryrord is ontinuous,and it
an datdy fossilsand y magntirvrsalmasumnts, indiatedin th
masntohronsalat th ottom.
ount th gntis and th amount of morphologial diffrntiation of
rlatdxtant spisand thn xtnding omparal statistialtstsof mor-
phologial diffrntiation to th fossil forms. Linag splitting jn trrb-
dolls sms to hav n rapid and puntuational in haratr. Spition
was spiallyrapid from 8 to 7 million yars ago' with nin nw spisap-
paring in that tim, although sampling may a prolm in roks of this
ag.Howvr, th intrval from 8 to 4 million yars ago' rprsntd largly
by information from Dominia, has n intnsly sampld. So, although
thrar qustions ovr th origins of th nin asal spis within this intr-
val, th origins of th rmainder (te,nw spis 10, and nw spis 8)
ar mor onfidntly doumntd s ing puntuational.
Th plurality of volutionary mods (gradualism without stasis, gradual-
ism plus stasis,puntuation plus stsis)might ral, and thr might n-
vironmntal ontrols that work in a somwhat unxptd way. Shldon
(I996) proposd that gradualism might hratriztaa that liv in stal
nvironmnts and hang in lin with slow nvironmntal hangs, whras
98 Plontologd thistor of Lif

tenu 9roup A qulitativ rgun

nothing Inuh nw h
-----.--.--
was Stillin 1859 rgu
urian rptils and D.
rturn to arth. DarW
olls' nd h knw
+U o m l n l n
t^
vonian, amphiians ir
SmplingInterv|
in th riassi, zrnd m.
,
V 1 8 5 9 , a n d d s p i t t h
hav simply adjustd
has n pushd l
phibians from th ar
rptils forn tlr r
of mmmals frorrr tht
has rmaind unhan
lssly unrprsntativ
to tim.
Th notion of a got
by awll arrd Bntr
of th mrin animal l
knlwledg from 1890
oth ass th sum tot
Figur 6. Puntuational spitionin th rlzoanetrrbdo,os' Th thre
l i n g i n 1 0 0 y a r s , u
living spisr doumntdy fossils,ut a furthrnine spisar xtint.
to tim, and sl th
Intensolltingthroughoutth arianhas rvaldth pattrnof
spiation.Samplingin th tim from 25 to 8 rnillionyarsago is sporadi,ut rmaind unafftd.
th tim from 8 to 4 million yars ago, th Dominian Sampling Intrval (DSI), whth nw fossil dis
is xtremly wll known' asd on a fin and ontinuous fossil rord on th Th found th form
island of Dominia. Spisdistintionsr asdon a omintionof th ompltness of tl
morphologialand gntistudis. mor oftn thn ret
h hav n tw
sampling ffts zrrrdl
stasis might b a fatur of taxa that oupy unstal nvironmnts' whih tht th fossil rord
an vary rapidly and dramatially, ut thos taa do not volv in lin with light th fat tht th
v r y n v i o n m n t a fIlu t u a t i o n ' wll known it is, th {
odid ogarrisrnsad
h z r p sl i v i n g i n u n u s u l
Quality Th ritiisms of sa
(2001,2007) argud
Th quality of th fossil rod has osssdand ontinus to ossspaloiol. ok rord, whih itl
ogists. vr sin Drwin (1859, 279) considrdth ..imprftionof th go. posd tintion vl
logial rord,'' palobiologists and othrs hav osillatd in thi onfidn. thr is a major rgrs
Som hav laimd, prhaps rathr wildly, tht vrything th fossil rord says parntly disappar frt
is orrt, whil othrs, prhaps qually wildly, hav ritd th fossil rord ut it is mly sdir
as bing nxt to uslss.Th truth prsumaly lis somwhr twn' and vr, that th divrsifil
most popl taitly vr to th positiv nd of th sptrum. A numbr of ma. is poaly ral ar
jor hallngs hav arisn rntiy' howvr, and I shall touh on ths brifly. v r s i t yi s n o t d r i v n

unguiculaturn group
n.sp.
Interval
i *l"Plins
lipolnum
zoantrrbdotos.Th thr
a furthrnin spis tint.
has rvaldth pattnof
8 million yarsago is sporadi,ut
)ominianSamplingIntrval(DSI),
l ontinuousfossil rord on th
asdon a omintionof
upy unstalnvirnmnts,whih
os taa do not volv in lin with
l nd ontinusto ossspaleoiol-
idrdth ..imperftionof h go-
hav osillatd in thir onfidn.
hat vrythingth fossil rord says
ldly, hav ritdth fossii rord
rably lis somwhr twn, and
of th sptum.A numr f ma-
' and I shall touh on ths bifly.
Plotologd tb Histlr of Life 99
A qualitativagumntfirst, in favor of th fossil rord, is tht sin 1859
nothingtnulrnw has <lrto liglrt. In rh histoial ontt, harls Lyll
Wasstill in 18.59arguing for nonprogrssion,that h might xpt to find Sil-
urian rptilsand Dvonian mammals, and that il-rthyosaursmight on day
rturntO arth. Darwin agud for progrssion of lif forms thror-rghtim, of
oufs'nd h knw aorrttriloits in th Silurian, armord fishsin th D-
vonian,amphiiarlsin th art>lrifrous. rptils in th Prmin, dinosaus
in th Triassi,nd martnaisand irds in th Jurzrssi.In th 150 yars sitr
1859,and dspitth input of millions of hours of sarhing,palontologists
havsimply adjustdth rrrid-Vitorianpitur: th origin of agnthan fishs
has rrpushd k from th Silurian to th mrian, th origin of am-
phiiansfrom th arly aronifrous to th latst Dvonian, th origin of
rptilsfronr tlr arl rmian to th rrrid-aronifrous, arrd th origin
of mammals from th mid.Jurassi to tlr lat Triassi. Th origin of irds
.!7r
hs rmind unhangd in th latst Jussi. th fossil rord hop.
lsslyr.rrrrprstltativ'r-rwfinds should provid mjor surpriss from tim
to tim.
Th notiln of a gotrd fossil rod was onfirmd in quantitativ analyss
y awll and Bnton (|990) lf th vrtratrord arrd Spkoski (199)
f th milrin nimal rord. Th first authors lookd at th aumulation of
knowldgfrtlm ] .990 to 1987, th sond t hngs from 1982 to |992. In
othassth sum totals of divrsity through tim inrasd,ssntillydou-
ling in 100 yars, ut th inrsswr randomly distriutd with rspt
to til1-l,and stl th ovrll pttrns of divrsifiations and tintions
rmaind unfftd. In a furthr study Bnton and Storrs (1994) tstd
rvhthrrrw fossil disovristnd to fill prditd gaps or rat nW gaps'
Th,vfound th formr: in 25 yas of study, nw fossil finds hal improvd
th ompltnssof th ttrapod fossil rord y som 5% plugging gps
mor ofttthn rating llw saps.
hr hav bn two rnajor hallngsto this somwhat omplnt viw:
sampiingfftsand mlular phylognis.In both ss th ritis apt
that tlr fossil r<lrdas d<lumtrtd is wll undrstood. Howvr' thy high-
light th ft that th fossil rord itslf is a poor sampl of lif. So, howr,r
rvllknown it is, th fossil rord an nvr dourrrntth volution of sott-
bodiloganisr-rrs adqtrtly,zrrrdlthr groups-prhaps mirosopi' pr-
haps living in unusual nvironmnts-my similaly not fossilizal.
Th itiismsof sampling, ltlrough linkd, hav had diffrnt foi. Smith
(2001' 2007) argud tlrat th fossil rord is losly tid tl th sdimntary
rok rord, whih itslf is linkd to sa-lvlhangs, and that many sup-
posd tintil>nvnts r nothing mor thn lrengs rf nvironmnt. If
thris a mjor rgrssiln(rtrtof th sa),sh:rllowmrin organisms p-
parntlydisappar from stions. This ould rordd s an xtintion'
but ir is mrIy sdinlntryrtifat. Snrith (200t ,2007I poirrtdout' how-
vf'that th divrsifiationof lif in th sa durirrgth last 250 million yars
is proablv ral bussa lvls wr falling duing this tim: th ris in di-
vrsit1, is ntrt drivn y a is in sa lvl.
1 0 0 Pleontolog d the Histor of I,if

Ptsand Foot (2001,2002) wnt on stp furthr, arguing that virtually p p r o a h s :f o s s i I d t t

all th fossil rod is losly dpndnton th sdimntaryrord. Supposd first fossil of a group).
rxtintions' divrsifiations,and otlrr iollgialpattrns ar l1 apparntly dts tend to too o|
drivn y th volum of psrvdrok. Ths authors found that th divr- spaking fror th mol
sity of lif apparntly maths th numr f namd rok formations in molular ag doulin
North Amria. So, thy argud, whn lif pprs to divrsifying, it is h i p o s i t i o n . o n h l l
simply bausthr is mor sdirntaryrok and lrnmore fossils.IWhn d v s t t i n g p o s u r l
a mass xtintion is idntifid' it is nothing mor than a loss of apprtlprit n shown to rst on
roks in whih to prsvfossils. D o h s d t sj u
Ths viws hv n sn as prhaps rthr trm (Bnton 2003; P- v r r i d g t h g a p t
trs 2005). Th orrlation of numbrs of namd formations and divsityof to good) and th pr
lif rpotd y Ptrs and Foot (2001,2002) ould rvrsd:it is just s don to sol tn
likly that whn lif is divrs nd fossils a aundirnt' gologistsrogniz Novak 1l992) r|i
and nam mor formations. In gnral' of ours, as w always th oLlr stu. sours of data on th(
dnts, orrlatio does not imply ausatiotl.Furthr, assurningthat rok vol- logny. By fossils thy
um, or som proy for rok volum, is pur rror in our undrstandingof in th roks. laiistis
th fossil rord and should thn b applied as a Orrtion ftor is havy n W p p r o h st o d r l
handd-maior iologial vnts suh as divrsifiatilnsand mass tin- phologial o molulz
tions ar rmovd at a strok. As Ptrs (2005) pointd out' th ls no ra. Norll nd Novak
son to rjt th proposal that mrin rok volum and marin divrsity ompard for rngu
might vary in onrt with a third fator, suh as sa-levllrng.Aundant th fossils or th trs
marin roks nd aundant marin fossils might atually rflt high sa lv- hin, o omnisin
ls and aundant marin lif: in suh a as' to divid th pak in mrin f o s s i l s q u n se n d t l .
divrsity y th pak in marin rok volu would fftivly rmov that grunt' thn proaly
(tru) iologial signal. Muh mor work is rquird to invstigatfurthr h i s r o yo f l i f . L a k o f
why th marin rok and fossil rords ar <rrrltd, ut th trrstrialrok i s w r o n g o r t h t r sa l
and fossil rord appars not to (Fara 2002). and thy show good <
Th sond major urrnt hallng to th informativnssof th fossil vk l 992; Bnton ar
rord oms from rnolular studisnd datsaour th tir-rrirrg of origins at tht all grups .
of major groups. Som molular stimatspl th origins of tazoa (an- rord' that thr is no
imal phyla), grn plents, ngiosprms, nd modrn ordrs of irds and fossil rords,nd thl
mammals at points up to twi as old s th oldst rprsntativfossils (.g.' r r sp l O r t da g i t l s ts t
Wray t aI. \996; opr and Pnny 1997;Kumar and Hdgs 1998; Wray
2001; Hdgs and Kumar 2004). Th rang of molular stimatsfor th
origin of mtazoans is 0.6 to 1.2 illion yars ago' witl-rrroststimatslosr Conlusion
to 1 illion yas ago than 600 million yars ago. Th range of molular s-
timats for th origin and bsal spliting of plantal mammals and modrn Palontolgy is an a
birds is 130 to 70 million yars ago' again with mor stimatsnrrr120 tnth nturis. o m
million tan7 million. Th first fossils dat, rsptivly'from around 600 happnd sin yon
and 70 million yrs ago. potdwkly in Si
Th mismath of first fossil dats and first molular dats ould indiat nd th fat that (ds
major rrors in on or th othr sour of data or oth. any ommntators ing is strong vidn
(.g.,asta| 7999; Wray 2001; Hdgs and Kumr 2004) hav argud that th history of lif.
th fossil dats r almost rtainly wrong' and tht th molula dats ar But th history of li
losr to th truth. Othrs (.g.,Bnton 19991 Bromham and ndy 2000; n o t h .g r o u p A g i v l r
Bnton and Ayal:r 2003) hav suggstd potntial prolrns with both n a r r a t i v i s i m p o r t t l t

p furth,arguing that virtully
t sdimntaryrord. Supposd
gial pattrnsar aIl apparntly
s authors found tlrat th divr-
. of namd rok formations
in
ppasto divrsifying, it is
".kand hn mor fossils. Whn
morthn loss of ppropriat
Lthrtrm (Bnton 2003; P-
.mdformationsand divrsity of
2 ) o u l d v r s di:t i s j u s t a s
aundant,gologistsrogniz
urse' s w always talr our stu-
Furthr'assumingthat rok vol-
ror in our undstandingof
J as a orrtion fator is hvy
i v s i f i a r i o nasn d m a s s t i n -
0 5 ) p i n t d u t ' t h r i s n o r - Norll arrd Novak (1992) proosd that ll thr pprolrs ould
;k volum and marin divrsity
;h as sa-lvlhng,Aundant
rightatually rflthigh sa lv-
;' to divid h pk in marin
r would fftivly rrov that
s rquird to invstigat futhr
orrlatd,ut th trrstrialrok
02).
:h informativnss f th fossil
atsaout th timing of origins
plath oigins of tzoa (an-
rd nrodrlr odrs of irds ar-rd
oldstpsntativ fossils (.g.,
Kuma and Hdgs 1998; Wray
of molular stimatsfor th
s ago' with most stimatslosr
l ago. Th rangof molulr s-
plantalmammals and modrn
with mor stimatsnarr 120
, rsptivly,trom around 600
t molulr dats ould indiate
ta or oth. any onrmntators
l Kumr 2004) hv argud that
nd tht th mlr-llardts ar
)9; Bromham and ndy 2000;
potntil prolms with oth
Plaotrllogd tb istt' of Lif 101
approahs: fossil .latsar always to() young, of ours (on nvr finds th
firstfossil of a gror.rp),ut not outragously too young' whil th molulr
datstnd tl too old, somtilsy a long way. Graur and Martirr (2004),
spakingfronr th rl-rolular sid, prsnt a roust attak on th vidnfor
moluiirrag doulirrg just notd, whil dgs and Kurar (2004| dfnd
thirposition.orl balrr,howvr, th nrolular lindings that snldlik
a dr,astting ep()surof th friltis of th fossil rord in 1996 hav norv
bnshorvnto rst on fr wk grould thn was first assrtd.
Do ths dats just rdu to irssrtilnand ()untrasstion?an w
vrridgth gap twnknowldg of th fossil rord (whih is aptd
t o g o o d ) n d t h p r o l m o f u n p r s r v e dl i f o f t h p a s t ? P r h a p st h i s a n
b don to som tnt. ln a rillint ampl of latral thinkrng, Norll and
Novak (|992\ ralizd that volutiolristsuniqulv had thr indpndnt
soursof dta on th history f lif: fossils, ladistis, ad rnolular phy-
logny.By fossils thy mant th stratigrphi lrdr of ourrt-t of fossils
irrth roks. laiistisand nroiular phylogn1'rOnstrutiotrlr rlativly
nWapproahsto drwing voltiol-rary trs' rvlrtlrron th asis of lrr<lr-
phiogilo mlulr hararrs,nd thy ae irrdpndntof statigraph1..
omprdfor ongun(agrmnt).Up to tht oint no on knw rvhthr
th fossilsor th trsrvr itr any lIylos to tlr trutlr: without a tim ma-
hin,or mnisil,how ould on tll? Thy argud, though, that if th
fossilsqunsand th squnsof rnhing points in th trs wr ()n-
grunt'thn proaly th fossil ord did rprstth tru pattrn of th
historyof lif. Lak of ongruIlould indit that ithr th fossil rord
is wrong or th trsr Wrong' or oth. Studisan dorrgroup y group,
and thy show good ongrrrnin s mrly s 7 5, of ass (Norll and No-
vak 1'9921Bntonand Stors l994; I}ntorrt al. 2000). Ths studis indi.
at that all gr<lu1.ls xalrrind ir<lutqtraliy wll prsrvd in tl.rfossil
rord,tlrat this no sustntiaIdiffrnl-ltwrr marin nd ltrtinntl
fssil rords, and tht tlr is no tim ias (for rod_slphyltlg'Ilti
trspllttdiinststag-lvldivisions t;f tim),
onlusion
Palontologyis an nint sujt, dating ak tl th sitnh or svn-
tnthntttris.To ]any' it had its hyday in Vitorian tims, and littl has
happndsin yond th disovris of nw dilrosurs and lronrinids r.
portd wkl l'nSi a'd Ntltr' In a sns,as I hav ilrgttd, tht is trr.t,
nd th fat that (dspittlr lain-rs)ths t-twdistlvrisa rtrrly slrok.
ing is stngvidrrthat tlr fossil rord tlls us solnthin]truthfr.llirlrtlut
tlr histtyof Iif.
But th histoy of lif is mor than just nrrativ of first on fossil, thn
anothr'group A givillg Wav to grltrpB' and so On through to mankind. Th
narrativis important for what it tlls us aout th stonishin!]invntivnss
1 0 2 Plotolog d th istor of Lif
.$hih
of lif, its aility to volv and do unprditlthings. Dvonian o- Bntn'N4.J.' and F. J
srvrwould hav prditd that th multilimd inststhat rpt round th B n t o n ' . J . , a n d D . -
l.ongman.
watrsid plants would on day tak to tlr air? \Whih Jurassi osrvr
208.Itrodu
would hav gussdthat th insts sh saw flitting and daning in tl-rsun- Blakwll.
light around th muddy lgs of th dinosars would volv soiality ilrrd thrr Bntorr'. l., nd P. l
driv thir iodivrsityand gloal iomass t<> untold lvls?urrt-ttiodivr- L|rlllgltd uol
sity is highr thn it vr has n.!hr light lif volv nt? Thr r in- Bntln, . J., anl G.
Palontlgial kr
timtions of miros that liv in i and i boiling Watrs, and sor that liv
Bnt<ln'\4. J.' . A. \
phaps thr kilomtrs down within th roks nath th sa floor. Ths
thrugh tim. Nat
ampls strain rdulity, but thy hav happned and ar happning. Brot-nhrn,l-. D., and l
Thr is mor to th study of th history of lif, though, than mr dou- dts to th amI
mntation of fossils. Fossils rprsnt tint groups. Dinosus annot S e r i sB 2 6 7 : 1 0 4
prditd from moluls, bt thy istd arrd show us organisms tlrat did htlrm, A. H., nd
sltion vrsus
what no living animal dos. Som dinosLlrswr 10 tims th nrass of th
rwin and R. L..
lrgst living marrlals. Som ptrosaurs ad wingspans thr or four ims Rlrd, 184_20 t-
thos f th largst irds today. Som tinr arthropods wr largr than onway orris, S.20
any aliv today. Ths organisms ar all truly wondrful, and thy pos in- amridg: amt
trstingprolms flr iomhaniistsand physiologists. oop, A., and D. P
rtiary undari
Divrsifiations and mss tintions n prditd (rtroditd?)from
D z r r w i n , . 1 8 . 5 9 .o
studis of modrn phylognis,ut th dtails lrav always rmaind mystri-
irstirl'S. 1999. ]Vtol
ous. Palontologists hav tlr grat plsur of irrg al to dissr suh B i r l s s 1 ' s2 1 : 1 0 :
vnts in fin dtil, and as fossil-olltingthniqus'th prisiorrof dat- l d r e d g ,N . , r r n Sl . . |
ing, and nvironnrntalanalytial tols irprv, th lvl of undrsranding grdulism. In T.
will inras. Frniso: Frm
rwin, D. H.' and R. l
Spiation an studid in th laoratory and in long.trm studis in th
and R. L. Arrsty
fild, ut thr is always a tim limit. Now' with improvd ollting and dat- 11_28. Nw York
ing thniqus,palonrologial studis of spisvolution and spissplit- F:rra, Fi. 2002. Sa lv
ting an tid samlsslyto th prsntdy, :rrrdwe ar ginningto s for .|rrl of th Gt
th first tim marovolutiorrir.ration. G o u l d , S . J . 1 9 9 0 .I s a
6:119-10.
Nw finds aptur th hadlins, ut nw insights into ionhanis,
Gould, S.J.' and . B.
larg-slvnts,and marovolution ar vn mor imprssiv.Tlr study
th night. Plobi
of th history of lit has nvr n mor iting than it is now. Graur, D., nd W. ar
of volution nd t
|BLIoGRAPHY Hdgs, S. B., nd S'
Gnetis 20:242-
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811-814. p21ttrs from m
1987.Progrssnd omptitionin llrirrovolutit. Bhklgil R,t,itlls
62-: rstitllts t|-tl
3 0 5 - 3 38 . Kumr' S., and S. B. I-
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limdinststhat rpt round th
th air? \hih Jurassi osrvr
rw flitting and daning in th sun-
rrs would volv soiality and thn
to untold lvls?urrnt iodivr-
night lif volv nxt? Thr r in-
Loiling Watrs' and som tht liv
roks bnath th sa floor. Ths
p p n da n d a r h a p p n i n g .
y of lif, though, than mr dou-
int groups. Dinosaurs annot
l and show us rganisms that did
LrrsWr 10 tims th mass of th
rad wingspansthr or four tims
tint arthropds wr largr than
r u | y w o n d r f u l .a n d r h y p o s i n -
lphysiologists.
n prditd(rtroditd?)from
ails hav always rmaind mystri-
ur of ing al to disst suh
g thniqus'th prision of dat-
lprov' th lvl of undrstanding
lry and in long-trmstudis in th
with improvdolltingnd dat-
;pisvolution and spissplit-
ry, and W ar ginningto s for
nw insights into iomhanis,
) vn mor imprssiv.Th study
liting than it is now.
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:rovolution'B iologil Ruitas 62:
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divrsity. l,hillsophil rstit-lslf th Rol Soiet, Sris B 345:13-2.
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aynard Smith,J., nd . Szthmir. 1995.Th ior Trsitionsi uoltio.

Oxford:.W.H. FrmanSpktrurn.
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Norll' . A., and . J. Novk.|992. Th tssiIrordnd volurin:
omparing ladistiand palontologividnfor vrtbrathistory.Siec
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2002. Dtrminantsof tintionin th fossilrord.Ntre 416:420424.
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Biodiursitriss,77-100. Nw York: olumbia UnivrsityPrss.
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330:561-563.
1996.Plus ga hng-A mdlfor stasisand volutionin diffrnt ing ird ggs hv(
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Ploglgrpll, s n a k s ) ,l l o u t h s w ]
Smith,A. B.200 l. Larg-sal htrognityof th fossilrord:Impliationsfor ra hind th nt
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2007. in divrsitythroughth Phnrzoi: Prlmsnd prospts.
how rlrsmaths
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priat f()r thir
moirs4. Plobiology 33 (Suppl.):1-55.
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loations (.g.,o
Wray, G. A.' G. s. Lvinton,and L. H' Shapiro.1996.olularvidnfor dp
n 6 n d z I s l n d : i s
Pramriandivrgnsamongltazoanp|a.Sienc274: 568_573.
d it smd rs
othrs through tl
that natur Ould
h tlror of a
th lnvitiorrth
intrvnd throu
th filturs of Pl.
and pts or m
,he
jor Trsitils i uolutil.

983. Pattrnsin vasulr land plant

ssil rord ad volutio:

Adaptation
dn for vrtrathistory. Si

maovolutionary history of marin

lm oi Sis IJS |2:

n t Phanrozoi: rintprttion. JosphTruisd DuidN. Reznik

fossil rord. Nture 416 420424.
anozoitxonomi divrsity. III.
ns. Plobiolog 10: 246-267.
n s i n r h P h l n r o z o i h i s t o r yo f
stemtis,llog, d th
olumia UnivrsityPrss.
angs in taxonomi dat ass fft
Pleobiolog 19: 4_5 1.
Th doul rvdgrvisitd. In D.
Js., uolutio r Pl l b iolog,
Prss.
tion of ordoviian triloits.Ntr
sis nd volution in diffrnr
ftolog, Pleoolog 127 : 29-227.
.th fossil rord:
Implitions fo
bil Trstios of th Rol
anrozoi: Prolms and prospts.
-745.
lA. Hoffmn' l998. ldgnti
' of Rhizosoli prbrgoii
v th spis|eve|.Pr<lcdigsof th
6-650.
:in nimal divrsity. PalobioIog
i.
lntation of spiation in nozoi
29:4744.
: of Lif using DNA. Gom Biolog
' 1996. o|ectrlar vidn for dp
phIa.Sci 274: 568-57.
What Is Adaptation?
Th farurstlf orgzrnislsar wondrfully suitd to thir nvironmnts and
lifstyls.In sonl ass th mtlr of fatur nd nvironmnt is simpl: a
browr,mlttl]moth is rypti to its prdators whn it is rsting on a dif-
fusly slradd <lwntr trunk. ln othrs th math involvs svral trits
tlrat onrl.lintl prform an imptrtnttask: rat snaks tht spializ in irt-
ing ird ggs lrv uniqu jaws that opn traordinarily wid (vn for
snaks),mouths with srooth gr()Ovsinstad of tth, and spilizdvrt-
ra hirld th rrk tl-rtlrlp pulltllr ggshlls and ush and fold th
mpty shlls fo pulsiln. Ths maths' whih w all dpttios, are
frrndamntzrl osrvationsthat iologistsWant to plain, and undrstandirrg
how tlrstnathsr tnd is l fundnrntalpzzlthat volutionry iolo-
gistswnt tl stllv.
Th list sintifi plantin-tht is, an planation that ould
tstd rvith osrvtior-rand primrrt-plsitd that maths lik ths
mrgdthrough a pross of limination. In this viw animals and plants
rvrrtdwitlr vrit of fatures;thos spistht hrrd faturspr()-
priat f<lrthir partiulr iurrrstansprsistd,whil thos that did rrot
Wnttint, laving us r.ithonl thtls spiswll mathd to thir iruln-
stans.his tlrory ws drivd in prt fom osrvatirrsof th prstsrn
or tintion of vritis of dorstiplents and animls introdud to nW
lotions(.g.,Townsn]'s1786 dsription of glirtsintrodud to Juan F-
njndz lsland: isly l958). Sorrrvaritisthrivd, whil othrs languishd'
and it smdraslnalto onlud that natur sltdsom varitisovr
othrs tlllugh th rmlrinaionsof faturs that distinguishd tlrm, alrd
that natur ould sltamong spisirr th sarrrway.
Th tlrory of daptation vi sltivfiltring trf spiswas prdiatd on
th lnr,itiontlrt th fturslf a spiswor"rldnot hrrgrrnlsshumans
intvndthrough onntratdsltivbrdirrg.Thr was no dout that
th fatursof plants and animals wr mallal;rop plirnts,farm animals,
and pts bor ampI lvitnss that sltivbrdingould hang almost ny

105
106 Adpttil

fatur of a plant or animal. But thr wr ovious limits to mallaility- th gns rsponsil fo
hikns nnot b rd to ly two ggs pr day evry day of tlrir livs-and thy wr in th prviotr
natur smdal to distingurshonly sustantialdiffrsamong spis dvlopmnt did with .
or stoks (Provin 1971; Rus 1979)' Ths substantial diffrnssmd quny of th favord gt
apal of ing produd only y sltivrdingof fam stok (if natur ltion to inras;mor
Was sorting among varitis of donrsti goats introdud to an isl:rnd) or avrag th nt gnrt
original ration (if natur was sorting among spis). ltion ours onsistnt
This xplanation of adaptiltion am inrasingly umrsome s mor ing sltd h:rngsu
and mor fossil organisms wl.disovrd.Fo on thing, layrs of rok that tiotr ats on it vrit
inludd fossilizd spis did not inlud fossilizd matrial from xtant g r a t r h u m r r l e t i v
spis;if natur wr mrly filtring spis'whr Ws th vidn that An daptation is any
xtant spis had vr oistd with tint ons? For anothr' although nti rsons to spi
many fossil organisms or liml rsrblanto xtant ons (.g.'dirrosaurs), pl' w all rpir'1dvlo
othrs smdvry similar to xtant spis(.g.'fossil floundrs).It was un- tmporary ponds. Adap
lar why on spisshould hav gon xtint wlril anothr with vy simi. aus it xplins th
lar fatursshould prsist. v o k i n g p u r p t l s f u ld s i g
Th sintifi rvolution in th stud of adptation is th ida of natural s- mnt for how nw sp
ltion' as suggstdy harls Darwin and Alfrd Russl walla. Although popultions down div1
both mn offd th ida, it ws Dwin who artiulatd it ror ompltly om so tnsivth
and intgratd it mor thoroughly into a largr thory of iologial volu- Sparlrtspis.
tion. As a rsult, w us th trm Drii uolutil to rfr to th pross Darwin's ritial argt
through whih adaptations mrg'a prossalso lld uolutio b trl ould irrdd disringurs
seltiotl or dptiue euolutil. t r a v l s h h a d s n 't l
In Darwinian volution, natural sltion, upld with inhritan of tions of th sam plan
small trit diffrns'is th gnt of adptation. Natual sltionis a statis- ass ths diffrnsr
tial filtring Pross; individuals with partiular haratristis r mor Arlriplago; in oths l
likly to surviv or produ mor offspring than individuals with othr har- i n t h s a m i r t ls p i
atristis.A mor formal dfinition of natural sltionis th ausal ssoi- mad rhs kinds of o
ation of a fatur or trait with fitnss. B fitss w man th numr of H saw all ths 1iffr
offspring that an individual lavshind; individuals that di without rpro- n t u a s f i I t r i l l g i s t
duing hav zro fitnss. The usl ssoitio of feture or trit tuith fit- individul diffrns t
ness mans' oprationally, that a fatur or vlu of rrait is dirtly sustntilulultiv
rsponsil for whthr an individul livs or dis or why som irrdividuals a m o n g p o p u l a t i o n s . }
lav mor offspring than othrs. tion of whthr natural
For xampl' many frogs brd in tmprary plnds tht hold watr for a whethr trait diffrn
rif priod and dry vry sason.Th tadpls in thsponds must orplt and ultirnatly wht
thir dvlopmnt into juvnil frogs for th pond dris and thy ar i n t h i r s | t i r 'r g i m
killd. Tadpols with vry slow rats of dvlopmntdi austhy annot a half.
omplt mtamphosisfor th pond dris. Ntural sltinfiltrs indi-
viduals with diffrnt rats of dvlopmnt so that only thos with mor
rapid dvlopmnt mrg from th pond. vidn for Adap
This filtring would hav no lasting efft if irrdividuals who dvlopd
rapidly wr not gntially distint from thos that dvloped slowly. Th Biologists hirv onttn
fftivnss of natural sltion mgs from th hrital natur of trait mnt and hav tn
variation; in th nt gnration th gnti varints rsponsilfor th fa- lvl. Thy hav als
voral trzritvalus will inras in rlativ aundan. In tlr frog arpl, mk sintifi xpla

obvious limits to mallaility-
dy vry day of thir livs-and
;tantialdiffrnsamong spis
s substantial diffrns smd
rdingof farm stok (if natur
;oatsintrodud to an island) or
,ngspis).
nrasinglyumrsom as mor
For on thing' layrs of rok that
: fossilizd matrial from xrant
:is,whr was th vidn that
int ons? For anothr, although
:to xtantons (.g.,dinosaurs),
(.g.,fossil floundrs).It was un-
n r w h i I a n o t h w i t h v r y s i m i -
laptationis th ida of natural s-
.!7alla.
Alfrd Russl Although
ho rtiulatd it mor ompltly
argr thory of iologial volu-
l uolution to rfr to th pross
;s also alld uoltio b turl
n, oupld with inhritan of
. t i o n .N a t u r l s l t i o ni s s t a t i s -
rtiulr haratristisr mor
than individuals with othr har-
rral sltionis th ausal ssoi-
fitss we man th numr of
rdividualsthat di without rpro-
tion of feture lr trit tuitb fit-
or valu of a trait is dirtly
or dis or why som individuals
rary ponds that hold watr for a
lls in thseponds must omplt
'r th pnd dris and thy ar
:lopmntdi austhy annot
:is.Natural sltion filtrs indi-
rt so that only thos with mor
:t if individuals who dvlopd
hos that dvlopd slowly. Th
om th hritabl natur of trit
varints rsponsil for th fa-
aundan.In th frog ampl,
dDttio 107
th gnsrsponsil for mor rapid dvlopmnt ar mor ommon than
thywr in th prvious gnrationausth gnsrsponsilfor slowr
dvlopmntdid with th tadpols that arrid thm. Th inras irr fr-
qunyof th favlrdgnsauss th avragvalus of th traits r.rndrs-
ltionto inras;mor opis of gnsfor rapid dvlopmntman that on
.Whn
avragth nxt gnrtionof tadpols will dvlop fastr. ntural s-
ltionours onsistntlyovr tim, th avragvalus of th traits that ar
bing sltdhang sustantilly.Th long and mor onsistntl sl.
tion ats or-rtrait varition and th mor gnti vaitior-rin th trait, th
gratrth umulativ hang in trait valus.
An adaptation is any fatur of an organism that has vtllvd through g-
ntirsponsto a spifiologial agnt of natural sltiln.ln our m.
pl' w all rapid dvlopmntrat an adaptation to th risk of dsitionin
tmporaryponds. Adaptation is ntral to Darwin's argumnt for volution
bausit plins th maths twnfaturand nvironmnt withut in.
voking purposful dsign or spial ration. It also provids th vital argu-
mnt for how nw spis aris; whn adaptiv volution taks diffrnt
populationsdown divrgntpaths, th diffrnstwnthos populations
omso tnsivthat thy no longr an intrrd,and ah oms a
sparatsp1s.
Darwin's ritial ergumnt Was that natur's filtr, his natul sltion,
ould indd distinguish srall diffrns among individuals. Dr"rringhis
trvls h had sn, tim and again, diffrns in fatus mong popula-
tions of th sm plant or animl spis in diffrnt loirtions. ln som
assths diffrnswr larg and dramati, as h sw in th Galipagos
Arhiplago; in lthrsthy wr smallr and mor sutl, s in diffrns
in th sam ird spisin diffrnt loations in South Amria. )thrshad
mad ths kinds of osrvations, ut Darwin intrprtd thm diffrntly.
saw all ths diffrnsas points along a ontinuum; rathr than srng
natur as filtring xisting larg diffrns,h saw natur s filtring small
individual diffrns that, lupld with inhritan and tim, produd
sustantial umulativ hangs in fturs and sustarrtil divrgn
a m o n g p p u l a t i o n s .T h t s t i n g o f D a r w i n ' s a f ! ] u m n t ,t h a t i s , t h s o l u .
tion of whthr natural sltion ould as powrful as artifiial siti<rtl,
whthr trait diffrns that wr undr sltiorrwould prov hritabl,
and ultimatly whther populations would dir,rg through diffrns
in thir sltiv rgims, has oupid iologists for naly i1 ntuy and
a half.
vidnfor Adaptation through Sltion
Biologists hav ontinud to dsri th math of tlrganism to nvlron-
mnt and hv tndd our knowldg of that math to th molul
l v l .T h y h a v a l s o u n o v r d s t r i k i n g p a t t r n s i n t h o s m a t h s t h t
mak sintifi xplanations tlth than l)arwinian volution untnill.
1 0 8 Adpttiotl

And through osvtionaland xprimntal str-rdisthy lrav providd th a diffrnt hrnoglobin

answrs to qustionsthat Darwin and his prs ould lnly pondr. mogloin (Fdr and B
Som of th nrost st
paralll adaptation. o
v A R | |E s o F A D A P A I o N
is found in unrlatd ol
Natur prsnts a wid varity of adaptations. arnivorotts plnts our in ganisms display th sa
nutrl.nt-poorsoil and us thir aility to aptur and digst inststo otain ass adaptations rfl
th nitrogn that otlrr plants asorb through thir roots. rtin fish and xampl, two unrlt
rustaeans that our in tmporary ponds and floodplins hav ggs that onditions on two dii
an tolrat -rpltdrving nd an rst in th soil until floodwatrs rak Amria and h i rlmt;
th animals' stivation and indu hthing. Svral spisof fish liv in polar rathr than trra, as do
wts that would fzothr irnimls solid' rlying on a spializd ntifrz on-twntith of th v
protin that tnhiits i-rystal formation within thir bodis (Hhalrka arrd adzrptivsolution to th
Somro 2002). A drirmati ampl
Adaptrions also om in lss dramti vritis.Damslfly spis in th h i s o i so f g u p p i s [ r
genas nllg1|l t'at nountr dragonflis as thir primary prdators tht is rpatd in sv.
hav diffrnt pattrns of movmnt and avio than spis that our aross th island. Gup;
with fish as th primary prdator. Th havilrstht prott against drag- matur at latr ags aI
onfly prdation filitat fish prdation nd vi vsa' so th haviors ar tions' and upstram ft
wll mathd to th prvailing prdator (Pk 7995). Th fungal Ths diffrns ar
pathogn olltotrichm lidemthittm grows on two losly rlatd guppissufi muh hi1
spisof an plants oftrrfoul-rdin th szrmlotions, Phseolus tlltlgris and, in gnrirl,must .
and P' occius, and produs th sam lsions nd sarrirrgon ah spis. from diffrnt drainag.
But th fungal srrains on th two htlst plants ar spializd at surmounting downstram guppis ln
ah host plant's hmial dfnss,just s diffrnt spis of damslflis twrrth upstram
ar spirlizdat ovroming diffrnt prdators (Siard t l' 2007). Pop- a g a i n ( R z r r i ka n d T
ulations of th fish Fudulus heterlclits from mor northn li.rtituds
hav diffnt forms f the nzym latat dhydrognasthan populations
DMoNsRA|
from mor southrly latitr.rds. Th strutur and th rgultion of xps-
sion of ah form is tund to th prvailing thrml rgims to produ sirn- N{any biology txtook
ilar ration rts ross widly diffnt tmpraturs,whih allows th d u s t r i a l m l a n i s mt o i l l
spis to thriv aross wid rang of thrmal nvironmnts (Shult 2005). As tr trunks
2001). oration of th pppr
Biohmial adapttions an partiularly striking zrusrhy oftn in- lak <lvra larg ara
volv th prodution of difft-ltmoluls y th sam individual at diffr- tions of prdation on 1
nt stgsof its lif, suh as th xprssirrof hmoglobin in frogs. Irr most grotrnds; prdtors fou
vrtratsth hmoglobin molul in th lood that arris oxygn to lls and lak moths on lil
is lss abl to hold oxygn undr aidi onditions. This proprty nlrans valu lf th osrvatl
its funtion; lls that ar starvd fr oygn ar aidi nvironmnts,and h- thos osrvatilns wt
mogloin will radily gir, up irs oxygrr to thos ils. But many tadpols mrgd from Kttl
that liv in warm, shallow ponds hav a hmogloin molul with th r. mrkd larg numr
vrsproprty: it holds tightly to oygn undr mor aidi onditiorrs.Ths o u n t d r h n u m l r ro i
:rnimalsliv in vy aidi watrs, and this hmogloin failitatsthir aility This work dmnstrat
to xtrat oxygn from th Watr vn whn thr is littl lf it availal. rats, and tht th m
Thi lls a lss aidi thn th \^ratr'so oYgn is radily rlasdwhr m o l d d 1 's l i l {r s
it is ndd.Upon mrtamorphosis th adult frogs, whih rathair' prss volum).
 dpttio 10')

rtal studisthy hav providd th a diffrnthmogloin gn whos protin havslik most vrtrath-
prsould nly pondr. nroglobirr(Fdr and Brrrggn 1992).
Som f th most striking patrns of adaptation involv onvrgnt or
parallladaptation.onvrgnt a1aptationours whn th sam adaptation
is found in unrltdorganis-rs;paralll adaptation ours whn rlatd or-
:ions.arnivorous plants our in ganismsdisplay th sam adaptatin through indpndnt origins. In oth
:apturand digstinststo btain assadaptationsrfltrpatalmaths of rg:rnismto nvironmnt.For
ough thir rtlots. rtain fish and xampl,two unrlatd frogs that liv undr similaly drrrandinghot dsrt
ds and floodplains av ggs that onditions on two diffrnt ontinnts' Phlllmds suugii in South
in th soil until floodwatrs rek Amri ad birlmntisrmpili in South Afria ah ert uri aid
. Svralspisof fish liv in polar athrthan ura' as do othr adLrltfrogs; uri aid xrtion rqr-rirsaout
i, rlying on a spializdantifrz on-twntithof th watr rquird for ura rtion and rprsnts an
':ithinthir odis (ohahka d adaptivsolution t th nd for wtr onsrvation (Shmidt-Nilsn1990).
A drmati arpl of paralll adaptation is th ontrast twn th lif
vritis.Dmslfly spisin th historisof guppis from upstram and downstram loations in Trinidad
rflis as tlrir prinrary prdators that is rpatdin svrl drairrgsfrom on muntitl ang to anothr
havior than spis that our arossth island. Guppis from upstram loations om rprodutivly
haviors that prott against drag- matur at latr ags nd largr sizs than guppis from downstrm loa-
ld vi vrsa' so th haviors ar tions, lnd upstram fr-rrals
produ fwr, largr ais in ah rood.
or (MPk 1995I' Th fungal Ths diffrns ar mathd to thir diffrnt nvironmnts; downstram
m grows on two losly rltd guppissuffr mu high prdation rats, hav lowr poprrlation dnsities,
am loations, Phseols ulgris and, in gnral,must omplt thir lif yl quikly or not t all. Guppis
;ions and sirrringon ah spis. from diffrnt drainags ar muh lss losly rlatd than upstram and
1 t sa s p i l i z d. l t s u r l . I 1 [ l n t i n g downstrmguppis in tlr sanr ivr, rvhilr indiats that th distintions
as diffrnt spisof damslflis twnth upstram arrd downstram popultions hav volvd tim and
' d a t o r s( S i a r d
t al. 2007). Pop- again (Rznik and Trvis 1996).
zs from mor northrn latituds
: dehydrognasthan populations
DoNsRA|oNs oF NAURAL sL|oN
,u:rnd th rgulation of prs-
g thrmal rgimsto produ sim- any iology ttooks still rploy . B. D. Kttlwll's lassi study of in.
t tmpraturs'whih allows th dustrialmlanism to illustrat th powr of natural sltion (.g.,Futuyma
f thrml nvironmnts (Shult 2005). As tr trunks in Britain turnd dark fror-nir pollrrtion, th ol.
oration of th ppprd moth' Bistl btulri, hangd from mottld to
rrly striking austhy oftn in- lak ovr a larg ara. Th ttooks usully dsriKttlwll's osrv-
s y th sarrrindividual at diffr. tions of prdation on tthrd moths of diffrnt olors on diffrnt ak_
n of hmogloinin frogs. In most grounds;prdatsfound it asy to dttnrottld moths ln dark tr trunks
: lood that arrisoygn to lls and lak moths on lihn-llor]trunks in unpollutd forsts.Although th
lnditions. This proprty nlrans valu of th osrvationson ttlrrdroths has n qustiond y ritis'
n ar aidi nvironmnts,and h- thos osrvations W not th ritiirl vidn. Th onvining data
to thos lls. But mny tadpols mrgd from Kttlwll's mtiulous mark-nd-raptur studis; h
rmogloirrmolul with th r- markd larg nums of moths of diffrnt <llors,rlasd thm, irnd
tdr mor aidi onditions.Ths ountdth numr of raptursof ah typ of moth in diffrnt loations.
hmogloinfailitatsthir ility lris work demotrstrtdtt th diffrnt olor typs had diffrnt nrortlity
'hn thr is littl of
it availal. IatS'and that th math twn motlr olor and bakground was ing
) oygn is adilyrlasdwhr ..Industrial
moldd by sltion(sth alphatialntry mlanisrn'' in this
r f r o g s ,w h i h r . r t h l r , x p r s s volum).
1 1 0 dpttil

Thr ar thlusandsof dmlnstrtionsof rratural sltion in ation. A In th most thoroug

wid varity of ologial fors rt diffrnsin mortality rats or lvls l u t i o n a r y t r j t o yo f
of rprodutiv sussamong individuals with diffrnt trait valus or dif- th most thoroughl st
frnt ominations of faturs.Frlm prdators and pthognstl old strss lss of stram or mou
and drought' frrm studis of snils in British flrststo Darwin's finhs in hav fwr' largr offs
th Gal6pagos' thr is no room to qustion th rality of natural sltion a t i v s d o w n s t m . r
(Hrford t al. 2004). in upstram loations
That natural sltionours is lar, ut an it a lriving for of vo- than half as many offs
lution? Thr ar twtl qustionsaout natural sltiontht ar ruil to s- nworns that r 50-
talishing th validity of Darwinin volution. First' an sltion among pis.Thr ar a num
individuals suffiintly strong and onsistnt to mld th ftr.rrs of or- in whih guppis ar
ganisms? Sond, is thr nough gnti vriation f()traits undr sltion opportunity y introt
to produ sustantial hangs in thos faturs?Two lins of vidn an- t i o n s , i n f f tr p l i a
swr oth qustiolsin th affirmativ. hbitat (Rznik t al.
Th first lin of vidnapitalizson nvirlnmntalltrationsmad y ay Path that in lss
humans and th rsponssof organisrnsto thos altrtions.Th dvlopmnt thos sn in natural
of antiiotis in th 1940s and 19.!0s allowd humans to rat novl nvi- stram nvironmnt,
ronmnts for atrialpathogns.To a atrium' an ntiioti is toin; to d i f f n s m o n g i n
surviv,it must squstrth ttlxin forit plisonsth ll lr dnaturit and moldd nw populti
rndr it harmlss.This sitution rts powerful sltivfor flr altr- xpimnt that Darr
nativ biohmial pathwys within th ll; individuls tht ould dlay or
stymi th toxin's fft would divid nd rprodu mor oftn than thos
that ould not. Slowly at first, ut at an alratingpa, spisaftr spis How Do W Rl
of atrial pathgn volvd fatusthat nald thm to tak on or an-
othr path towrd surviving in th prsnof antiiotis.Although th lls A l t h o u g h s t u d i so f s t
of Strptllcs|ook th sam as thy did in th -l930s, thir ilhmistry lution ours' th full
nd llular mtaolism ar vry diffrnt (Dzidi t al.2008). ar joind with pains
Sustantial iohmial volutitln in rsptlnsto mrdrnhmisty is not ogy. of ours,th [
onfind to tria. ovr 1,000 spisof insts hav volvd iohmial not studid vry stril
rsponssto instiid(Roush and Knzie 1987) nd wdy plant spis m o s t s r u t i n i z d a s
hav rapidly vllvdrsistnto hriids,vn glyphosat'th .,onin a T h f a t u r st h t d
ntury'' hriid (Powls 2008). Many aquati organisms hv vllvd tiularly strong alx
adapttions to anthropognilly produd havy mtals in wtr (Klrks squn of fatur a
and \/is1987) ald many spis of plants hav undrgone tnsiv io. rdution of tos, an
hmial and morphologial volution in rsponsto th prsnof havy th molars. Biomhz
mtals in th soil produd y th fftsof rr-rining, ldd gsolin,and vn p r t i u l f a t u r s ; l
th zin in galvnizd fning (MaNir 7987I. riods, and highr m
In th sond lin of vidn,ilogists hav studid hundrds tlf ass in vgtation lik grass.
whih individuals f th sarspisfrom diffrnt ltlations hiit rdily mt nd th mrgr
osrvl diffrnsin oloratior.r,m<lrphology,lif history, or fatursof lutionary pthWay t
iohmistry nd physiology ut rmain apal of intrrdirrg.Thy hav family of grazing ds
shtlwn that ths diffrnsar hrital and, through a varity of pri- risk from vrity l
mntai approahs (oftn dirtly hanging irrdividualstwnlotions), Suh a ronstru
hav dmonstratd that divrgnt sltilnin th diffrnt lltitlnsls r. longtionof th lim
sponsil for maintaining thos diffrns(Rznik arrd Travis 1996; Tavis a dvlopmntlly
a n d R z n i k 1 9 9 8 ; R z n i k r r dG h l a m o r 2 0 0 l ) . vlopmntal mhan

ns of naturl sltion in tion. A
iffnsin -rortalit1,rts o lvls
rls with ditlrnttr:lit valus tlr dif-
datorsand pathogtlsto oli strss
,ririslrforsts to Dilrwin 6r"rhsirr
ition th rality of natural slti<ln
ut an it b a drir,ing fo of vl-
turl sltionthat ar ruiill to s-
I u t i o n .F i r t . e n . l t i t l n i l l n O n l
nsistntto mold th faturs of o-
: variation flr traits undr sltion
ftus?r'vtl lins of l,idrran-
nvironmtalaltrationsmird y
) thosltratilns. h dvIopmrrr
owd humns to rat novl nvi.
ltrium' n arrtiiotiis toin; to
it poisonsth ll tlr dnaturit and
a powrful sltiv for for altr-
;ll; individrralsthat <llddlryrlr
d rprodu mor otn tharr thos
lratingp' spisaftr spis
a rrldtlrnrto tk on o an-
of antibiotis.Although th lls
id in th l 90s, thir iohmistry
( D z i d i t l . 2 0 0 8 ) .
s p n s t t l n l t l d r n h m i s t r yi s n t l t
lf instsar.voIv]iohn-ril
nzte |987):rnd wlyplant spis
is, vn glyphsat,th ..lnin a
r aquatl oganisms hv r,olvd
:d havy lntals in watr (Klks
nts hav unlrgontrrsivil-
rsponsto th i]rsnof havy
f rining,lddgas<riin, nd vn
9871.
; h a v s r u d i dh u r r . l r dtsl i a s si n
r diffrntllationshiit radily
lholog,lif history, o ftursof
rplof intrr1ing. Thv hav
and, through a varity of pri-
ing idividuIs trvnJoatior.rs),
,n in th diffrnt lotions is r_
; ( R z n i ka n d T r v i s l 9 9 6 ; T r a v i s
r r2 0 0 1 ) .
dpttiolt 111
In th most thorough studis iologists hav r-ratdth prsumd vo-
lutionarytrajtory of th diffrns.Th Triniddian guppis off on of
th most thoroughly studid xarnpls.Rall that upstrm guppis, rgard-
lssof stram or mountin rang, liv llngr,matur ltr and largr, and
havfwr' largr offspring at ah or-rtof rprodutiorrthan thi los rl_
ativsdownstram. Th magnitud of ths diffrns is not trivial; fmals
in upstam loations ar-r mor than 20" |argr at mturity, hav fwr
than half as many offspring as downstram guppis, and produ individual
nwornsthat ar 50_75% largr than tos produd y downstram gup-
pis' Th ar a numr of upstram loations aov sustantial watrfalls
in whih guppis ar asnt.Jolrn ndlr nd Dvid Rznik ploitd this
opportunity by introduing downstram guppis into som of ths loa.
tions,in fftrplitingtlr likly ours of guppy oloniztion of upstram
habitat(Rznik et a|.1.997).Th introdud guppis followd an volution-
ary path tht in lss than si yars produd trait valus omparal with
thos sn in naturl upstrarn popultions. Natural sltiorrin th up-
strmnvironrnt' ating on th gnti variation that was produing small
diffrnsamog individual downstram guppis trnsplantd upstram'
moldd nw populations of upstram gtrppisirr vry short ordr. [t was th
primntthat Darwin would hav likd to do.
How D.W RonstrutAdaptiv volution?
Although studis of sltion and rspons dmonstrat that Darwintn vo-
lution ours,th full pitur of adaptation mrgswhn stdisof sltion
ar joind with painstaking work in grrtis,dvlopmnt, arrd plontol.
ogy. of ours' the pitur is nithr omplt nor prft; iologists hav
not studidvrv striking faturof vry organism, and vn for som of th
most srutinizdass' thr is still muh to larn.
Th tatursthat dfin th modrn quids, horss and zbras, offr a par-
tiulrly strong mple (MFaddn 1992). Palontology has rvld th
squnof fatur apparan from th longation of th lowr lims, th
rdution of tos, and th mrgnof hoovs to th inrasing hight of
th rnolars.BionrhniaIstudis hav pinted to th dvntagsoffrd y
partiularfaturs;longatd lims failitat rapid running for xtndd p-
riods, and highr molars ar fiial for grazing touglr, silia-ontaining
vgtationlik grass. Plaing ths hangs in th ontt of a hanging li_
mat and th mrgnof savatrnasand plains, w an ronstrutth vo-
lutionary pathwy through whih small anstal rowsrs produd
family of graztng dsndants mtlldd y living irr opn Ountry nd ing at
risk from a varity f larg prdators.
Suh onstrutionpsumstlrat th transitionsw ronstrut,lik th
lngationof th lim and th rdution in th numr of digits in th foot,
ar dvlpmntallypossil.That is, th rtlnstrutionPrsums that a d-
vlopmntainrhanis-rftlr produing suh hngs n dsrid, and
1 1 2 dpttil

th th gnti vri:rtions in tlr lrnntshrlr-rglr wlrih thr mhtlism qustions has inspird
tltrld t r'"'tluldhv n radily vailal. Although W anllor tst ths ian volution' vn a
prsumptionsirr tint spis,W-ilIl quir insights frtlm str-rdis of ist-
ing spis.
A4lsttrrstrialvrtrtshav fil, digits ln th flt at tlr nd of ah ontrovrsis in l
limb, ut th position of thos digits on th fo.tt,thir lativsiz, and vn
|s HE RAW
thir nr-nr ,.1nviv. Stu1islf ftlltand ligit dvltlpr-nrrt
il] tratal -
READILY AVAII
primnt:rlnimals lik salamandrsnd hikns hav shown that th num-
r, siz, and position of digits ar dtrmind},moltrlrsigr-rals harrgd At first glan th ans'
1,mryoni lls ]urirrgal,vlim dvl<lprnt (Shtrin2002)' Rlativly ttion is rdily avail
small liffnsin rh timing, inttrsity,and pattrrringof thos signals guppis offrs ompt
hrrgth sr.rs1ullt pattrn of digit dvloprrlnt.Dlays in signaling lvill sstantial adaptiv
it-llpdth forr;rtilnof th third, fotrth,and fifth digits, and if thy dl d- x p t i o n a l s :h u .
vllp, tlros digits an quit srll. Dlys irr signaling, opld with ellular onntrtio
gr()\^,,thin th foot itslf, will ltr th ositi<lno digits on th flot.Thr"rsl- h i n d l i m l n g t ho f t r
thtlr'lghW nn()trtath prist1vltlpnlntal pth that ld to hrofin found apprial lv
fossil quids' W l111lffr :r plausil rnodl through whilr umulativ viduals within a singl
hllrtgsin signaling rrd gn prssilttor'rlll-ravtrltrrsformda fiv_t()d B u t h i s r n o l
ftlot inttl :r sillgl-todhoof. that natural sltin
\zilullglrti vrition f<lr t ky dvl<lpmntlimnts hav lln shar dvlopmntal
vilirl?T.hst llalog,v()lllsfrtllrl stulisof rlativ lrindllnl lngth.in valus to inhritd
frogs. Frtlg spisvary normously in th lngtlr tlf th hidlil lmpard to sltion. In svr
with th:rtlf th ody; rikt frogs hr,hindlims sustantiall1,longr tlran lis, swordtails, and
ti tld1.lrlgths,r'r,hrsspadflor-ttladlrindlims ar raly mo than an ndorin pathw1
:rout lrlf th ldy lngtlr.vn losly rlatd spislik th arking tr opmnt (Kllmnl9t
frog nd gl1tr frog lf th slutlrlrstrn Urritd Sttsan hr,sustn- soition btwn r.t
tiallr- diffrltlativ hindllm lngths.Ths diffrt-ls ris from diffr- h i s a s s o i a t i o n r a
ns in th timing of lim-ud initition and th rat of lim.ud growth in largr ody sizs at
th tir1p<lls (Blouin |99 I). Gnti vlrriatiorrtilr th rlativ lrrgthof th If slowr maturation l
hi]lim app:rrsradily l'irilal \, in sitlgl frog popultions (Blluin solution; tlr gnti
1992I. This work suggststhat th raw matrial for hindlim volutitln in maturation is not imr
fr<lgsrr.sponlrlv not liffiultfo sltilnto find in tl-rpast arrd, ,- - Th ky point v
tnsitln,suggsts silnila lrrlusionflr othr anin-rals. Straint hold in th l
lt is sy to ritiiz lnlttsionslrout adptation in htlrss whos pr- suffiintly long, wor
sllnlpti()nsl111 tstdonl\, tlrr<lugpri-ltltl studis o irds or fogs. ths traits? If sl
Suh trapoltion u'ill alw1,s nssarywlrn w attmpt t() ronstrut m u t t i o n s f i l i t a t t
adaptation orr a larg sal, suh as might f<lundin th fossil rord. It is and rapid maturatio
alstl asy rt>ritiiz't gnrality rlf onlrrsionsillut ada1rtatilndivd though it ay wll h
frll study on a snrall srl,suh as th as tlf industrial lnisrn. l a m aris, it is imp
riti, th rid lrIrgin th frqunr,of altrrativforms of :r singl gn wld favor suh
ftl lllratilnis lradly vidn tht naturi-llsltilt-l an div profound T h p o s s i i l i t yo f .
transformtionsin orgnismi f:lturs-l(lharrgingollr pttrnsin moths of adaptationsthat ;
tll rrs ntluglr tl undrstand h<lwvolrrtilnmol]sfish musls ir-rtolrri xampl' most asS
()rgns<lrtnsfln]splant lavs itlto taps ftl atlring and digstig in- h a v si n a p a r t i u
sts? h diffiulty of alaning what an ahivd xprimrrtallvat sum a partiular p
smallr sls ig:rillsr th l:rrgr.sals of th rrlost importnt volutilrlary oloration ar filrr
 dpttil r1)

ts througlr whih that mhanism

qustionshas inspird and sustaindonsidralontrovrsyout Darwin-
ll. Although W annot tst ths
ian volution' vn among sintistswho study it.
qLririnsightsfronl str-tdis
rt-is-

gits on th foot at th nd f ah
ontrovrsisin th Study of Adaptation
t foot, tlrirlirtivsiz' and vn
i digit dvlomntin tratal - |s H RAW MARIAL FoR ADAPA|oN
:hiknshar'shlwntht tlr num- RADlLY AVAlLAL?
ld y mtllrrlrsignlslrngd
At first gln th answr to th qustion whthr th rw nrtril for dp.
l t l p m n t( S h u i n2 o o 2 I . R l a t i v l y
tation is radily availal would sm ovious. Th study f th transplantd
, :rnd pattrning lf thtls sigI-rals guppis offrs omplling vidn that th grrivariation rquird for
.lopmtrt.Dlays in signalirrgr,l.ill
sustantiladaptiv volution ws avilal. This dos not appar to an
nd fifth digits, and if thy do d-
ptionalas;hundrds of studisof a varity of haratrs,ranging from
)lys irr signaiing, oupld u,ith
llular onntrations of alohol dhydrognas in flis and th rlativ
tion of digits on tlr foot. Thus al-
hindlim lngth of fogs to th olor of swallowtail uttrfly pupa, hav
pmntirlpath that ld to a hoof in
-rrodl found appriallvls of gntivariation for individual traits among indi.
through ,lrih umulativ vidulswithin a singl population.
:ould hav trnsflrml fiv-tod
But th istnof hrital variation in singl haatsdos not imply
that natural sltionan mold fatrsaritrarily. Diffrnt haratrs an
vlopmntIlmnts hav rr
shar dvlopmr.rtal pathways that aus partiulr omintions of trait
-rdisof rlativhindlim lngth in
vlusto inhritd togthrand thry onstrain th immdiat rspons
. lngth of tlr hirldlir
mpard to sltion.In svral spisof pliliid fish-th fmily of guppis, rnol-
i n d l i m ss u s t a n t i i r l livo n g t a )
iies,swordtails' arrd mosquito fish-th pross of mturation is govnd y
d h i n d l i m sa r r a r l y m o r t h a n
an ndorinpathwy that rdusgrowth rat as it alratssual dvl-
:latdspislik th arking tr
opmnt (Kallman 1989). Th gntivariation in this pathway ratsan as-
LjnitdStatsan hv sLlstlr-
.hs soiation twn rapid nraturation rat an] sm:rll ody siz at mturity.
diffrnsis from diffr- This assoiation mans that th rspons to a sltion pfssur that favors
nd tlr rt tlf linr_dgrowth in
largr ody sizs at maturity will invitly inlud slowr maturation rts.
tion for tlr rlativ lrrgthof t
If slowr maturation rats ar dtrimntal for an rason' thr is no ovious
n singl frlg popultions (Blouin
slution; th gnti variation for th omination of larg siz and rapid
rtril ftlr hirrdiim vllrtion irr
maturation is not inrrnditlyvailal.
on to find in th past lrnd, y -
Th ky point rvolvs around th word immedit'l; dos su a on-
t h ra n i m a l s .
straint hold in th long run? /r w to wath suffiir-rtlylosly and wait
adaptation i hoss rvhtls pr-
suffiintlylong, would W se mtlttios in individual gns that dissoit
rimntalstudison irds or frogs.
thstraits? If sltion atd with suffiint Onsistny'would ths nW
y rvhnW attmptto rostrut
mutations failitat th vntual mrgn of th orninatiorr of larg siz
flundin th ftlssil ror]'It is .W
and rapid maturtion? hv no simpl answrs to ths qustions; al-
l u s i o n sa o u t d a p t a t i o nd i v d
though it may wll that givn nough tim, a mutation of th right kind
s of industrial mlnisnl. Tl a
may aris, it is impossil to prdit if th ologial irumstans that
altrnativftlrms of singl gn
would favor su a mutation would still prvil whn it finally aros.
rral sltionan driv prlfound
Th possiility of onstraind gntivariation provoks a ronsidration
hrrgingollrpttrllsin moths
of adaptations that a uilt on assoi:rtionstwn individLrl trits. For
n rolds fish musls irrto ltri
xampl,most ass of rypti oloratiln in animals work only if th animal
ps fo athing nd digsting irr-
bhavsin a partiular fashion, wlrthr it is to main motionlss or to as-
n alrir,dprinrrrtallyt
sum partiula postur or intatio.isrnaths twn bhvior and
th most importnt volutionarv
oloration ar failurs. T adnt advoat of Dar,inian volution might
1 1 4 dpttio

argu that natural sltion hs assmldth omponllts of suh adapta. possibl solution ontl
tions via mtiulous molding of individual fi1tursuntil th st omina- s t i l l u s t r t i o n so f t h i
tion mrgd from all th possiilitis.At th othr trl n riglrt gu frnt poptrltins of t
-
that th ontd inhritan tlf th onrponntsoffrd natural sltiona ologial hallng.
limitd rng of rarv mtrial frorr-rwlrih only a fw good maths ttr ir- ids; diffrnt popul
umstanswr pssil. i o h m i l n d p h y s
Th sam argumnt n prov<>kd y onsidring th volution of ny (atsumur and Voss
puttiv rldapt:rtionthat involr.s rultipl haratrs, it ;l iohmil ltri fishs o-pt
patWay or a suit of assoiatdhavitlrs.Th ontrovrsy is not Whthr o g n s ( B n n t t1 9 7 1 )
suh fturs rprsnt adptlltirns;th ontrovrs is vr h xtnt t Th othr part { th
whih adapttion is onstraind y th natur of th vailal raw matrial. ural slti<ln filtrs r,
This is L.linglrpprolrd fr-l sval prsptivs, fror rinirrg th na- prior volution. As a l
tur of nW mutations for ky haratrsto disstingth ssoietionsamong tion, arrd history an g
individual fatursof an adaptation inttl thir gntiontrrland sard d. proably not what al
vlpmntal pthwavs. wr bing dsigrrdd
lind spot ausd y t
i n f f i i n ts t r u t t t r i s
ls ADAPATIoN PRF?
organ and susqunt
Th rrrarvlousmaths of orgnisrrrto nvirtlnmnt offr a strong t]pta- tion. An asir nd bt
tion to onlud that adptation is prft. Indd, whn ontmplting in- not through th rtin
triat adaptatins lik ag-spifihmogloin gn prssion,th amzd Thr is anothr Wi
sintist is rr<ltvr,v diffrnt frorn th awstruk plt or th inspird rd by volutionary
thologin. Brrt th sintist must tak this qustion es tstbl id. faturs of organisms
Thr ar two oprationl wys in whilr w rnight us th tert prft; it prforrn a spifi funt
ould rf to whthr vry fatur of an lrganism hs adptiv signifi- that aptur and fous
an nd/or wlrthr t matl-rrf orgirnism ttl tlvirtltlIntis as gool as molular signls intt
possil. bn onsidrd am
Nlt vry fatur of an organism is adaptivly signifiant.F.or olr thing, of ths omplx trait
onrtd inhritanof haratrs,as dsrilprviousl1,,mns tht som of Darwiniirn vo|uti
faturswith no adptiv signifianmay prsist austhy ar inhritd vry thm in th stu
throtrgh shrd dvlopmntl pathrvays rvith othr fturs that rrsnt
an adaptation. onrtd inhritann also pr()ttftursthat r dtri.
AN DARWlNI
mntal y thn-rslr,s if thy ar inhritd alorrg wit adVanta1]rous ha.
xPLA|N oM
trs and th nt fitnss fft of tlr mination is plsitiv. This is th
ssIlof rnthtlr for th volution of snsn;gnswith dtrimntal Darwin himslf appl
ffts at latr ags. aftr a sustantial rprodutiv priod, rmairr in th dilmma is tht all sr
population austhy onf a larg fitnssdvatagarly in lif, for boli pathway lik tht
or rly in an <lrgnism'srprodr-rtiv priod. psit f rany indil
Is th mt of fatur to nvironmnt as good as possil?Oviously th proprly intgratd f
answr to this qustion dpnds on lrorn,w dfirrptssible.Ad tlrrin lis ring so lany things
th nswr, at last in part. Ntural sltilndos not dsign an orgnism or ntural thologian's z
its fatus;it mrly filtrs xisting variation. Th nd produt of Drwirrian sam position tirkn l
volution is lways :ls good s possil, ut hr plssiblis dfind as th tnt thsis offrd y
st of th availal options' whih ar dtrmind in tun by th gnti th y or th Krs l
varition that is availal nd .hat th onsturintson that vaiati<lr-r might ltitln. Chnging th
. Put anothr way, adaptation is ontingnt pross;it onstrtttsth st nothr (Aala 2006

d th omp<lnnts of suh adapta-
al ftursuntil th st omina.
tir othr tmln nrighr agu
ponntsoffrdnattlral sltion a
h only a fw good nrathstO ir-
y onsidringth '',<llution
l hatrs, it iohmial
s' Th ontrovrsyis not whtlrr
ontrovrsyis tlvr th tnt to
Lturof tlr availalraw matrial.
sptivs'from amining th na.
. rd i s s tl lrg
t h s s o i l t i t i l l 5. l t . l t ( ) n g
hir gntiontrol nd shard d.
vironmntoffr strong tmpt-
t. Indd,whn ontmplatirrgin-
;loingnprssion,th mazd
awstruk pt or th inspid
this qr-rstionas tstl ida.
Lw might us th rcrm prfet; it
n orgnism has adrrptivsignifi-
tisnr to nvir<lnmntis as good s
aptivly signifiant. For on thing,
:ridprviously,mns that sol.l-l
l prsistaustlr ar inlrritd
with othr fatursthat rprsnt
also prottftursthat ar dtri-
long with advantgoushara-
ninatiln is positiv. This is th
snsn; snswitlr dtrimntal
produrivpriod, mirr irr th
rssadvarrtagarly in Iif' for
od.
rs good as p<lssil? oviousl}. th
,,dfin possibl. And thrin lis
o n d o sn t d s i g , a
n n r g a n i s n lr l r
orl. Th nd produt of Drwirrin
)ut hr possibl is dfind s th
Jtrmindin turn y th gnti
nstraintson that variation might
'gntpross;it onstrutstlr st
dpttion , t.)
possilsolution ontingnt on th rw matril providd y mutation. Th
stillustratinsof this ntingny our whn diffrnt spisor vn dif-
frntpopulatilns of th sarrrspis rspon.lidisynrtially to tlr sm
ologialhallng. This has ourrd in volution of rsistan to psti-
ids; diffrnt population of th two-spottd spid mit volvd diffrnt
iohrialnd plrysiologial mhar-rsims to op wit th sam pstiid
<lfany (tsr.rmuraanl Voss 1964| and, on a largr sal' whrr diffrnt spis of
ltri fishs o.optd diffrnt musls for transformation into th ltri
organs(Bnntt1971).
Th othr part of th nswr is mor sutl: adaptatin is historial. Nat-
url sltion filtrs variation in faturs that ar thrnslvs produts of
prior volution' As a rsult, thr is a signatur of history on vry adapta_
tion, zrndhistor,vn gnratfatursand lrninationsof fatursthat ar
proaly not wlrt an intlligent dsignr would produ if th orgnism
wr ing dsigndd novo. For ampl, th huran y has a sustantial
lind spot usd y th thrding of lood vsslsthrough th rtina. This
inffiintstrutllris a historial artifat of tlr y'sorigin as a light-snsing
organ and sus1unt loration of th rtina as a Srnfor imag rsolu-
tion. An asir and ttr dsign would wind th lood vsslsaround and
not through th rtina.
Thr is anothr way in wlc perftio i adaptation has n onsid-
rd y volutionary irlogistsnd ritis alik. ompl adaptations-
fatursof orgarrismsth:1tar uiIt fom may haratrs tht ombin tl
prform a spififuntion' lik th vrtraty, with sustsof haratrs
that pturand fous light, that dtt rightnssnd olor, and that rsolv
molulr signurlsinto an aurat irg of rh trnal world-hav long
..prftion.''xplaining th volution
nonsidrdampls of natur's
of thsomplx traits has always n a ntrl hallngfor th hampions
of Dwinin volution, an] th answrs to th;rt hallng illustrat narly
vrythm in tl.rstudy of adaptation.
AN DARW|N|AN voLUloN RALLY
PLA|N oMPLx RAls?
Darwin himslf appriatd th h;rllng posd y ompl traits. Th
dilrrrrnis tlrt all sulr faturs,whthr n organ lik th y r mta.
oli pathway lik th Krbs yl usd y lls to produ nrgy' ar a om-
posit of many individual adaptations' all of whih n-rust psnt and
proprly int ] atdfor th adaptatin to funtiorr.ow l-lnatural sltion
ring s many things togthrall at on and mak thrn work proprly? Th
natural thologian'sanswr in th nintnthntury was that it annot' th
samposition takr-ry mo1rnproponnts of IntlligntDsign' Th onsis-
now, is tlrat th lrnplxity of
tnt thsis offrd y Darwin's ritis, thn r-r1
th y or th Krs yl is too grat to hav n uilt by mutation nd s-
ltion.hanging th olrrr of moths is on thir-rg. ut uilding n }.is quit
anothr(Ayala 2006).
I16 dpttit'l

But is it? Drwin argud that ni]turalsletiolroul1produ tlrrrpladap- hagfish). A sond intr
tations if thy volvd through a sris of srall stps, ilh of whih had sm ians, rptils, nd irds t
daptiv advantag ov its prdssor.Th v<llr.ltitln of th 1-ltrldonlv fish, and th dditilrra
valdy ronstutingth hist<lrialslutlof thssmll hangs,and from ons assoiatdW
an undrstandingf lvhy an y lllksas it dos rquirsan ttndrst:rnding of Ths would rmark;
tlrat sar.nsignatur of history (.Wlrrand Dpw 2004). To th modrn volu- tion drov th wholsal
tionary iologist,th ferilurto appriatth onringntand historilnaturs shaps and rting ntl
lf adaptation is th signal w,knssin th ar]ulnntstilr IntlligntDsign. z 0o7l .
Darwin illustratd lris argmntflr th v 1,pointing otrt th divrsity of But th diffrns rn
potosnsitivstruturs in diffrnt spis tlf gastropods (snaiIs,russls, st. Gill arhs ar a (
squid, and otpuss).Ths struturs reng frot.nlr-rstrs of lls in sor s a llows. Fish us t
lams that an disrimirrt twn light lld dak to t spistiatd, avity. .Whn a fish opl
mra-lik ys of an Otopus' with a rng of phltosnsitir, ogns of in- oral avity inrass,an'
rasing mplxitv in twn. Drr,vin grrdL.l1. nlogv, ssrting that bllows ontrts and r(
tlr modrn divrsity of phtlsnsitivtlrgans in gastropods illustrats th watr through th gill sl
stpsthr ould hv orlfror ansrort() dsndantto uild tlr most stors of all iawd vrl
omplietd of ths orgns' th otopus y, with a omparal plssiility trorne prtils into t
for uilding th vrtrat }l.In this r]umlltsn-rallhangs agrinstth in th oral vity, whi
akground of what hs ll for n aumult into a rrrrarkalrl also funtions as a rsp
trirnsforrrrationof lls nd tissus. wll supplid with blo
Dar."vin'sargumrlt,whil rvalinghis grat knowldg of anirl divrsity thm, and th thin tissu
and his rathtakingrtivity' did rrltnvin th ritis d, in f;.rt,dls from th watr into th
ntlt lrrd itslf to a rodrn vliuti<;naryirnalysis.Th spisinvllvdin the drivd from th gill a
argumnt ar too distantly rlatdtO on nothr for skptialt:rsts; th fssil gasp and pross fod
ord arrd th inrpliatinsf omplrrativr-nolulardta slrgstthat it hs n h a n st h t r n s m i s s
n tns of millions of yars sin many of thm shard ommon anstor on th outr surfa of 1
(prsumirrgthr on vn gir'srlno th ida f vltrtion).Th vllu. gion of our skulls'
ti>naryPathway to th Otoplls y has long sin n lost. ]Wwould hv t-to ow an suh divrs
bttr luk trair-rgth pthwirv to tlr vrtrt1';ntrnlrv tlffrs larg rats, volutionaril
group of organismstlrat all lravwll-dvlopdyspt fo thos living in fish ar prt Of a rp
vry drk nvir<lnmtrts that sl to hav son]rilvolv]rdud ys. musls that opn arrd
vn in vtetsth progrssiolrfrom simpl to trtllplys ourrd irr that pand and ontra
som unknown st of ilnsttlrsthat livd ovr 400 milliln yrs ag(). arhshav th funrio
Th lilrnrafor rrndrstarrdingth vtllutiol-lof tllplity is that it o- grasping and grinding
urs on a timsal that is vry long onrpard rvith th timsal on whih t h t r a r l s f o r m a t i o nf o
mutationl vrition irl sirrglhirratrriss or natLrralsltionhausts so dramati as to s
th vailal variation in a singl haratr. But vn whn w rsort to hangs.
brad ornprisons <l indit infrns,it is oft tlr s tt w arl Th trnsformation ]
mak slmvry strong infrnsin partiular ss.!7 rviw two lf thos is lss dramti than it
ass :rusthy illtlstrt htlw ,an omin dt fr<lnldiffrrrtlif- spis.Th fossil ro
sin disiplins to dviop insights int<lth tlrigin of or-nplstuturs' isms that rprsnta tl
vn thLlghthis origin is not dirtly osval. th rptils tl-ratpp
omposit of svn ol
mpl 1': Fro gill rhes to er ossiles d js skull was formd by t
An old infr frtlm th ftssiltlrd is tht vrtIit jr'vswr drivd on on th skull (Figr
from th gill rhs tlf j:rwlss fish (rprsntdtoly y lan-rprysnd dntary, and artiula
 dpttion ft/

tion ould produ onrpl adap- hagfish).A sond infrn is that th singl middl-ar ossil of amphi-
;rnallstps,h t>fwhih hd sllr.r ians' ptils,nd irds rvas drir,dtrom an lmntof th jaw suspnsionof
r volutionof th y ould only fish, and th additional twtl middl-ar ossils of mammals wr drivd
]qunlf rhssmtrll hnges, lrd fronronsssoitdwith th |aw joint of tir rptilian anstors(F-igurl )'
t dos rquirsan undrstandingof hswould rmarkal voltionary vntsin whih mutation and sl-
Dw2004)' To th modn vrllu. tion drov th wlrolsaltransformation of haratrs' molding ntirly nw
.htlrrtingrrt
and lristoril naturs shapsand rating ntirly nw funtions (!7ak 1979; Radinsky 1987; Luo
fgumntsft, IntlligntDsign' 2007\.
y y pointing ollt th divrsity lf But th diffrns in funtion rrrak sh a progrssion sm fniful at
- . i so f g s t r o p o d s( s n a i l s 'm u s s l s , st.Gill ahsar omposit of ons, musls, and nrvs that funtion
ang from lustrsf lls in som as a llows. Fish us thm to xpand and ontrat th volum of th oral
ht nd drk to th spistitd, .!7hn
vity. a fish opns its mouth, th llows xpands, th volum of th
n g o f p h o r o n s i t i vor g l r n s l i i n - oral vity inrass,and wat is drawn irr. !7hn a fish loses its rrrouth,th
argued y analog1',asstin]tht bllowsontrats and rdusth volum of th oral avity as th fish pls
rgans in gstop()dsillustrats th Wtrthrough tlr gill slits. ln living iawlss fishs(and prsumaly in th n-
tor to dsndantt uild th most storsof all jawd vrtrats),this pump is a fding dvi' drwing wa-
y, rvith a omprrlpossiiliy trornpartilsinto th mouth that adhr to a stiky sustanprodud
.gumntsrnall
hrrgsagainst th. in th oral avity, whih is thn onvyd to th gut. Th ntir apparatus
n aumult ir-rtoa rmarkrl also funtions s a rspiratory orgn. Th tissusthat lin th gill ahs ar
wll supplid with lood; th pump passs frsh, oxygntd watr ovr
3raknowldgof aninrzrldivrsity thm,and th tlrin tissstlrt lin th arlrsallw oygn to trrrsportd
nvin th ritis and, in fat' dos from th watr into th lood vssls.By ontrast, th struturssupposdly
rrlysis.Th spis irrvllvdin tlr drivd frol th gill rhs lrav mrrr iumsrid funtions. Th jaws
nothrfor skptiaItasts;th fossil grasp and pross food. Th first rniddl-arossil in trrstrialvrtrats
nrtllulrdt sr-Iggst tht it hs nhansth transmission of th nrgy in sound wavs fronr th ardrum,
f thm shad a ommo anstor on th outr surfaof th ody, to th innr ar, whih is in th tmporal r-
l th ida lf l.oltrtitln).
Th volu- gionof our skulls.
g sine nlost. ! would hav no L{ow an suh divrs struturs'not all of whih ar prsnt in all vrt-
'rtl.;nttlr nlu,offs
a lag rats, volutionarily linkd to ol1anothr? Th iaws in th aly jawd
iopdyspt1thos living in fish ar prt of a rpatd sris of ons that inluds th gill arhs. Th
' slndariiyvolvlrdued
ys. rnuslsthat opn and los th jaws funtion in a similar fashitln to thos
iinrpl to ornpl .ysourrd in that pand and ontrat th gill arhs.In som living spisof fish, th gill
'vr400 million yarsi1go. arhshav th funtiollal quivallrtof tth and funtion as do our jaws y
lluti<lnof omplity is that it tl- grspingand grinding food. Givrr th pattrns of musl nd innrvrion,
rard rvith t timsalln whih th transformationfrom gill arh tO jaW suggstdy th fossil rord is not
rrissor ntural sltiorrxhausts so dramti as to sm an unlikly urrrulativfft of small, attain:rl
:tr. But vn whn w sot to hangs.
, it is oftrrth lls tht w n Th transformation from jw support in rptilsto r ossii in mar.rals
ulr :rss.\/rviw two of thos is iss dramati than it might appr from mparisorrsof only th isting
on-rindilta flr diffrnt lif- spis.h fossil rord offrs striking pattrns of lrng aross th organ-
' th origin of ompl srruturs' isms that rprsnta transition twnrptils and mammals (Figur 1). In
rval' th rptils that appar to anstrl to rammals' tlr lowr jw ws a
ompositof svn ons and th artiulation twnth lowr jaw and th
ls lld itus skull was formd by th artiular bon on th lowr jaw and th qudrt
that vrtratjaws wr. drivd onon th skull (Figur 1A). In mammals th lowr jaw is a singl bn, th
sntdtoda v lmpys irrrd dntary, arrd rtiulation of th j. tlss th dntary on itslf nd th
1 1 8 dpttitl

D{ondy| sqrramosal on on th
a0uI
delhmeil
t/-
sids (th sintifi trm
jaw hings that onsist
surangu|r
squmos|
I
middle
th lowr jaw and th q
on!at
Adutt
ear
1B_). This dual artiu
quadrtojugaI
. Monodlphis faturs of th sklton,
mbryoni rptils and mammals. I
and squamosal ons fil
{= rplad th artiular ar

=_ on th artiular an
o
of jaw and skull, thy v
fossils nd from th ml
mals indiats that th
and orintation' and
l<
l
I
| I

) o () Brsilith rim rep
.
; Triassi through th lat
3s.*. Pchgnlus (B) but rt
D
(D) Sinocoldon was '

from th arly Jurassi,tl
q though th individualo

f
GI (Lft) orguo
late Triassi thrugh th
a 8 r a t l yr d u dp r t i
To th right of ah ol
tht th stapdialpros
prossrtiulatswith t
( ) ( R i g h t ) T h i si s a v
D.ond|e
orgucodoz, illustrt
SO-glenoid
srisof ons inluding
quaoratolug|absenl
bons hav a diffrntn
animals and thy an ls
Figr-rr1. This figur, modifid from Luo 2007, illustrats th volution tlf th
totympaniis a ring o1
manrmlian jaw, livlfrom :r sris of ftlssils in th synpsid linag (th linag irl
twn th outsid wor
wih mmm:rls mrgd) nd fro.l th mrylni dvltlpmnt of t living marals.
(F) Ths ar adult an
Th lrawings on tl-rlft (A-D) rsnt th l<lwrlfi vrrtral (Ottom) suri of tlr
rnitb orhchus. |n te
aniu.t,as illustrtd in th drawing in th low-lft orn at th otttlrn of th fisur.
mdial surfaf th lov
Th figurs tln th right (_H) rprsnt th mdial (irtnr)nd vrrtral surfas of lr.
totympani,mallus'
ltlu.r jw. h numrs rprsllt rltiv ags, wit tlr lowr numrs indiaring oldr
onfiguration sn in th
flssils and th highr numrs inditing rnt mammals.
(G) Th lowr drawin
() Prlirlgthlls ws iat Triassi Synapsid tht is r.rotin tl-rdit
anstral lin adult jaw of Ylood
to mammls; thr was a tl.rplartiulatitlr.rof ltlwr jirw nl rniunr involving th
.,mllus'' Th ondition of th
(tl-rrodifid artiLllr orr) ln th low jaw and tlr ..inus'' (th rnodifid
duk-illd platypus.
quirirt orr),quirdatojrrgal,d squamosl ons of th uppr jw,.
(H) Th lowr drwin
(I\| h1'gelsrpsnts a difrrrtrnh of th Syri;lsids tht dats tl th lt
oPossum' ol delph us
Trissi_rlJurassi priol. This group ht-rd duill rtiultion in whih th ..n-ra]lus',
mryoni jw is vry sr
On th loWr jirrv artiultd with th ..inus'' on th Llpr jaw and th dntary on th
Ynoonldoz, with th
lowr jaw artirrlat1with th squarosal on th upp jw. Thr Wils l1o 1uadrtljugal
th lowr jaw, whil th
on.
kl lmntsing

ossified
M.L
sepltion
3g;,".^
lustrtsth volutiln of th
n th synpsid lirrag(th ling in
Li dvltlpmnt<lfth living mmmals.
r lft vntra] (tlttom) sr-rrfaof th
r-lft ornr t th ottom tlf th figur.
al (inn)r.rdvntral surfs of th
vith th lw numrs rndiating ldr
nmmals.
sid that is not in th dirt nstral lin
: lowr jaw
nd rniurrl ir-rvolvingth
w jaw and th ,,inus'' (th mtldifid
tlnsof th uppr jaw.
of th Sypsids tht dts to th lat
lul artiultion irr whih th ..mllus''
th uppr jaw ni th dntary on th
uppr jw. Thr rvas no quadratojug1
dpttio 119
squmosalon on th skull. Som of th advnd nonmammalian Synap-
sids (th sintifi trm for th tint group of mammal-lik rptils) hav
jaw hingsthat onsist of a omination of th dntary and th artiular on
th lowr jaw and th quadrat and th squamosal on th uppr jaw (Figur
1B-). This dual artiultion of th jaw, in omination with many othr
fatursof th sklton' indiatsthat thsanimals formd a ridg twn
rptilsand mammals. In th as of th jaw joint, it appars that th dntary
and squamosalns first wr inludd in th jaw joint and thn ompltly
rpladth artiular and quadrat ons as th jaw joint.
on th artiular and quadrt ons wr rdundant in th artiulation
of jaw and skull, thy wr availal for othr purposs. Th vidn from
fossilsand from th mryoni dvlopmnt of th most anint living mam-
mals indiats that ths ons am rdud in siz' altrd in position
and orintation, and onvrtd into th middl-ar ossils, th artiular
() Brsilithrium rptsentsa third linagof Synapsids' found from th lat
Triassithroughth lat rtaous'that had th samdual artiulationas
Phgnels (B) ut rtainda quadratojugai.
(D\ Sinooldonwas a mmr of a fourth Synapsid linag,known only
from th arly Jurassi,tat had a dual artiulation lik tht of Pngeluseven
thoughth individual ons hd diffrntsizsnd shapsthan in Pgnelus.
(\ (Lft) orgnucodo rPrsntsa fifth Synapsid linag,known from th
latTiassithroughth mid-Jurassi,that also had a dul artiulation,ut with
..inus''and no jugl.
a gratlyrdudpartiipationof th
..inus.''Not
To th right lf ah of thsfigursis a dtailddiagramof th
thatth stapdialprossis prsntin (C) and () ut asntin th othrs.This
prossartiulatswith th staps'whih is th third middl ar ossil.
(BI igha Tis is a vntraland latralviw of th lowr jaw of
..Mkl'slmnt,''whih is assiatdwith a
orguodoz,illustrting
srisof onsinludingth totympani,th mallus,and th inus.Ths
onshav a diffrnt mryoni origin from th othr jawons in living
animalsand thy an also larlyrognizdin som fossils.Th
totympaniis a ring of on that suppqrtsth ardrum,whih is th intrfa
twnth outsidworld nd th middl ar ossils.
(F)Thsear adult and mrylnijaws of th duk.billdplatypus'gnus
ornithorhhus.Inth mryo,th kl lmntsar attahdto th
mdialsurfaof th lowr jaw' as in organuodon.Latr in dvlopmntth
etotympani, malleus,nd inus spartfrom th lowr jaw and produth
onfigurtionsnin th middl ar of th dult.
(G) Th lowr drawing is a vntralviw, th upPr on a mdial viw of th
adultjaw of Yooodon,an rly mammal that datsto th arly rtaous.
Th onditionof th kl lmrrts a similar to thoslf an mbryonl
duk-billd platypus.
(H) Th lowr drwing is th mdial viw of th mrylnilowr jaw of th
opossum'lodelphus,whil th uppr drawing is th adult jaw. h
emrynijaw is vry similr to th adult jw lf orguodon and
Yooodo,with th kl lmntsingattahdto th mdil surfaof
thlowr jaw, whil th adult ondition is typial of living mammals' with th
kllmntsingompltlydtahdfrom th lowr jaw.
1 2 0 dpttl

into what is now alld th rallus and th quadrat into what is rrow alld m u s l s a s s o i t dw i
th inus. drivd from what is
Th drivr for this transformation would hav n natural sltiln'sfa- nrvs that srv ths
voring nhandaility to dttvirations. Rsarhsuggststhat th jaw in nrvs.
ths transitionai animals transmittd virations from th ground to th mid. All ths osrvation
dl ar, as it dos in som modrn snaks-think of a small, nlturnalanimal r h sa n d t h t t h m l
that pikd up virations y touhing its jaw to th ground and transmitting mutation irnd sltion
virations from th jaw to th staps.Ths virations would hav travld ponnts. But hw nd
through th artiular (mallus) and quadrat (inus) ons to th staps.In \|977) on dsri
two linags of nonmammalian Synapsids, nithr of whih is anstral ttl nring; a tinkr uss v
mammals, th inus has an outgrowth alld a stapdial pross' whih is a tions, whil an ngin
lvr arm that transmits th for of th sound wavs from th inus to th that adapttion is a hrs
staps. Its prsn in ths two linags stlggststhat oth of thm wr s t u t u r sf o r n w f t l n
volving an nhand paity to transmit sound from th jaw to th staps (ffiint feding) and fl
and thn to th innr ar. into largr, mor ativ
Th arrangmnt of btlns in th linag that inluds mamrnals is diffrnt tant funtion of th ph
than in ths othr linags ut dos inlud n inus with a stapdial pro- a n a s s o i r i o nw i t h f .
ss.Tlr final stagof ours is th sparationof th ossiIsfrom th jaw it- pharyngal pump hav
slf. \/hnth ossils am sparatdfrom th jaw' thy ould transmit los th jws, so thy
sound viations ffiintly from th adrum to th middl ar. Th om- funtion.
plt fossil rord rvls that, rathr than thr ing a dirt transforma- Although th transit
tion from the primitiv synapsid lndition to th mammal ondition, thr drstand, th assoiatt
was a divrsity of arrangmnts lf ns in th jaw-ar rgion and a divrsity trat.Stik your fillg
of rarrangmntsovr tim. Th anatomy f living rptils nd mammals will s tht th jaw j
rprsnts two ndpoints of transformation that followd divrs paths in you an fl th joint r
twn,with most of th variants having gon tlnt. v r t r t s .i n | u d i n g:
Although th fossils Suggstth progrssionof fatursin this transforma- p tlpik up vi
tion, dvlopmntal iology offrs us twl important additional lirrs of vi- and thn to transmit th
dn. First, whn th ossilsfirst ppar in mammalian mryls,thy ar snaks and small mamr
part of th jaw assmly.During dvlopmnt'thy sparatand f<lrmth fa- t i o n s a r n o t u r n |, r n
miliar struturs of th middl r (igur 1F-H). Whil th notion tht an m n so f t h j w s t o l
organism's dvlopmnt rapitulatsits vrlutionaryhistory has n dis- l a r g l ye o n s q u n
rditd in gnral, this is a as in whih dvlopmnt offrs a lu to an th prior rl tht th j
volutionary history that is othrwis traalin th fossils. vironmnt to th innr
Sond, gill arhs, jaws, and ar ossils hav a ritil rlationship to
on nothr that sets thm apart from othr faturs of th skull. Th vr- rple 2: FIorI ul
trat ody is a omposit of thr diffrnt layrs of tissu, th todr- our sond ampl, v
mal (outr), msodrmal (middl), and ndodrmal (innr) layrs. Ths nti analyss ontribu
t h r l a y r s a r a s i l y d i s t i n g u i s h df r o m o n a n o t h r a r l y i n d v l o p m n t t a t i O n s( S h m s k n d
ut om lndd in many parts of th ody as dvltlpmnt progrsss. w fous on shp and
Most of th vrtratskltlnis drivd frtlm th middl, Or msodrml' rs flr thir auty n<
lyr of tissu. Howvr, th ons assoiatdwith tlr gill rhs, jew, and rprodutiv strutur
middl-ar ossils hav a omlnon dvlopmntal origin diffrnt from polln (th mal gam
that of most of th sklton;ths ons ar drivd from spializdlls of mal gamt).
th outr, or todrnral,layr of tissu. Likwis, th musls nd nrvs Th llowrs that insp
assoiatd with ths ons ar rlatd to on anothr in thir origin; th animal pollintors likr
 dpttion 121
h quadratinto what is now alld
m u s l sa s s o i a t dw i t h t h g i l l a r h s ' j a w s , a n d m i d d l - a ro s s i l s r a l l
drivdfrom what is alld te brhiomric sris of musls, and th
rld hv n naturl sletion,sfa.
nrvsthat srv ths musls all dvlop from th sam group of ranial
ns. Rsarhsuggststhat th jaw in
nrvs.
rtionsfrom th ground to th mid-
All thsosrvationssupport th rgumnt tht jaws ar drivd from gill
-think of a small, noturnal animal
rhsand that th middl-ar ossils ar drivd from jaws, and thus tht
j w t o t h g o u n de n d r e n s m i t i n g
mutationnd sltionan indd m<lldompl fatursfrom simpl om-
rsviratiorrswould hav trvld
ponnts.But how and why would suh a transition our? Frangois Jao
lrat (inus)ons ttl th staps.In (1977\ one dsrid volution as a pross of tinkring rathr thn ngi-
Js, nithr of whih is anstral to
nring;a tinkr uss whatvr is at hand and modifis it to srv nw fun-
Lllda stapdialpross,whih is a
tions,whil an nginr dsigns somthing nw from srath. Jao mant
sound-avsfrom th inus to th
that adaptation is a historial pross ausit involvs modifying xisting
s suggststhat blth f thm wr
strutursfor nw funtions. In this as gill rhs gn with on funtion
it sound from th jaw to th sraps (ffiint fding)nd failitatda sond (rspiration).As vrtratsvolvd
into largr, mor ativ animals' rspiration am an inrasingly impor-
that inludsmammals is diffrnt
tant luntion of th pharyngal pump. Th antriormost gill rhs rtaind
lud n inus with a stapdial pro-
n assoiationwith fding. Th ons' musls, and nrvs that pow th
ation of th ossilsfrom th jaw it-
phangalpump hav th sam ation pattrn as thos that opn and
from th jaw, thy ould transmit
losth jaws, s thy wr rdily modifid to spializin on or th othr
drum to th middl ar. Th om-
funtion.
ln thre ing a dirt transform-
Although th transition fr<rmjaws to middl.r ossils is harder to un.
)n to th mammal ondition, thr
drstand,th assoiation twn th jaw joint and haring is asy to illus-
n th jaw.arrgion and a divrsity
trat.Stik your fingr in your r and thn opn and los your mouth. You
ny of living rptils and mammals
will s that th jaw joint is right nt to your outr nd middl ar so tht
Ltionthat followd divrs paths in
yoll an fl th joint mov s you opn and los your mouth. Som living
] gon xtint. vrtrats,inluding small nturnal mammals, taans,and snaks, all
;sionof ftursin this transforma-
apparto pik up virations from th trnal nvironmnt through th jaw
l important additionl lins of vi-
and thn to transmit thm through th jaw t th middl and innr ars. Th
r in mammalin rryos,thy ar
snaksand small mammals that rly on this mhanism for piking up vira-
nnt'thy sparateand form th f-
tions r noturnal and do so through th tip of th jaw. Th ryling of l-
1F_H). \Whilth notion that an
mntsof th jaws to om sound-transmissionons in th middl ar is
volutionaryhistory has n dis.
larglya onsqunof th proximity of th jaw joint to th innr r and
:h dvlopmnt offrs a lu to an
th prior rl that th jw playd in transmitting sound from th trnal n-
lalin th fossils.
vironmntto th innr r.
ls hav a ritial rlationship to
thr fatursof th skull. Th vr-
mple 2: Florl euolution in moke flors (Ivimulus)
rnt layrs of tissu' th todr-
our sndxampI,whil diddly lss swpingin sop, rvalshow g-
ndodrmal (innr) layrs. Ths
ntianalyssontriut to undrstndingth arhitturof ompl adap-
on anothr arly in dvlopmnt
tations(Shmskand Badshaw 19991'Bradshaw and Shmsk2003). Hr
: ody as dvlopmntprogrsss.
w fous n shap and olor of flowrs. Although vryonappriatsflow-
l from th middl, or msodrmal,
rs for thir auty and fragran, from an volutionary prsptiv thy ar
iatd with th gill arhs, jaw' and
rprodutiv struturs that ar spializd to produ and dissminat
llopmntal origin diffrnt from
polln (th mal gamt)and riv and dlivr polln to th ovul (th f-
Lrdrivdfrom spializdlls of
malgamt).
Likwis' th musls and nrvs
Th flowrs that inspir our gratstafftion ar thos spilizd to attrat
:o on anothr in thir origin; th
animai pollinators lik s o hummingirds. Ths flowrs ar atually
122 dpttitl

ompl ominations of haratrsthat togthr ar an adaptation for dis- the flowr. Th flowrs
prsing and ollting polln. Flowrs pollinatd y animals are ompara- produtiv organs onta
tivly larg' olorful, and shapd in ways that failitatth transfr of polln. thy ar nountrd y
Som flowrs ar highly spializd to attrat spifi pollinators, usually y of th insid of th flow
a olor and/or fragran that r partiularly attrativ to a pollinator. Th flowrs that lak th hol
sugary ntar is th rwrd offrd y th flowr to th pollinat,or, a rward y hummingirds, whil
that draws th pollinator's intrstto th insid of th flowr and ststh pol- du larg quillltitis Of
linator in pla to pik up and dposit polln. any flwrs hav mrkings nthrs (ml org:rns)t
that dit th visitor toward th ntar,som of whih a visil only to an- that polln is dpositd.
imals lik insts that s in th ultraviolt (lak-light)frqunis.othrs stamns (fmal ()rgan
hav modifid ptals or spals tht srv as a soft of landing strip on whih t h r nn s r h i r d f d r .
th pollinator an st' nd som spishv spializdstrutursto attah ah flow is wll m:
th polln to th pollinator. diffrns twn tlr
Two losly rlatd spis of monky flowrs, imulus letaisii and . pl alptations for p
rdilis, our along riparian hitats in th wstrn Unitd Stats (Figur 2). math of florvr tl polli
Thy sgrgt y lvation, with . ltuisii ourring at highr lvations, Organs' and ntar pro
and thr is narrow rang of lvation in whih th spis ovrlap. Gnti and sltion ould hv
analyss show that ah spiesis mor losly rlatd to th othr than i- a Ommon nstor. O]
thr is to any othr spis of monky flowr, so any diffrns etwn ussd arlir tht gn
thm volvd aftr thy last shrd a ommon anstor. diffrns rprsntth(
Th two spies hav quit diffrnt flowrs. Th spis at th highr l- hratrs. lf this wr
vations' . lisii, is pollinatd primarily y s. It has pink flowrs that plind vry simply
ar rlativly wid and whos ptals form horizontal landing strip; yllow t th two flowrs; shi
struturson th landing strip appar to guid sto th appropriat part of nario, not srrstantially
mottld to lak. At t
mati diffrnsar a
ah of whih has vry l
or lngth of th anthr
n v y m t i u l o u sa l
th floral traits. Of or
trm vlWs.
O u r k r l o w l d g f p l
p t i n g h y p o t h s s .A l t t
of spals, ptals, st:rm
g n s 'i t i s u n l a rh o w
of th rprodutiv str
dotion rat.
Th rout to th ans
nd thir ollagus y
Figur 2. Studisof th olor, siz,shap,and fatursof th flowrsof two
lular mthods (Sh
loslyrlatdspisof imulus hv illustratdth gntintrol of
2 0 0 3 ) . S h m s k a n d i
rmpladptations.Lft: Te lu flow of . letuisii,whih ftursa
..landing pltform'' usd ,v Mndl whn l
for th inststht transfr polln twnflowrs and
..guids,' tnth ntury. Thy pr
that dirt th inststoward th ntarand th rprodutiv
strutursof th flowr. Right:Th rd flowr ot . rdilis,whih is a long, rids rarly our in n:
narrow' tuular flowr with ntat th as,rahaly th tongusof th ar quit hardy in gr
hummingirdstht srvas th pollinators. lagus nt rlssd h
lza Ad|lttil

sond gnration of so-alld krossd plants with a varil mix of A fundamntal tnt of I
gnsfrom ah original spis.Thy thn turnd to molular iology' m- trs that nhan th fitn
ploying molular markrs tht enald thm to ssoiatptiular varia- wll-ing of spis.I:
tions in floral lraratrsas sn in th vrious orninations of plants with in prtiul:rr for th g<l
diffrnsin molular maks on spifi lotions of hromosoms. Th gon from sintifi dis
diffrnson th hromosoms rfltth loations of gnsthat ontriut nfit to th individu:rl.
to th vition in th flowrs' But altht>ughthis may
Twlv individual haratrswr idntifidarrd studid with this mthod, planatit-rfor all fatu
whih rvald that th truth rvas indd somrvlrr twn th wo x- dividual sltilnoms
trm ids :rout gnti ontrol. A singl grr dtrmind whthr th m i t s ,i n w h i l r n a r l ya l |
flowr was d or pink. By plaing f.lowringhyrids and akrosssin na- It is diffiult to onstu
tur, Shmsk and ollagusfound that th olor diffrnlon had an individu| with :r fitnss.
normous impat on whthr th flowr would visitd y a or a hum- not prolu.
-I.h Th ky to rsolving th
mingird. sintists also found that whil anothr singl gn mad a
larg ontir.rtionto th diffrnsin 9 of th 12 floral traits, thr wr that th ol<lrryworkrs l
many gns tlrt mad smllr ontributions to on or a small sust of th th srifi in individua
traits. rais th offspring oi .l l
From thsrsultsW an offr som plausil ronstrutionsof this om- ommitt-t-lllt'Thr ar
pl adaptation. Th fft of flowr o1or o visitations suggststhat a from th ooprtiv ra
hang in a singl 8n' that fr olor, ould initiat th volution of a sp- providd v oldr offspr
ialized flowr. Diffrnt pollirrators attratd to th diffrnt olors would hirs n <lsrvdin sv
rt diffrnt sltionpressurson th rmainirrgtrzrits.Although it is un- Ths ass :rramp
lar whtlrr th larg sirrgl-gnefft on nin traits prddo followd tiorr. In kin sltilnindi
th hangs in th many othr gns that afftd on or a fw traits' th dution lf rlatd individ
ovrall pitur is of a singl, simpl gnti hang (olor) inititing a stp- omination trf numr
.his
wis pross of modifiation and rfinmnt whos nd produts ar th two th srifi. is a por,
..SlialBh
omplx adptations that w now s.Th math of fatur to irumstan main ssy
..
was not built qrritas asily as hanging th oloration of moths ut did not th alphatial ntris
rqui so mtiulous a molding as to strain duliry' I n f l r t ,k i r r s l t i l l rils
A finl not out th historial natur of irdapttion is that it dos not for th lvl t whih th
sm pssil that th hangs in floral shp and rpodutiv haratrs thr is i1 usal rlations
.!ithout
ould hav prdd th olor hang. variation in olor nd th r- suh as th family lvl,
sultant diffrnin spisof pollinators, thr would hav en no onsis- ilis or lrgr groups f
tnt sltion prssurto mold divrgnt vlus of th othr htrs.But goup sltior-rto our
on th olor diffrnSppard,tlr divrgnt foraging of s and hum_ phistitd thoris, n
mingirds proaly ata|zeda rpid divrgn'This ampl illustratsth to diagntls how oftn t]
ontingny inhrnt in th volution of omplx faturs in whi a singl thn that of th individu
hang in a fatur an promot asad of susqunt hangs that rfin A wid varity of {a
its adaptation to nw irumstans.To rturn to th ritiqu bf Intllignt othr thn that of th rr
Dsign, wihout appriatingth rol of history, w would hrd prssdto rsarh.Bsidsth ml
plain how n{why ths flowrs divrgd so sulrstntially. of grup living arrd
h a v i o r r lS o i o i l o g
..Group sltion'' in th
W H A A N I N D l V | D U A Ls L | o N N o X P L A | N ?
al rol in undrstandin
Our disussion of adaptation hs mphasizdhow th ftursof individr'rals a n d h l s t s( s t h m a i l
ar moidd y th ation of sltion on th avirilal gnti V;rrii1tion. winin diin'' y \

d plants with a varial mix of
l turndto molular iology, m-
hm to assoiatpartiular varia-
rious omintionsof plants with
fi loationsof hromosoms. Th
: ioationsf gnsthat ontriut
ifidand studidwith this mthod'
somwhr twn tlr two -
gl gn dtrmind whethr th
n g h i d s n d k r o s s si n n -
th olor differnalon had an
'ould visitd y a
or a hum-
whil anothr singl gIrmad a
of th 12 florl traits, thr wr
|nsto on or a small suset of th
usilronstrutionsof this om-
lor olr visitatirlns suggsts that a
uld initit th voiutirnf a sp_
td to th diffrnt olors would
mainingtraits. Although it is un-
,n nin traits prdd or fl]lowd
' afftd on o a fw traits, th
i h n n g ( o I o )i n i t i a t i n g s t p -
rt whos rrd pduts irr th two
: math f fatur to irumstan
l oloration of moths ut did not
n dulity.
, of adaptation is that it dos not
hap and rpodutivharatrs
horrt vaiation irr olor nd th r-
thr would hv n no onsis-
,alusof th otlrr haatrs.But
rgntforging of s rrd hum-
gn.This ampl illustratsth
omplx faturs in whih a singl
of susqunthangstht rfin
turn to th ritiqu f Intllignt
;tory' W would hard lrssdto
.d so sustntirrlly.
oN No PLAIN?
d how th ftursof individuls
l t h a v a i l a | g n t i v r i a t i o n .
dpttio I !.\
A fundmntltnt of Dwinian vtllution is that sltion favors hara-
trstht nhanth fitrrssof individuais; it dos not nssarilnhan th
wll-ingof a spis.Indd,th notion thilt volution hs d<lnanything
in prtiul:rrfor th good of th spisis, as fundrntlpinipl, long
gonfrlrnsirrtifidis<lurs; arry nfitto th spisis silfftof th
nfitto th individual.
But ltlrtlr"rgh this may .a usful first prirlipl, it annot th Orrt
planationftlr all Aatursof organisms. Tlr mtlst striking lrzrllngttt ill-
dividual sltitlnoms from th s<lialinsts' ants' Wasps' s, and tr-
mits,in rvhih nlrrly all mmrsof th olony forgo prodution ntirly.
It is diffiult to rOnstruthoW sltioll 11l-t1()tl] individuals wluld far,or n
indivrdulwith a fitnsslf aso|utzro, whih is wht happns if on los
not rprodu.
Th ky t<lrsolvirlgtlris parado fr s'ats, and wsps is to apprit
that th olony w<lrksar raising thir nis and nphws. It appars that
th sarifiin in]ividualfitnss is ltpnsatdy th nhnd ailit to
ristlr offspring f siling, providd that rlstindil.iduls sustain this
ommitmnt.Thr ar many anrpls of lss trllforms of ltruism,
from th ooprtiv raring of offspring y sistrs in lion prids to th id
providdy oldr offspring tl thir pntsirr raising y()ungroffspring, as
has rrosrvlin svrlspisof irds.
Thsssar ampls <lfsltiont th lvl of th family, or kin sel.
tion. In kin sltiorrindividul sirritiis firr,ordrvhrrit nhnsth po-
dutionlf rlatd individtra[s'with th dgrof saifiproportional to th
omintionof numr and rlatdr-rss of th individrralsthat nfit from
thsarifi.This a polr'rfulplanation filr nrirn1'fornrsof soiaIity(sth
is
main ssav ..Soil Bhvior and S<lioiollgy''y Danil Runstin and
th alphatialntris ..Altruism'' and ...W.D. Hmilton'' in this volum).
In fat, kin sltion is or-rlf svrl vaitisof sltion tht ar nnrd
for th lvl at rt,hih tlry our. T'hsvaritis of sltion mrg whn
thris us:rlrlationship twll fatursprssdat a partiuiar lvl,
suhas tlr familv lvl,anl tlr sussof th units ar tht lvl. thv fnr-
ilis or lrgr groups of r-rnrlatdindividuls. Th onditions for kin nd
group sltiont() our hv n xplord xtnsivly in a v:rrityof so.
phistitdthl>ris, nl h ritizrlissu in nlodrn volrrtionaryiology is
to digrroshow <lftnth nditiolls that promot sltion t lvls thr
than tht of th individual irr mt.
A w i d v r i t y t l f f t l r sa p p r l i a l o n l v 1 , s l t i o I - lt l v l s
o t h rt h n t h a t O f t h i r r d i v i d u l ,i r r r dt h i s i s v r y a t i v t o p i o f m o d r n
r s a r hB. s i d st h m a n y a s s o f r p r d u t i v l t r u i s m , t h r r : r s s
o f g r o r ' r pl i v i n g a r - r d o o p a t i v h v i o r ( s t m i n s s a . . S o i l B -
h v i o ri r n d S o i < l i o l o g y '' y D a n i l R u n s t i n r r d t h l p h a t i a l r r t r y
. . G r o u ps l t i o n ' 'i n t h i s v o l u m ) .
S l t i o na t s v r a ll v l s p l a y s i t i -
a l r o l i n r t n d r s t n d i n gt l - r v o l u t i r l r r a r i n t r t i o n s t w l -pl a t h l g n s
a n d h o s t s ( s t l r m a i n s s a y , . v < l l u t i l n a rByi o l o g v o f D i s s a n d D a r -
r v i n i a nM d i i n ' ' y i h a l A n t o l i n n d t h a l p h a t i l n t r y . . H o s t
1 2 6 dDttil

parsit volutiot-t''in this volum); a pathogn tlrat nrultiplis too quikly s that thr ar long
nd kills its lrost tlo quikl for it an b trr-rsmittdto nothr host not at ll, thn rif in
loss out aus sltion among familis Or groups (rprsntdy th drdg dsrid th l<
pathogen popultiorr within an individual host) ovrwlrlms sltion short intvals as ptrnl
a m o n g i n d i v i d u a l s ( p r s n t d y t h s t r u g g l f o r p r o l i f r i r t i o nw i t h i n n ssoition twn pu
individual host). th trm ptltuted q
Gotrld rgrrd that ot
with nturl sltiona
AR sUDls oF MlRoVoLUloN RLEVAN o
on th prmis that if s
U N D R s A N D IN G M A R o v o L U | o N A R Y P A R N s ?
a prd<rminantPattrn
[t is onvnint to hartrizth produts of evolution as ithr mlrovo- haratrs suggsts rs
lution or marovrlutirln.iroeuoLutioll rfrs ttl small-sirlhangs that suggstdto thm that l
ar dirtly osval, lik many o{ thos distrssdlir (mlanism irr a susqunt puls lf a
moths, disas rsistanin atri, or instiidrsistan).As w hav Although Gould nd
disussd, mirovolution has n vry thorouglrly dourntd and has that nsud aftr thir {
n studid irr ways tht mt any tst of sintifirigi<lr.As rsltlt,it is nr First, neturl sltion
ontrovrsial,vn among n-ranyfaith-asdritis of volution (sth main isms to small-salhar
ssay ..Amian Antivolutilnism: Rtospt and Prospt,' y ugni tlran natural sletion
Sottin this vlum). qunt assoiation tw
crleuollttio rfers to th Iargr-salvnts tl,ratw attriut to volu- suggststo Gould and
tion, sulr as tlr origin of nw spis,th diffrntialprtllifrarion or diffr- phnomna, ut thy h
ntial tintion of rtain typs of spis, and th mrgn of highr d u r i n g t h 3 6 - y a rh i s t
lvls of th txonomi hirahy.This is th aspt of volution that is on- Th ountrargrrm
rovrsialusit rprst-tts vnts that otrr on a timSllltht is -ruh hs thr parts. First, v
longr than our livs :lnd hn not dirtly osrval. As w hav dis- tion y ntural sltr
ussd, sintistsmust turn to indirt mthods to draw infrensaout th thn th rats i]ssolat
undlying mhanismsthat aus marvolution. G16pags finhs an
Th most rvolutionary ftur of Darwin's th rigi of Spcies i n a p l p u l t i o ni n s i r
(1859) was to popos naturl sltion as th singl unif1'ing mhanisrn fall. In dought yrrs
tht auss oth miro- and marovolutiln.Drwin argud that mrovo. sdswith thik sl<
lution is just mirovolution writ larg, or that th possW s nd study ftiv at harvsting su
as th aus of mirovolution will, givn suffiit.lttitn, lso aus vry- Baus tlrr is a gn(
thing that w attriut to marovlution. H ergud thi.rtnrural sltion, finhs xhiits largr,
whih alrssth volution of th ad:rptationsdisussdthroughout this s- to put this rat of vol
say, is also rsponsilfor th origin of :r1llvls of biologial orrrplityand sil rrd' it is l0,000
for th origin of iologial divrsity, or all th spisthat hav n found tprt as puntuation
on th arth thoughout its history. inonsistnt witlr tlr r
otrr inaility to osrvmrovoltltionin tion has inspid ontrovrsy tion' [f anything,w r
ovr whthr ntural sltionrally is th unifying mlrnism of all volu- fossil rord.
tionry hang. StphnJ. Gould and Nils ldrdg ar th uthors of wht S o n d , s l t i o l li s
is urenly th most prominent sintifirgulnnt against ntural slti<ln This, of ottrs, is tlr.
as a singl, unifying aus of miro- and nrrovolution (ldrdg and ut hr it is importan
Gould 1972; Gou\d and ldrdge 1977). Ths authors Wr motivatd y tion an onl mak fi
th pttrrrof hang rordd in th fossil rord. If w look t th fossil his. tn to hang, mltst
tory of th volution of most orgnisms, suh s th inrasof ody siz in Darwin's lins of argu
horss or rain siz in th hominid linag lading to our spis,w o[tr-l t that domsti dtlg

1 o g nt h a t m u l t i p l i s t o o q u i k l y
r transrittdto nothr host
]s or groups (rprsntdby th
ual host) ovrwhlms sltion
uggl for proliftilnwithin an
oN RLVAN o
|oNARY PARNs?
s of volutionas ithr miroevo-
rfrsto smirll-salhangs that
; disussdrlir (mlanism in
.stiidrsistan).As w hav
:horoughlydoumntd and has
:intifirigor' As rslt,it is not
l ritisof evolution (sth rnain
spt and rospt'' y ugni
vntsthat w ttriutto volu-
liffntialp<llifrzrtion
or diff-
s, and th mrgrrof highr
l asptof volutilnthat is on-
)ur on a timsalthat is rnuh
:tly osrval.As w hv dis-
Lodsto drw infrnsaout th
lr-rtion.
win's o th origil.t of Spies
s th singl unifying nrhrrisnr
r. Darwin argud that marovo-
hat th prossW s nd study
sLrffiinttir, also aus vry-
H argud that ntural sltion,
lns disussdthrouglrout this s-
vls of iologialornplity nd
th spisthat hav n found
n tion has inspirdontrovrsy
nifyingmhanismof all volu-
Idrdgar th authos of what
gumntagainstnatural sltion
marovolution (ldrdg nd
hs autlrorsWr motivatd y
ord.If w look at th fossil his-
:h as th inrasof ody siz in
ladingto Our spis,w oftn
dpttkl 127
sthat thr ar long intvals whn th tait in qustion hangs littl or
not at all' thn rif intrvls wlrn th tr:lit volvs rapidlv. Glr-rldd l-
drdgdsridth long, stal intrvals as stasis or quilirium and th
short intrr.irisas puntutions. Thv also pointd out that tlrr is oftn an
assoiatirrtwnpuntations :rnd th lrigin of nw spis and oind
th trm punctuted equilibrium to dsri th ntir pattrn. ldrdg and
Gould agud that oth phass of pr'rntuatdquilirium ar inons,istnt
rvithnaturl sltionas a singl urrdrlyingaus.hir argtlmnt,s asd
on th prmisthat if sltionwr suh unifying aus' w should osrv
a prdominrltPttnof graduI.otrtinuoushang in hatrs.Stsis in
haratrssuggstsrsistanto hang. Th disontinuity of puntuations
suggstd to thm that somthinghappndto rak this rsistanand aus
susquntpuls lf alatdvolutiorr.
Althoug Gould nd lddg'sargumntshv volvd during th dt
rhatnsudeftr thir first .ssay,thir argumntshav two unifying faturs.
First,natralsltionis plsitds ing fftivonly in fir-r_tuning organ-
is-rsto smirll-salhangsin thir rrvironmnt.Snd' som pross othr
thn naturirl sltion must om into pla to aus puntlliltions. Th fr_
qunt assoition twrrpuntuirtionsarrd th apparan of nw spis
suggststo Guld and ldrdg tht this pross' whtvr it is' auss oth
plrnomna,ut rhy lrav ltrd tlri prtlpsd mhnism tor this pross
during th 36-yar history of puntuatd quilirium as an ida.
Th ountrargumntin favor of natural sltirnas a unifying Pross
has thrpars.First, w hv mpirial stimatslf h r:rtat whih voiu-
tion y natural sltionn our' and w know that it arr muh fastr
thn th rats assoilltdwith puntlltions irr th fossil rord' For ampl,
Gal6pagosfinhsan volv a |0" hang in avragody siz or ill siz
in a populartionin a singl var in rsponsto ithr droughts or hvy rain-
fall. In drought ylrrs ftlod avilaility is low' th ids mrrst rly orr larg
sdswith thik sdoats and individuals rvith wid, havy ills ar mor f-
ftivat harvstingsuh sdsand ar mor likly to surviv and rprodu.
Bausrhis ir grrtisis t<lill siz :rnd shap' tlr nt gnrtion of
finhslriitslrgr,hvir ills. This is lassi mirovolution. If w wr
t put this at of volution on th sam timsal as what we s in th fos-
sil rord,it is 10'000 ()r mor tirsfastr than wlrt Gould and ldrdg irr-
trprtas puntuations.Th apparent suddnnssof puntuations is thus not
inonsistntwitlr tlr rat of volutilnthr is attainlrlund naturl sl_
tion. If anything' w migl-rtask wlry volr-rtirn appars to so slw in th
fossilrod.
Sond,slrion is apal lf using sustnilhngs in orgltrlisms.
his, of ours' is th fundamntal hallngw rviwd irr arlir stions'
ut hr it is inrpotant ausGould and ldrdg rgu tat ntural sl-
tion an only nr.rkfin rdjustnrnts to orgnisnrs.and tlrat stasis' tlr sis-
tilnto hang,nrust rokn y som othr pross.'Wan follow on <l1
Drwin's lins of ar]umnt'alit lvith mor informtion. Gnti dta indi-
ttht lomstidogs wr drivd from wolvs, ginning tns of tlrlusands
128 dpttio

of yars ago. Th fist apparanof morphologially distint domsti dogs th ontr<lvrsyrmalns t

dats to around 10,000 yrs gO (Savolainnt al. 2002). rom suh rnt shot trm with th nor
ginningsw hav rdsthat rang in siz from th hihuahua to th mas- sk to xplain, lrnd th t
tiff nd in shap from th Pkins to th lislr wolfhound. If w fous on an uSttld frtlntir rft}
ody wight alon' th rang of avragwights mong rds is fom around
1 kilogram for hihuahuas to 80 kilograms for mastiffs. This rang ds
th ntir rang of ody sizs f othr spisin th family anida' whih Horizons in th Stu
inluds dogs, wllvs, fos, and othr doglik arnivors (Finarlli and
Flynn 2006). It talls slrot of wh:rt w s in th ordr anivora, whih in- Adprirtion is ing sttrd
luds th family anida along with othrs' suh as th Flida (ats)or Ur- of adaptation is vidnt,
sida (ars). Tn thousand yas of artifiial sltion hav thus n r ginniIrgto dissttl
.!
suffiint to gnrat a rang lf morphologial variation that is quivalnt to daptations. hav f
th rang w Swlrn w lok thr or fottr stps up th tironomi hirar- ut rr. llnot onlr'lt1
hy (spis'gnus' family, ordr, lass' phylunr, kingdom). found ar gnrlll. Gn()
Third, w must oIlsid whthr th prsnof stsis' or long intrvals s r l r , n d e v s i . l t.ll f t
lf littl or no hang, an Lrplaind y natural sltion.Th Gal6pagos v r y l if f r n t ( ) n | u 5 i O
finhs again provid a good lnptual altrnativ to plain why thr Th <lnstrairrts on th
might b long intrv:rls of no apparnt hang in th fossil rord. In irn ways. In slm:ls
drought yars thr is intns sltion for inrasd ill and ody siz. In tatiOtls. In othrs th'vaI
th iny yars that aompnv l Nino vnts' th rsour s hngs in ntur lo sl lrirusl
aus thr is an undan of small, thin-shlld seds from diffrnt individual gnsth21tPro
spis of plants. Sltion now favors smllr individuals with narrow, that tlr ar inl,rritdar
mor forps-lik ills. This mans that priods of sltion for inrasd p | u s i l r ( )p o s i t t n s
ill siz altrnat with priods of sltion for drasd ill siz, and in in ft th:ra frvstu
..vollttt
som yars th is in fat nl sltion at all (Gibs nd Grzrnt 1987). ovr th ririn ssay
th narly thr dads in whih ths irds hv n studid, thr has rah harlswoth and th
n no nt hang in th ill siz of th populaion in spit of strong vi_ suggststht th !]n()m
dn for intnse sltion and rapid volution on a yar-to.yr asis (s not mrly a oDstralllt.
.!inr
1994 for nonthnil rviw of studis of Drwin's finlrs).Th important qustion thilt
fossil rord ould rprsrrta frgmntd,long-trm history of suh short- our olltiv vir'vo1
tm flutuating sltion. Th fossil rord dos nrt rpsnt ah nd has hangd rnakl11 it
vry yar' and, in fat, a dfinl priod in th fossil rord will rprsnt a portnt for undrstnd
random rnitur of ths diffrnt pisods of sltion.T atual rt of this air is lding to [
volution that w s in th fossil rord is a long-trm avrag of th in- th vllutitln <lfsoi:rl
rass and drass and will invitaly undrstimat th true rt at l:rrg phr-rt>mntl sUh 1
whih orgnisms an volv. A fssil rord of th Gal6pags finhs may th volr-rtiln f th ll
w | |l o o k l i k s t a s i s . tiv ras of rsah l
In tlr final nalysis thr is nothing in th fossil rord that inhrntlyon. rvlllttion hs rrald
tradits Drwin's daring ida that natural sltion is th unifying mh- n W ( ) n st o a s k d . .
nism. Organisms volve muh mor apidly tlran tlry appar to volv and tlr diffirrlty lf a
during puntuations,and th apaity of organismsto hang undr sltion tht hilppnd ovr a l
far xds th onstraints that ar apparnt during stasis. Th asrof otl-rrttlpis, ut irr a dr
hang during stasis lay instad a statistialartifat of flrttuatingvolu- o u r l p r i e t i < l on t n t
tion tht traks a flutuating nvironnlnt. But our argrrmntsdo not prov vrr, diff-rntthan th
that marovolution is mirovolution writ larg;thy mrly support th vi- h s t t r d y l f D r ' i n
aility of naturl sltion as a singl, ovri1rhingplnation. Th rot of ftr (Jll Ib rigitt ,

lhologiallydistirrtdomsti dogs
nn t al. 2002). From suh rent
ze from th hihuhua to th mas-
Irish wolfhound. If w fous on
lightsamong rdsis from around
ns for mstiffs. This rang xds
:isin th family anida, whih
doglik rtrivors(Finarlli and
in th order Cnivora, whih in-
s,suh s th Flida (ats)or Ur-
tifiial sltion hav thus n
4ialva;:itiontht is quivalnt to
)ur stpsup th taonomi hirar_
rylum,kingdom).
rsnof stasis,or lorrg intrvais
ntural sltion.Th Gal6pagos
ltnativto plain why thr
hang in th fossil rord. In
'r inrasdbill and ody siz. In
vnts'tlr rsour as hangs
thin-shlldsds from diffrent
nllr individuls with narrowr'
riods of sltion for inrasd
ln for drasdill siz, and in
l l l ( G i b b s n d G r a n t 1 9 8 7 ) .o v r
irds hav n studid, thr has
p o p u l t i o ni n s p i t o f s t r n g v i -
ution on a yar-to-yarasis (se
studisof Darwin's finhs).Th
, long-trmhistory of suh short-
lrd dos not rprsnt ah and
r th fossil ord will rpsnra
:s of sltion.Th atual rat of
is long-tm vagof th in.
. undrstimat th tru rat at
rd of th Galdpagos finhs may
: fossil rodthat inhrntlyon-
sltion is th rrnifyirrg mha-
Lpidlythan thy appar to volv
anismsto hangundr sltion
rnt during stasis.h asn of
ltial artifat of flutuating volu-
But our rgumntsdo not prov
larg;thy mrly Support th vi-
rhing xplntion.Th root of
d7lttil l2L)
thontrovsyrrnainstlr difIiulty of roniling wlrat W an study in th
shorttrm rvith th normously liffrnttirsalof th miri<lrpattrns W
rmit-ts
skto plin, r-rdth ]ividtwn miro. d nl:lrlr,oir'rtion
an ullsttl]
frontir of th disiplin of volutionary ioltlgy.
Hrizonsin th Study of Adaptation
Adapttion is ing stulidt last as intnsly as vr. Althtlugh th rality
of adaptatiolris ','idnt,trrtrn1, fi1g15 of it rrnain unlr. Fo itt1ll,rr,
arginnirrgto disstth gntiasis of dptatins' ;lartiularlv orrrpl
adaptatins../ hlrr, f.lassi studis lik tlros ()n th mnky flwrs,
ut w annot lr-rlud whthr th pi-lttrnsthat Slrrnsknd ollagr'rs
found ar gnral.Gnomi and protomi nrtl-rods ar alatingthis r_
sarh,and a vrsion of tlris sslr1. wittt-ra dd frttltnorr,rrrylffsorrr
vrv diffrnttlnlusilnsaout this tlpi.
Tlr tlnstrints on th raw mtril f<ladptlrtion ing sttrdidin dif_
frntways. In som ass iologists a studying th pr<lprtisof nw mu'
ttiOns.Ir-rothrs tlry ar xarir-ring wlrthr hirratstht tlu ttlgthr
in tttur1lso usthry ar <lntroIldy th san1gl1stlr usth
individualgnsthat produ onrtdinhritn:rrlignd irr suh wav
tht thl' r inhritd s units lr lrloks lf gns.Although it nla1'stlriln-
plausiltl posit tnsivlinkag of gnsthat ontrol orrlatdharetrs'
in fat thr.ar frv studisthat show a surprising lvl of suh linkags (s
th mzrinssay..vllutionof th Gnorrr''y Brian lrirlsworthrrd Do-
rah hrlsworth lrnd th alphatialrrtry..Minriry'' in this volurn).This
stlggststlrt th gnonl itslf irn prodtrt of Dilwirrilln volr.ltitlnatl]
nt mrly a onstraint aginst whih suh volution -rustoprat.Tl.risis :rn
important 1r'rstion that hs ttratd mnv sil-ltists.
our olltivviw lf sltilrrat lvls othr than thos lf tlr individr.rl
h a s h a n g m
l a r k d l y i n t h p a s t 0 y a r s . u l t i l v l s l t i l ni s l a r l y i m .
potantfo undrstandirrgmrr1'lldptirtions i divrs systlls;rsrh itr
this ara is lading to fa mor nuand and sophistiatdintrprtati()ns()f
th volrrti<lrr of soilitr,,rlf pllthogn-lrostrltionships,ntl vn of r.},
larg phnornirsuh as multillularity and phaps th ll itslf. Indd,
th volution <lfth ll rld th ]ivrsitl.of ll struttlrsr trmly i.t-
t i v a r a s o f r s r h( \ / t l s20 0 4 ; K u r l n d t a l . 2 0 0 6 ) . T h n r < l l u l . r r
rvtllutionhas rlirldlong-standing qtlstionsto lrnswrd nd ntirly
nw ons to rskd.lr spi:rlizdrrturlf th nrthtldsirrgrrrploy1
and th diffiulty of app|ying lultilvl sltion thory ttl phntnnon
that happnd lv a illilll y2lrsag() tllak poerss slowr thn it is for
othrtopis' ut in dild Or two our knowldg of llr.rlarvlution ni
our appriationof multilvl sltiontrsa ritiill vtlllttionvfor rvill
vr diffrnt tlrilrrthy ilr tod.ly.
Th study lf Drwinin volutiorl rm:rins virnt topi narly 150 yrs
aftr n tha rigitt tlf Sp,isllrrd.AlthtlLrgh Dru,it-tlil out th
10 Adpttil

argumnt in road outlin, it fll to his sussors to disovr whih lmnts aFddn, B. J. |992. l"rl
the Fmil quidc. N
of his argumnt Wr possil in thory, whih wr dmonstral in pra-
N{Nair,. R. 1987.Ha
ti, and whih wr roust to varitions among spis and irumstans.
sstm' rtJs ilt tll
T'hat so muh of his original rgumnt mains standing is a triut to his g- tsurnura, I]', and G. Vss
nius; that so muh rmains to disovrd is a triut to th mrvls that i n h r r , r ' t l - s p t t sJp i t
adaptation offrs and th divrsity of natur th:rt it has atzrlyzd. tomtllog 57: 9| |-9
N,lPk,. ,\. l99.5. or1
damslflis rf tw<ltltlt
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Rznik,D., nd.|. T.r,is.1996. rnpiri:rlstudisf irdpttion.In ' Ros nd
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 P U r p Ibe t e i

Molular volution ynobteri

FrnisoJ. Al Flavobatria

Thermotog
|es

Figur 1. Th univrsaltr
Molular biology hs mad it possil to ronstrutth ontinuity of su- Batria,Arhaa, and uk
ssion from th original form of lif, anstral to all living organlsms' to oganismsar rlatdy o
vry spisnow living on arth. Th univrsal tr of life, whih mrs y th tr's..trurrk''(thst
using slw-volvingrilso
all known sorts of organisms, inluds thr sts of ranhs' Batia, Ar.
haa' and ukarya' whih all mrgd from on form of lif, LUA (Last
Univrsal ommon Anstor; Figur l). Al1 living organisms ar rlatd y
This has mad it possil l
ommon dsnt from a singl form of lif, rprsrrtdy th tr trunk in
ously unknown and to or
th figur. Lif may hav originatd mor than on on our plant-w
Th prision with whih l
prsntly do not know-ut only on form of lif prosprd and lft dsn-
vidn from molular b
dants, th organisms that nw populat th arth.
h molular <rmpo
Batria and Arhaa ar prokaryoti mirosopi lrganisms.ukarya ar
natur of th omponnts
organisms with omplx lls that ontin svral orgnlls,on of whih'
n d u s d .I n a I | a t r i , r
th nulus, inluds th hrditry DNA. Most ukaryoti organisms r
sists of a diffnt squn
singl lld and mirosopi. Animals, plants, and fungi ar multillular or-
sntd as A' , G, and T; Fl
ganisms, thr of th many rnhsof th ukary st.
ontaind in th DNA of t
h main ranhs of th univrsl tr of lif from LUA, svral illion
th sam in all organisms
yars ago' to th prsnt hav n ronstrutdon th whol and in many
sorts of organisms ar syntl
dtails. Mor dtails out mor and mor ranhsar pulishd in sors
varial lngth, of th sam
of sintifiartils vry month. Th virtually unlimitd volutionary infor.
a l h u n d r d o r h a m i n i
mation nodd in th DNA squnof living organisms allows volution-
of biohmial rations-
ists to ronstrutany volutionary rlationshipsthat hav ld to prsnt-day
nrgy and to mak up th
organisms, or among thm, with as muh dtail as wntd. Invst th ns-
This unity rvals th g
sary rsous(tim and loratory pnss),and you an hv th answr
ganisms. Thr is no oth
to any qury with as muh prision as you want.
formity whn numrous
likly. Th gnti od st
thr nulotids in th u
olular volution: Uniformity and Divrsity atly th sam amino z
t h a n i t i s f o r | a n g u g t t
In its unviling of th natur of DNA and th workings lf organisms at th snt patiu|a objr.
lvl of nzyms and othr protin moluls, molular iology hs shown plnt, tr' woman-ar(
th unity of lif. orovr, molular iology has shown that ths mol- diffrnt oks, on an b
uls, DNA and protins, hold informtion aout an organism's anstry. of ommon origin.
t.)z
 olulr uolutio l

ARlA ARHAA UKARYA

Euryrhaeot
G nn o n s u I f u r ^/|thanosarina Anim|s
bteri N4eth0bteri
Um
Fungi
Purp
btri Gmpositives Mthanoous
S l i m em o l d s
.lr
utl0n
P|nts
Crenrhaeot ntamoba
(ynobatri hermoproteus iIiate5

Pyoditium
F|g||ates
Iavobatria
rihomonads
Thrmotg|s |\4irosporidi

DipIomonds

Figure1. Th univrsaltr of lif. h thrmajor groups of organisms,

to ronstrutth ontinuity tlf su-
n s t r a tl o a I I I i v i n g o r g a n i s m s ,r o
tnivrsaltre of lif, whih mras
thr StSof ranhs,Batia, Ar-
from on frrn lf lif, LUA (Last
. All living organisms ar rlatd y
lif, rprsntdy th tr trunk in
nor than on on our plant-w
lrm of lif prosprdand lft dsn-
th arth.
mirosopiorganisms.ukarya ar
ain svralorganlls,on of whih,
tJA. ost ukaryoti organisms ar
llants, and fungi ar multillulr or-
th ukarya st.
of lif from LUCA, svral illion
'nstrutdon th whol and in many
or ranhsar pulishd in sors
rtually unlimitd volutionary infor.
lf living organismsallows volution-
'ionshipsthat hav ld to presnt-day
:h dtail as wantd. Invst th ns-
nss)'and you an hav th nswr
you want.
'y and Divrsity
nd th wrkings of organisms at th
uls'molular iology has shown
iology has shown that ths mol-
ation aout an rganism's anstry.
Batria,
Arhaa,and ukarya' ar rprsntdy th thrmin ranhs.All
organismsar rlatd y ommon dsntfrom a singl frm of lif, rprsntd
by thtr's..trunk''(thstraight.uplin t ottom).This tr was onstrutd
.Wos
usingslow-volving riosomalRNA gns.(Adaptdfrom 1998.)
This has mad it possil to ronstrutvolutionary vnts that wr prvi-
ously unknown and to onfirm and adjust th viw of vnts alrady known.
h prisionwith whih thsvntsan ronstrutdis on rason th
vidnfrom molular biology is so omplling.
Th molular omponnts of organisms ar rmarkaly uniform in th
naturof th omponnts' as wll as in th ways in whih thy ar assmld
and usd.In all atria' arhaa' plants, animals, nd humans' th DNA on-
sistsof a diffrnt squn of th sam four omponnt nulotids (rpr-
sntds A, ' G, and T; Figur 2). Th gntiod, y whih th information
ontaindin th DNA of th ll nulus is passd on to protins' is virtually
th sam in all organisms (Figur 3). Th normously divs protins in all
sortsof organisms ar synthsizdfrom diffrnt ominations, in squnsof
variallngth, of th sam 20 amino aids (listd in Figur ), although sv-
ralhundrdothr amino aids xist. Similar mtoli pathways-squns
of iohmialrations-a usd y th most divrs organisms to produ
nrgyand to mak up th ll omponnts.
This unity rvals th gnti ontinuity and ommon anstry of all or-
ganisms.Thr is no othr rational Way to aount for thir molular uni-
formity whn numous altrnativ struturs ar in prinipl qually
likly.Th gnti od srvs as an xampl. ah partiular squn of
thrnulotidsin th nular DNA ats as a pattn for th prodution of
xatlyth sam amino aid in all organisms. This is no mo nssary
than it is for a languag to rrsa partiular omintion of lttrs to rpr-
sent a partiular ojet. If it is found that rtin squns of lttrs-
plant,tr' Woman-ar usd with idntil manings in a numr of
diffrntooks, on an sur that th lnguags usd in thos ooks ar
of ommon origin.
11 ollrF,uolutil

.''io
8roov
*j U
Ph
UU'l U

c
U
*^.l Leu
U

UUGJ U
o
o

! l phogPht str
t*r
U |
o *^l
id N ln ss

I Leu
L

*"1
AUUl

y)

AU l
I lle

I
^u^
AUG et

GUUI tl

GU l
G I val
GUA
I
.u"J
igur 2. Th doul.stranddhlialnfigurationof th DNA molul.Th
Figur 3. The gntiod:.
molulonsistslf two polynultidhains.Th outward akonof th
mssngrRNA and th am
molulis mad tlf altrnatingdoyriossugrs(S)and phosphtgroups
nitrogn as thymin dos
(P).Nitrlgnassonntdto th sugrsprojttowrd th ntrof th
th othr thr nitrogn as
molul.Th two nulotidhains r hld togthry hydrognonds
adnin (A)' ytosine (), aI
btwnomplmntirrypurin-pyrimidinss.Adnin (A) nd thymin (T)
protins (with th thr-lt
flrmtwo hydrognonds;ytosin() and guanin(G) form thr.
(Ala, A), arginin (Arg, R), '
(ys, ), glyin (Gly' G), g
( H i s , H ) ' i s o l u i n ( I l ,I ) ,l
)' phnylalanin(Ph,F),
tyrosin (Tyr' Y), tryptoph
Gns nd protins ar long moluls that ontin information in th s-
qun of thir olponnts in muh th sam way in whih sntnsof th
nglish languag ntain information in th squn of thir lttrs nd
words. Th squnstht mak up th gnsar passd on from parnts to th rational for th ror
offspring and ar idntial from gnration to gnrtion' xpt for o. nd outomswiIl rvi
sionl hangs introdud y mutations. losly rlatd spies hav vry As an illustation, lt ul
similar DNA squns;th fw diffrnsrflt mutations tht ourrd ooks are 2oo pags long
sin thir last lmmon nstor. Spisthat ar lss losly rlatd to on amination rvals that t
anothr hiit mor diffrnsin thir DNA than thos mor losly r- for word, xt that an r
latd ausmor tim has lapsd sin thir last mmln nstor.This is two ooks annot hav l
 olulr uolutil 1''

_^..-jhl,rnt Sond Psitio

*l
U A G
= UUl uu1 U

-".::&
UGUl
Phe
,,-- | I Tyr | y,
UU l ULL
I
UAl UG J
l&'
....c-
uu^l U UAA Stop UGA StoP
lu |

*ur
.".1
UUC J UG- UAG Stap UGG p G
'*-l
^,,
-" t*-l U
Fl
U | |
t
J

*^l l Lu l Po
*l
:::J
-r..4, A
l
J

Gln
- UGJ Gl Gl G
-l-
^Ul *ul ^"u-l

tW
AUU.l U
;i
U
I lle A | AAJ
Asn
AG J
Ser

|
l
^u^ ^.^
|
t^'
*".l Lys
Ac"l
Arg
A

UG et AGJ CJ AGGJ

"uur "*r GAUl ccul U

G
GU l
cuA I val "tt I ^"
G
GAJ

GAAI
l 'p
""'
GGI",,

A
I | I clu l
P.ent GUG- GG- GJ G6GJ G

3urtionof th DNA molrrl.Th Figur3. Th gntiod:orrspondentwnth 64 possilodons in

rins.Tlr outwardakbonf th
sugars(S)and phoshatgtlps
projttowrdth trof th
ld togthrby hydrognonds
ss.Adnin(A) and thvmin (T)
l guanin(G) form hr.
tht ontain information in th s_
samWay in whih sntnsof th
l th squnof thir lttrs and
]nsar passd on from parnts to
ion to gnration, xpt for oa-
. losly rlatd spis hv vry
rsrflt mutations tht or.rrd
i that ar lssloslv rlat{to o1-l
DNA than thos mo losly r.
: thir last ommon anstor.This is
messngr RNA nd th amino aids (r trmintionsignals,..stop'').Th
nitrogenasthymindos not ist in RNA, whr uril (U) tkes its pla;
hothrthnitrogen assin mssngrRNA ar th sm s in DNA:
adnin(A), ytosine(), and guanin(G). Th 20 amino aids that mak up
protins(with th thr-lttrand on-lttrstandard arviations)ar alanin
(Ala,A), arginin(Arg, R), asparagine(Asn, N), aspartiaid (Asp, D), ystin
(ys,), glyin(Gly, G), glutamiaid (Glu, )' glutamin(Gln. Q), histidin
(His,H), isoluin(Il,I), luin(Lu, [-)'lysin (Lys' K), mthionin(t'
), phnylalanin (Ph,F)' prlin (Pro, P), srin(Sr'S), thonin(Thr' ),
ryrosin(yr, Y), tryptophn(Trp' W), nd vzrlin(Val' V).
th rtional fo th ronstrution of volutionary history, whos mthods
and outoms will rviwd latr.
As an illustrion,lt us assum that w ar omparing two ooks. Both
ooksar 200 pags lorrg and ontain tlr sam num of haptrs. Closr
aminationrvais tht th two ooks ar idntial pag for pag and word
for word, xpt tht an oasional word-say, 1 in 100-is diffrnt. Th
two ooks annot hav n writtn indpndntly: ithr on has bn
16 N\llulrtlolt.ttion

opid fror th othr, lr <lthhav n opid, ditly or indirtly, from stalishd whn th squt
th sam oiginal ook. ln living ings, if ah omPoilnt nulotid of diffrnt organisms. Th s
DNA is rprsntdy on lttr,th omplt sqtlnof nr-rlotids in th vn twn vy diffrr
DNA of a lrighr organisnr ,,orrldrquir svrl lrudrd llks,ah with partiv anatomy' for
.Whn
hundrds of pags, witl.r svral tlrlusnd lttrs tln irh pg. th pin trs, and humn i
..pags''(or squnsof nulotids)in ths ..oks'' (gnoms)ar ant-
protin squns that ar
ind on on, th orrspondnin th..lttrs'' (nulotids)givs un- multipliity. ah oganis
rnistakalvidnof ot-nlrronorigirr. all rflt th sam evoluti
olulr biology offrs two kinds <lfargumnts flr voltrti<ln, irsdon gne or protin does not si
two diffrnt grounds. Using th alphat arralogy,th first rgumnt sa}'s a st of spis, addition
that languagsthat us th sam alphat (th sam hrditry molul,th mattr has n sttld to
DNA md r"rpof th snrfour nultltids,and tlr sanr20 atlrino aids in oov, th widly d
thi protins),as wll s th sam litionary(tlrsm glltiod),arrrrot opn up th opportunity
of indpndntorigin. Th sond argumnt ()nrllssimilarity in th s- nt dgrs of rsolution il
qun of nulotids in th DNA (and thus th squnof amino aids i volving gnS to reonst
th protins);it says tht ooks with vry similar tts annot of indpn- fastr-volvin$ $ns to r
dnt lrigirr. divrgd oganisms.
Gns that nod th l
volving gnS.Thy hav
Informationalaromoluls ships among groups of or
among tria, rha,;
Nuli aids and prorins hav n a||ed i|'ormtiol romoltlls ing world), whih divrg
as thy ar long linar moluls mad up tlf squns of units- among th mirosopi pl
nulotidsin th as of nuli aids, amino ids in th s lf prltins_ ompard with plants and
tirat rtin oirsidr.rlmoLlnts of volutionary itrflrmirtion.omparisor-l of orgnisms that divg
of th sqn of th omponnts in two momoluls stalishsth tiond alir, whih volv
numrs that ar diffrnt.Bausvolution usually ous y hanging on gns' is usd to diphr l
unit at a til' th numr of diffrr-rs is an ir-rdiationof tlr rny of suh as among humans, fi
omon nsty. th fiinopptids involv
Th dgr of similarity in th squnof nlotidsor tlf amino ids ing th volution of los
an prisly quantifid. For ampl, in humans and lripanzsth maaqus' himpanzs,a
protin molul alld ytohrom_.whih srvs vitl funtion in rspi-
r a t i o n w i t h i n l l s ' o n s i s t so f t h s a m ] 0 4 m i n o i d s i n a t l vt h s a m
ordr. It liffrs, horvvr, frm th ytohro-t_of rlrsr.rsrnonk1.s y olular Phylogni
1 amino aid, from that of horss y 11 additinl amino aids, and from
that of tuna y 21 additional amino lrils. DNA and protins providt
Th dgrof similarit rfltsth nylf ommon nsty.Thus in- ags from ommon nCeS
frnsfrom omp.riltiventrr'nd othr lisiplinslnrtlir-rg vllu. nti hang that hs o
tionary history an tstd in molular studis of DNA nd protins by sm at first that quantifyi
xamining thir squnsof nulrltidsand amin aids. T authority of impossiI ausit w
this kind of test is ovrwlrlming:irl-rof th thor"rsands of gns irnd thorr- isms that livd in th prst
snds of protins ontind in an orgnism pvids an indpndrrttst of past ar somtimes prs
that organism's volutionary histor1'. largly disintgratd. Nev
Molular volutionrystudishav thr notal advntagsovompar- vid informatin about a
iltiv anatoly and th tltlrrlassial lisiplins.on is that th informtion As a spifi ampl'
is mor radily quntifial.Th nr-rrrrr of Lrnitstht ar diftrlltis rdil hrom-, whih is involv

opid' dirtly or indirtly, from
,, if ah omponnt nulotid of
rpltsqunof nr-llotids in th
svralhundrd ooks, ah with
.Whn
rd lttrs on h pag. th
. . o o k s ' (' g n o m s )
'hs ar am.
h ,.lttrs''(nulotids)givs un-
argumntsfor volution, asd on
t analogy, th first rgumnt says
: (thsam hrditarymolui, th
ls,nd th sanr20 amino aids in
rary (thsamgntiod), annot
ul-nntonrns similarity in th s-
hus th squnof amino aids in
similar tts annot of indpn-
||ed iformtiol mromoluls
mad up of sqnsof units-
nino ids in th s of protins-
utilnary infornration. lmprisln
Wo maromolulsstalishsth
tion usually Oursy hanging on
; is an indiirtion of th rny of
; o f n u l o t i d so r o f a m i n o a i d s
, in humans and himpanzs th
l i h s r v s v i |f u n r i ni n s p i -
04 amino aids in atly th sam
tohrom-of rhsus monkys y
additional mino aids, and from
i.
ny of ommon anstry.Thus in-
othr disiplinslnrningvolu-
lr studisof DNA and protins y
and amino aids. Th authority of
f th thousandsof gns and thou-
sm providsn indpndnttst of
:notal advantagsovr ompar-
:iplins.on is that th informatin
of units that ar diffrnt is radily
olulr uolution 17
stalishdwhn th squnof units is known for a givn maromolul in
diffrntorganisms. Th sond advantag is that omparisons an mad
vntwnvry diffrnt sorts of organisms. Thr is vry littl that om-
parativanatomy, for xampl, an say whn organisms as divrs as yasts'
pintrs,and human ings ar ompard, ut thr ar numrous DNA and
protin squnsthat n ompard in all thr. Th third advantag is
multipliity.ah organism posssssthousands of gns and prtins, whih
all rfltth sam volutionary history. If th invstigation of on partiular
gnor protin dos not satisfatorily rsolv th volutionary rlationship of
a st of spis, additional gns and protins an invstigatd until th
mattrhas n sttld to th satisfation of th invstigator.
orovr, th widly diffrnt rats of volution of diffrnt sts of gns
opn up th opportunity of invstigating diffrnt gns for ahiving diffr-
nt dgrsof rsolution in the tr of volution. volutionists rly on slowly
volving gns to ronstrut rmot volutionary vnts but inrasingly
fastr-volvinggns to ronstrut th volutionary history of mor rntly
divrgdorganisms.
Gns that nod th riosomal RNA moluls ar among th slowst-
volvinggns.Thy hav n usd to ronstrt th volutionary rlation-
ships among groups of organisms tht divrgd vry long ago: for xampl,
among atria, arhaa, and ukaryots (th thr major divisions of th liv.
ingworld), whih divrgd mor tan2 illion yars ago (s Figur 1); or
among th mirosopi protozoa (e.g., Plsmodim, wih auss malaria)
ompard with plants and animals, th thr of whih ar ukaryoti groups
of organisms that divrgd aout 1 illion yars ago. ytohrom-' mn.
tiond arlir' whih volvs slowly, ut not as slowly as th riosomal RNA
gns,is usd to diphr th rlationships within larg groups of organisms,
suh as among humans' fishs, and insts. Fst-volving moluls, suh as
th firinopptids involvd in lood lotting, ar appropriat for invstigat-
ing th volution of losly rlatd animals, for xampl, th primats:
maaqus'himpanzs, and humans.
olular Phylognisof organisms
DNA and protins provid information not only aout th ranhing of lin-
agsfrom ommon anstors (ladognsis)ut also about th amount of g-
nti hang that has ourrd in any givn linag (anagnsis).It might
smat first that quantifying anagnsis for protins and nuli aids would
impossil aus it would rquir omparison of moluls from organ-
ismsthat livd in th past with thos from living organisms. Organisms of th
past ar somtims prsrvd as fossils, bt thir DNA and protins hav
larglydisintgratd. Nvrthlss, omparisons twn living spis pro-
v i d i n f o m a t i o na o u t a n g n s i s .
As a spifi ampl, onsidr th prviously mntiond protin yto-
hrom-,whih is involvd in ll rspiration. Th squnof amino aids in
I8 oleulr uolution

this protin is krrown for many rganisms, fom tria and ysts to insts volutionary trs aI
and hunrans;in animals' ytohrom_ollsistsof 104 amio aids. Whn th volutionary histoy of
amino aid squnsof humans and rhsus monkys ar ompard' thy ar as gnra' familis, or t
found to diffrent at position 58 (isoluin in humans' thronin in rhsus kinds of information r
monkys), ut idntial at th othr 103 positions. Whn humns ar om- nsis. Figur 5 illustrat
pared with horss, 12 amino aid diffrnSare found, and whn horss ar th rltiv rlationshi
ompard with rhsusmonkys, thrar 11 amino aid differns(Figu4). of th figur, humans al
vn without knowing anything ls aout th volutionary history of mam- to h othr than ith
mals, on would onlud that the linags of humans and rhsus monkys di- shorvs that th last o
vergd from ah othr muh mor rntly than they divrgd fom th hors mot past than the last
linag. orovr, it an onludd that th amino aid diffrn twn volutionary trs m;
humans and rhsus monkys mr-rsthav ourd in th human ling aftr its ah linag' or nagl
sparation fom th rhsrrsmonky linag (sFigur 5). last ommon nsto
h a n g d i n t h l i n a g t
t o r h s u s m o n k y s .T h i
,l that at position 58'
5 ]0 15 20 25 0 5 40 45 50
Figur 4) hav th snr
Uman tHGLtGRKTGQAPGYsYAAN
GDVKGKKItIMKSQHVKGGKHKTGPN
n t o n ( i s o l u i n )w
,h
Monke
aftr it sparatd fom 1
Horse

5 5 6 0 6 5 7 0 1 5 8 0 8 5 9 0 9 5 10 0 1 0 4
umn KNKGIlWGDLMYLENPKKYIP6TKMI
FVGIKKKERADLlAYLKKATN
olular Ronst
Monkey .....7
Hors Svral mthods ist f
opd for intrprtirrg
Figur 4. Amino aid squnof ytohrm-protinsin humans,rhsus
data; som an usd
monkys,and horses.Th squnonsistsof 104 amino aids,ah
rprsntdby a lttr.A dot indiatsth samamino id as in humns.S usd in molula vol
Fisur 3 for th namsof th min aids. maimum liklihood.

D|sAN M|

A ..distan'' is th num
Humn
diffrns ar msur
data) or to rtin maI
--t.,,.n",,, protins or th squn
monke
in Figur 5 was otain
amino aid diffrns
protin. Th amino aid
is rfltdin th num
plamnt of on am
Horse
lotid substitution in
ass it rquirs at last
im -__---.----.->
numbr of nulotid di
Figur 5. Th volutionf ytolrrom-among hurnans,monkys,and horss. ssary to aount fo th(
Th numbsinditth mino :ridrplamnts that hav tkn pl:rin h lutionary tr basd on
ranh of th phylogny. nulotid hngs in z

from atriaand yasts to insts
Lsistsof 104 amino aids' Whn th
us monkys ar ompard, thy ar
in in humans,thoninin rhsus
.Wlrn
positions. humns ar om.
]s ar found, and whn horss ar
1 amino aid diffrns(F-igur4).
t th volutionary history of mam_
of humans and rhsus monkys di-
y than thy divrgd from th hors
t th amino aid diffrn twn
;urrd iir th human linas ftr its
l (sFigur.5).
25 0 5 40 45 50
LHGLtGRKGQAPGYsYTAAN
KHKGPN
t'.D..
i 80 85 90 95 100 104
)6TKMItVGlKKKERADLIAYLKKATN
n-protinsin humatrs,rhsus
; of 104 aminraids,ah
;mamino aid as in huI-rrans.
S usd in molula volution ar alld dist. maimum parsimony. and
11"*u"
---l
r-_\
0
----
onuru.
monkev
orse
tong humans, monkys, nd horss.
:mntsthar hav tak pla in ah lutionarytr asdon tlr data in Tale 1, shorvirrgth minimum nus of
olculr uolution 19
volutionary trs ar modls or hypothss that sk to ronstrut th
volutionaryhistory of taxa (i..,spisor othr groups of organisms, suh
as gnra'familis, o ordrs).As pointd out :lrlir' th trs ma two
kindsof information rlatd to volutionary lrang'ladognsisand ang-
nsis.Figur .5illustratsboth. Th rarrhigltionshipsof th trs flt
th rlativrlationshipsof anstry, or ldognsis.Thus in th right sid
of th figur,hunrns and rhsus monkys 21sn to mor losly latd
to ah othr than ithr one is to th hors. Statd anothr way' this tr
showsthat th last ommon anstor to ll thr spislivd in a mor r-
motpast than th last ommon anstor to humans and monkys.
volutionarytrs may also indit th hangs that hav ourrd along
ahlinag,or anagnsis.Thus in th volution of ytohrom- si th
lst omnron zlnstor of humans and rlrsus monkys, lr-rarino aid
hangdin th linagthat Wnt to httmns ut none in th ling tha Wnt
to rhsusmonkys' This lnlusionis drwn from th osrvation (Figur 4)
that at positin 58, monkys and horss (as wll s othr animals; s
Figur4) hav th sam amino aid (thronin)'whil humans hav a diffr-
nton (isoluin),whih thrformust hirvhangd in th human linag
aftrit sparatd frolrl th monky linag.
olularRonstrutionof vlutionary History
Svralmthods xist for onstruting volutionary ts.Som wr dvl-
opd for intrprtingmorphologial data, othrs for interprting rnolular
dat;som an usd with ithr kind of data. Th main mthods urrntly
m a x i m u ml i k l i h o o d '
DlsAN oDs
A ..distan''is th numr of diffrnstwntwo kinds lf organisms. Th
diffrnsar masurd with rspt to rtain traits (e.g., morphologial
data)or to rtain maromoluls (primarily th squn of amino aids in
protinsor th sqttnof nulotids in DNA or RNA). Th tr illustratd
in Figur 5 was otaind y taking into aount th distan, or numr of
amino aid diffns,among thr orglisms with rspt to Prtiular
protin.Th mino aid squnof a protin ontins mor intrmtionthn
is rfltdin tlr numr of mino aid diffnsaus in som ass th
rplamntoi on amino aid y anoth rquirs no mor thurn on nu-
lotidsustitution in th DNA that ods for th protin, whras in othr
assit rquirs at last two nulotid hangs. Tal 1 shows th minimum
numrof nulotid diffnsin th gnsof 20 sparatspiesthat is n-
ssaryto ount for th amino aid diffrnsin thir ytohrom-. An vo-
nulotid hangsin h ranh, is illLrstatd
in Figur 6.
 -l
Tl th gns dirlg [r tlhrtlm- in 20 orgnisrs.
. Minimum rrttrrlf rrr'rlotiddiffrns trvt.t

Organisrn r0 1t 12 1-l \4 l5 T6 17 1u t9 20
1.umln - r t3 17 16 1-l 12 t2 17 t6 18 18 19 20 ) i
.) t -)-)
1/
-)o 6 56 66
2. onk1, _12 16 l-5 t2 11 1 16 1.S 17 t7 18 ZT 2 2 5 62 57 6_5
. Dog 10 8 4 6712 12 14 t4 l-l 30 29 24 28 64 61 66
4. Flors -1 5 11 11 t6 16 16 \7 \6 -'t-z- 27 24 3 64 60 68
.5.Donky 4 10 t2 15 15 15 16 15 .tl 26 25 2 64 59 67
6. Pig 6 7 ] 1 t |4 1 30 25 26 31 64 59 /
- .10
7. Rabit 7 8 1r 11 11 25 26 2 29 62 59 t)/
8. Kangarlo -14 t4 15 13 14 30 27 26 31 66 -t8 68
9. Duk -)
11
.) -)/ 2"4 26 2s 29 6\ 62 66
l0. Pigon -4 48 24 27 2.6 30 .59 62 66
11 . h i k n 28 28 26 26 31 61 62 66
12.Pnguin -6 28 27 28 30 62 61 65
1. Turtl 0 27 30 33 65 64 67
14.Rattlsnk 8 40 4| 6t 61 69
1 5 . u n a _44I 72 66 69
16. Srwwormfly _16 .'8 6 65
1 7 .M t h s9 60 6\
|8. Nurlspor <'7
61
19. Sbroms |
20. ndid

Slure: Aftr Fith and Mrgtllias 1967 .

B 6 ia ] a ! J-i

i-lo-;1:!QJ v=.-

_151E ] *.: -
- ;
,9

i e. >-;+P+
' a
d * (D :'

0.o.
iq.=.:1 ;J - 5
i' a
^w'^*:v
a.-; = a: o = L:.
r }
rl^'p:-

}3li91;fii
} g i * 1 a i i
. ^ F - I

;;*5n
i <

3 g 1; a .' }!
*
i., - . )
rz a =
- '' l '
,a
-
rN.t
 oleculruolutio 141

}* un

i - m - * ! a Sroworm
....ot-+ ||

!N.Od-..)l l rA"^
o.-tc '
9.9
''lN.j |

tl'

oooo r' Satro.

.'l 'l o /.nyCeS Neurospora

o
l

.t;

Figur6. Phylgnyof 20 <lrganisms, asdon diffrnsin th anrin<laid

squn of ytohrome-.Th minimum numrof nulotidsustitutions
rquirdfor ah ranhis shown. Although frtionlnumrsof nulotid
sbsituinsnnot our' th nurbsshown r thostht fit stth data in
Tabl1. (AftrFith and argoliash 1967.)
t

The rlationships twn spis as shown in Figur 6 orrspond fair|

wll to th rlationships dtrmind from othr sours' sllh as th fossil

- rord.Aording to th figur,hikns ar lss losly rltd to duks and
pigonsthan to pnguins, and humans and monkys divrgd from th oth
-
^: mammalsforth marsupial kngaroo sparatdtrom th nonprimt pla-
i..
ntals.Although ths xampls r known to rronous rlationships,
a

.6: s :
th powr of th mthod is apparent in that a singl protin yilds a fair| a-
= i
= O > -
t*
<.* uratreonstrutionof th volutionary history of 20 organisms that startd
l . . _ - / * > \
: : = : l R
]ti.,r>zd ,.
to divrgmor than 1 illion yars ago.
Th most ommon produr to transform a distan matrix into a
-j^i-; =t.] .od.
_***it+!
phylognyis alld lustr analysis. Th distan matrix is sannd for th
1 4 2 oleulr uolutirl

smallst distan lmnt,and tlr two taa involvd (say, human and mon- from th linags that .
ky in Tal 1) ar joinlat an intrnal nod, or ranhing point. Th matri vors' fr exampl' dol
is sannd gain fr th nt-smllstdistirn,nd th two nlv ta (sy' gthr aus they pol
hors zrnd donky) ar lustrd. Th produr is ontinud until ll taxa ommon anstor bran(
.!fhn
hav n join1. a distan involvs a taxon tht is alrady }rertof a and oyots.
pviluslustr, th avagdist:inis otiirltwnth n'ton irnl
th prxisting lustr (say, th avrag listantwn humurnand hors
MA|U-LlKL
and monky anl hors).This simpl prtldurssllmstht th rat of vo.
lution is uniforrn alorrg :rll ranl.rs. aximum-liklihood mt
Othr distan mthods (inluding th on usd ttl onstrut tlr tr in availal data. Thy r<
Figur 6) rlax th ondition of r"rniformrt and illlow for rrnqulrats of would mak it possiblt.
volr.rtionallng th brirnhs.or-rof th most tnsivl1,usd ll-rthodslf hang. For ampl, as
this kirrd is alld nighor-joining.Th thod starts' as for,y idntify- mor likly than transv
ing th smallst distan in th mtrix and linking th two ta involvd. Th proaility must assl
nt stp is to mov ths two ta nd alul:lta nw mtri- in whih p o i l i t i sf o r a h i n
thir distans to othr taa ar rplad y th distan twn th nod tr is th on with th I
that links th tw.o taxa and all otlrr taa' Th smallst distn in this nw all possibl trs'
matrix is usd for making th nt onntion' rvhih will twn trvl aximum-liklihood
othr taa or btwn th prvious nod and a nW taon. Th produr is numr of taa is larg
ptduntil all tax hav n onnt]with Oll an()thy intvning whih th proability m
nods. of taa. With 10 taa,
taxa, t numr of pos
A x l M U r . 4 . P A R s
l oNY MoDs vn with powrful o
h i i t i v i f t h n u m ro t
aimum-parsimony mthods sk to ronstrutth tr that rqrrirsth t i m r q u i r d .H u i s t i
fwst(i..,most parsimorrious)numr of hngsstlnrmdltlongll rrhs. sil ts is xmind,
This is rasorralassumption ausit usually will tlr rnost Iikly. But
volution may not nssarilyhav ourrd follorving a minimum path, -
sA|slAL AP|
us th sarn lrang irrstadnray hav <lurrdirrdpndntlr'alorrgdif-
frnt branhs,and som hangs mav hav involvd intrmdiatstps. Th statistial dgr o1
Not all volutionary lrangs,vn thos that invlv a sirrglstp, may and maximum-likliho
quilily proa bl. For ampit lnong tlr t<lurrrr.llltil ssirr DNA, y- strapping. It onsists of
tosin () and thymin (T) ar mmrs of a family of rlatd moluls data point at random a
alld pyrimidirrs;likwis, adr-rin (A) nd guanin (G) llngto a family random sampling pro
of rrrolulsalld pr-rrins.A hang withir-ra DNA sqlln om on ootstrap valu for al
pyrimidin to nothr ( *T) or from on purin to anothr (A *' G)' alld all spis drivd from
transition, is mor liklv to our than a hang fom a purin to pyimi. bootstrap valus aov
d i n o r t h l n v r s ( G o r A . ' o r T ) , a l i la t r n s v s i l nP. r s i m o n r ' thos with ootstrap val
mthods tak into ount diffrnt proailitis of ourrn if thy ar
known.
imum-parsimony mthods ar rlatd to ldistis' a vry formalisti lVlolular Phylog
thory of tonomi lassifiationthat is tnsivlyusd with morphologi.
al nd palontologial dl.rta.Th ritil fatur i ladistis is th idntifi. Th mthds for otai
ation of drivd shard traits,lld syn:1pomorphitraits. A synapomorphi mously improvd in r.
trait is shard y som taa ut nlt othrs l.lsth formr inhritd it gns hav bn squn
from ommon nstor thar quird tlr trait irftr its lirragsperatd has n squendin r

< ai n v o l v d( s a y ,h u m n n d m o t r -
d, lr ranhingpoint. h matri
tan,and th two nw t (say'
ldur is ontinud until all ta
3s taon tlrat is lirdypart of a
ltaindtwtth nw taxon arrd
istantwnhuman rndhos
duassutt-ts tht tl-rrt of vo-
on usd to onstrut th tr in
.at i1ndallor,vfo ltll'qual rats
most tnsivlyusd mthods of
thod starts,s for,y idntify-
lirrkirrgrh rrvo ta irr'',olvd.Tlr
j a l u l t a n w m t r i i n w h i h
y th distirntwtth nod
. Tlr srnallstdistarr in this rrw
tiorr,whih will twn trvo
rnd a nw ta()n.Th produr is
. J ' i h ( ) n : l I 1 ( ) t h rl ) \ . i l . l
s vn with powful omtrs' mximum-liklihood mthods an b pro-
onstrutth tr that rquirs t
:hngssummdalong ll rarrhs.
u s u l l yw i l l t h r o s t l i k l y . B u t
" df o l l o w i n g m i n i m u m p a t h '
ourd indplrdrrtli,along dif-
lve involvd intrmdiatstps.
tht involv a singl stp, may
: forrr nulotid ass in DNA, 1..
; of a family of rlatd moluls
rnd guanin (G) long to a family
rithin DNA sqrrnefrom o
purin to anothr (A.'G), alld
h n g , fr l m p u r i n t t l p y r i m i -
, alld transvrsion.Parsimon1,
labrlittsof lurrnif thy irr
td to ladistis'a vr formalisti
tnsiviyusd with nlophologi-
faturin ladistisis tlr idntifi.
lomorphi trits.A synapomorpi
lrs austlr formr inhritd it
h tait ftr its linag spatd
oleulruoltion 14
from th linages that wnt to th othr taa. In th volution of arni-
vors,for ampl, domsti ats' tigrs' and lopards ar lustrd to-
gthraus they possss rtratal laws' a trait aquird aftr thir
ommon ansto ranhd off from th linag that ld to dogs, wolvs,
nd oyots.
MAxlMUM-L|KL|HooD HoDs
Mimum-liklihood mthods sk to idntify th most likly tr, givn th
avilal data. Thy rquir that an volutionary modl idntifid that
lf would mak it possi1to stimatth poabiiity of ah possibl individual
hang.For xampl, as is mntiond in th prding stion' transitions ar
mor likly than transvsions among DNA nulotids, ut a partiular
proailitymust assignd to ah. Ali possibl trs ar onsidrd. Th
proailitisfor ah individual hang ar multiplid for ah tr. Th st
tris th on with th highst probaility (or maimum liklihood) mong
all possiltrs.
aimum-iikelihood mthods ar omputationally pnsiv whn th
numr of taa is larg aus th numr of possil trs (for ah of
V l l i l l g
whih th proility mst alulated)grows fatorially with th nmbr
.!7ith
of taa. 10 taa, thr ar aout 3.6 million possil trs; with 20
taa' th numr of possil trs is aout 2 followd y 18 zeos (21018).
hiitiv if th numr of ta is larg, aus of th onsidrl omptr
tim quird.Huristi mthods xist in whih only a susampl of all pos-
siltrsis xmind, nd thus an xhastivsarh is avoidd.
-
sAIs|AL APPRAIsAL oF VoLU|oNARY Rs
Th statistil dgr of onfidn of a tr an stimatd for distan
and maimm-liklihood trs. Th most ommon mthod is alld oot-
strapping.It onsists of taking sampls of th data y moving at last on
data point at random and thn onstruting a tr for th nw data st. This
random sampling pross is rpatd hundrds or thousands of tims. Th
bootstrap valu for ah nod is dfind y th prntag of ss in whih
all spisdrivd from that nod appar togthr in th trs. Nods with
ootstrap valus aoe 90o ar rgardd as sttistially strongly rlial;
thoswith bootstrap vlus elow 70o ar onsidrd unrlibl.
olular Phylogny of Gns
Th mthods fo otaining th nulotid squns of DNA hav nor-
mouslyimprovd in rnt yars nd hav om lagly utomatd. Many
gnshav n sqund in numrous organisms' and th omplt gnom
has bn squndin many spis,ranging from humans to atria. Th
144 oleculr uoltion

us of DNA squns has n partiularly rwarding in th study of gn

dupliations. Th gns that od for th hmlgloins in humans and lthr gs
myogIobin N1
mammals provid a good xampl.
Knowldg of th amino aid squns of th hmogloin hains and of
myogloin, a losly rlatd protin, has mad it possil to ronstrut th
volutionary history of th dupliations tht gav ris to th orrsponding
gns. But dirt xamination of th nulotid squns in th gns that \
od for ths protins has shwn that th situation is mo omple, and
also mor intrsting,than it appars from th protin squns.
DNA squnstudis on human hmogloin gneshav shown that thir
numr is grater than prviously thought. Hmoglobin moluls ar
ttramrs (molulsmad of four suunits)that onsist of two polypptids
(rlativly short protin hins) of on kind and two of anothr kind. In m-
ryoni hmogloin , on of th two kinds of polypptid is dsignatd ; in
ftal hmoglogin F, tt ls ; in adult hmogloin A, it is F; and in adult hmo-
gloin A,, it is 6(hmogloin A maks up out 98" of humn adult hmo-
globin, and hmogloin A, aout 2"|. Th othr kind of polypptid in
mryoni hmogloin is (; in oth ftal and adult hmogloin, it is a. Th
gnS that od for th first group of polypptids (, y, 0, and loatd
on hromosom 11; th gnsthat od for th sond group of polypptids
(( and a) ar loatd on hromosom 16.
Thr ar additional omplitis.Two y gnseist (known s G,and A/'
as do two 0 gns (t' and a,). Furthrmor, thr ar two B psudogns
(B, and VF,) and two gpsudogns (a,and {{,),as wll as a (psudo-
gn. Ths psudognsar vry similr in nulotidsqunto th orr-
sponding funtional gns, ut thy inlud trminating odons and othr
mutations that mak it impossil for thm to yild funtional hmogloins.
Th similarity in th nulotid squnsof th polypptid gns and
pseudogns of oth th a and B gene familis indiats that thy ar all
homologous-that is' that thy hav arisn through various dupliations and
susqunt volution from a gn nstral to all. Morovr, homology also
ists twn th nulotid squnsthat sparat on 8n from anothr.
Th volutionary history of th gns for hmogloin and myogloln is sum. Figur 7. volutionaryh
marizd in Figur 7. whih nstralgnsdt
p p r o x i m a t t i m sw h
y a s( Y 1 a g o .T h t i n
Nlultipliity and Rat Htrognity unrtin.

ytohrom- onsists of only 104 mino aids, nodd y 312 nulotids.

N v t h l s st, h i s s h o r t p r o t i n s t o r s n o m o u s v l u t i o n a r yi n f o r m a t i o n '
whih mad possil th fairly good approimation' shown in Figur 6, to of ytohrom- mol
th volutionary history of 20 ver divrs spis ovr a priod longr than twn losly rlatd
1 illion yars. ytohrom- is a slowly volving protein. Widly diffrnt ytohrom- in humz
vrgd 6_8 million ya
spis hav in ommon a larg proportion of th amino ids in thir
divrgd from thir ol
ytohrom., whih maks possibl th study of gnti diffrns twn
organisms only rmotly rlatd. Fo th sam rason, howvr, mparison only on amino aid r
 olulr uolutio 145

rly rwarding in th study of gn gns

hemoglobin
hmogloinsin humans and othr gns
myogIobin p

s of th hmogloin hains and of

nad it possilto ronstrut th
hat gav ris to th orrsponding
otid squnsin th gns that
r situation is mo ompl, and
r th protin squns.
globin gnshav shown that thir
ght. Hmogloin moluls ar
:s)that onsistof two polypptids
Ldand two of anothr kind. In m-
ls of polypptid is dsignatd s,.in
loin A, it is B; and in adult hmo-
aout 98% of human adult hmo-
Th othr kind of polypptid in
and adult hmogloin,it is a. Th
pptids(, 0, and ar loatd
,r th sondgroup of polypptids

r gensist (known as G,and A/,

lor' thr ar two B psudogns
, and ,\, as wll as a (psudo-
n nulotid squnto th orr_
rd tminatingodons and othr
n to yild funtional hmogloins.
rs of th polypptid gns and
amilis indiats that thy ar all
n through various dupliations and
al to all. Morovr, homology also 1,100N/Y
tatsparaton gn from anothr.
rmogloinand myogloin is sum- Figur7. volutionaryhistory of th glin gns.Th forks indiatpoints at
whihanstralgnsduplitd,giving ris to nWgnlinags.Th
appr.oimattims whn thsdupliations ourrd ar indiatd in millions of
yars(Y) ago.Th tim whn th dupliationof ' and r, ourrdis
)rty unrtaln.

ids,nodd 1'2nulotids.
iormous volutionary information,
rimation, shown in Figur 6, tl of ytohrome- moluls annot dtrmin volutionary rlationships b-
spisovr a priod longr than twn losly rlatd spis. Fr ampl, th amino aid squn of
volving protin. \/idly diffrnt ytohrom- in humans and himpanzs is idntial, although thy di-
rtion of th amino aids in thir vrgd6_8 million yars ago; twn humans and rhsus monkys, whih
tudy of gntidiffrns twn divgdfrom thir ommon anstor 5_40 million yars ago' it diffrs y
s a m a s o n h. o w v r , o m p a r i s o n only on amino aid rplamnt.
146 oleculr uolutio

Protins that volv mor rapidly than ytohrom- an str'rdidin or. .W.ithin
r|ir. ah
dr to stalish phylognti rlationships twn losly rlatd spis. v o l v a t d i f n tr t
Som protins volv vry fast; th firinopptids-small protins that ar vnts. Sintists n
involvd in th lood-lotting pross-ar suital for ronstruting th th volutionary phy
phylogny of rntly volvd spis' sh as losly rlatd mammals. n u m r o f g n si n v
othr protins volv at intrmdiat rats; th hemogloins, for xampl' tion twn gns o
an usd to ronstrut volutionary history ovr a fairly road rang of g n s o r h i v i n gd l
tim (sFigur 8). volving gns an
On grat advantag of molulr volution is its multipliity, as notd t h a r n o d s l t l w l v .
l u t i o n r yr l a r i o n s h i
rdundant odon su
th nodd mino i
PE
:' ;e ! l sprsd among th s
U }a.
;::>
sgmnts of th gns
G o d i n g p r t i r l n so f g
.9-aE
g . l o t i d st h e t s p i f vr
> nd
.!
s .J s
tt ttt t
vv v v rr}' Th }yIolulrl
0
160 O n l l n s p i u o t ' ts t t r i
h o m < l I o g o um s olul
140 pl. popotionsf nL
I v o l u t i o n a r y h a n g
1?n
I
t o D N A o r p r o i n st h
00
tion. Studis of mll
100 "r"*
maromrlulsmay s
.s,9,
80 It ws first osrv
ns twnhomolt
60 n a r l y p r o p l r t i o n a lt t
r o r . I f r h a t o f rt
smin th voIui
D N A s q u n sW o u
q u n s o u l d r h n h
ing vnts of a phyl
ourrd.
onsidr, for am
gn oding for yto}
o u l d d t r m i n t h t
0O simply y amining
0
O9UUO
o 0
r n h .o n w o u I dn
0 10 2oo 30o 40o 5oo 600 7ao 600 9oo 10oo
E<^ sour' suh as th tt
v
t illins of yrssin divrgne
t i m r h a t l p s di n r
Figur 8. Rats of molular volution of thr diffrnt protins. (Aftr T h m o l u l r v o
Dikrson 1971.\ lok, lik a wath
 ollr uolutio 147

ytohrom-an studid in or- alir.Within ah organism af tousands of gns and protins; ths
ps btwnlosly rlatd spis. volvat diffrnt rts, ut vry on of thrrrrfltsth snrvolutionary
ropptids_small protins that ar \.nts.Sintistsan otain gratr and gratr auray in ronstruting
-r suital for ronstruting th th volutionary phylogny of any group of organisms y inrasing th
suh as losly rltd mammals. numrof gnsinvstigatd.Th rang of diffrnsin th rats of volu-
tts;th hmogloins,for ampl, tion twn]I.lsopns up th opportunity of invstigatingdiffrr-rt sts of
histoy ovr a fairly road rang o{ gnsfo ahivil-lgdiffrnt dgsof rsllutiorrin th tr of lit; slowly
volving]nsn b usd to study rmot volutionary vllts. vn gns
olution is its multipliity, as notd thatndslorvly vlving protins an usful for ronstrutingth vo-
lutionaryrlationshipstwnlosly rlatd spisy amintion of th
dundantodon sustitutions (nr.rlotid sustitr'rtions
that do not hang
th nodd mino aids), th intrns (nonoding DNA sgmnts intr-
sprsdamong th sgmrrtstlrirtod fo mino ids), or othr nonoding
a

e sgmntsof tlr gns (suh as rlr squnsthat prd and tollow th n.

o. oding portions lf gns);ths gnrally volv muh fastr than th nu-
EE
: !
lotidsthat spify th amino aids.
\.:
t
9
U
li
tl
V\Y Th olular lok of volution

on onspiu()tlsattriut of molular volution is tht diffrnstwn

homologusmolulsan radih' quantifid and xprssdas, for am-
pl, proprtions trf nulotids o amino aids tlrat hav hngd' Rts of
volutiorrirrvlrangan thrfo mor prislystalishdrvith rspt
to DNA r pr<ltinsthan witlr rsptto phnot,vpitrits of forrrrand fun-
{\+'/ tion. studis o| molulr volution rats hav ld to th proplsition that
maromolulsmay srv s volutionary loks.
ryq"' -l
It was first osrvd in th 960s tht th nunrrs of amino aid diffr-
nstwntllntllogous protills of ny twrl iivl-lspis sm]to
nrly proportilnl to th tirrrof thi divrgn from a olnmon ans.
tor. If th rat of volution of protin or gn Wr appro-\imatly th
samin th volutionary linags that ld to diffrnt spis,protins and
DNA squns would provid a molular lok of volution. Th s-
qunsould thn usd to ronstrut not Onlv th squnof ranh-
ing vnts ot a phylgrl1, ut lso th tir whn tlr vilrious vnts
ourrd.

fE onsidr, for ampl, Figur 6. If th sustitution of nulotids in th
gnoding for ytohrom. ourrd at a onstant rt through tim, n
.' ould dtrmin th tim that lapsd along any ranh of th phylogny

simply y arirring th nrrmr lf nulltid sustitutions alorrg that
ranh.on w<luldnd onl,vto alirat th lok by rfrntl an outsid
50 600 700 80 9oo 10oo
sour'suh s th fossil rord, that would provid th atul gologial
ears sine divrgne
tim that lapsd in at last on spifi linag.
hr diffrnt protins. (Aftr Th molul:r volutionay Iok is not xptd to b nltronomi
lok, lik a wth Or othr titr-rpi that msurs tim tly' ut
148 oleculr uolutio

stohasti lok' lik radioativ day. In a stohastilok th proaility

of a rtain amount of hang is onstant (for xmpl, a givn quantity of
atoms of radium.226 is xptd' through day, to rdud y half in
1',62ears), although som variation ours in th atual amount of hang.
Ovr fairly long priods of tim a stohasti lok is quit aurat. Th
normous potntial of th molular volutionary lok lis in th fat that
..tiks',
ah gn or protin is a sprt lok. ah lok at a diffrnt
rat-th rat of volution haratristi of a prtiular gn or protin-ut 3so
ah of th thousands and thousnds of gnes or protins provids an ind-
pndnt masur of th sam volutionary vnts.
volutionists hav found that th amount of variation obsrvd in th vo. z Z3
lution of DNA nd protins is grtr than is ptd from a stohasti
..ovrdisprsd,''
lok-in othr words, th lok is or somwhat rrati.
h disrpanis in volutionary rats along diffrnt linags ar not s.
sivly larg, howvr. So it is possil, in prinipl, to tim phylognti 0
vnts with as muh auray as may dsird,ut mor gns or protins
(aout two to four tims as many) must xamind than would rquird
if th lok wr stohastially onstant. Th avrag rats otaind for sv- Figur 9. Rat of nul
15 dots marks th tim
ral protins takn togthr om a fair| pris lok, partiularly whn
anstor (horizltIs
many spis ar studid and th volutinary vnts involv long tim pri.
protin hangs,that h
ods (on th ordr of 50 million yars or longr). solid lin drwn from t
This onlusion is illustratd in Figur 9, whih plots th umulativ sustitution.(Aftr Fit<
nmr of nulotid hangs in svn protins against th dts of divr-
gn of 17 spisof mammals (16 pairings) as dtrmind from th fossil
rord. Th ovrall rat of nulotid sustitution is fairly uniform. Som
primat spis (rprsntdy th points blow th lin at th lowr lft of
Figur 9) appar to hav volvd at a slowr rat than th avrag for th volution is slowr, t}
rst of th spis.This anomaly ours austh mor rnt th divr- rasingly rmot.
gn of any two spis,th mor likly it is that th hangs osrvd will
dpart from th avrag volutionary rat. As th lngth of tim inrass,
priods of rapid and slow volution in any linag ar likly to anl on onlusion
anothr out.
volutionists hav disovrd, howv, that molular tim stimats tnd N4olular iology r
to b systmatially oldr than stimats asd on othr mthods and' in. th 1953 disovry
dd, to oldr than th atual dats.This is a onsqunof th statistial hmial. N4olular
proprties of molular stimats' whih ar asymmtrially distriutd. B- volution and maks
aus of han, th numbr of molular diffrns twn two spis muh dtail and pri
may largr or smallr than xptd. But ovrstimation rrors ar un- N4olular iology
oundd, whras undrstimation rrors ar oundd, sin thy annot unity of lif in th nat
smallr than zro. onsquntly, a graph of a typial distriution of sti- of nzyms and othr
mats of th ag whn two spis divrgd, gathrd from a numr of dif- lutionists, y making
frnt gns' is skwd from th normal ll shap, with a larg numr of that wr prviously
stimats of youngr ag lustrd togthr at On nd and a long tail of tionary relationships
oldr-ag stimats trailing away toward th othr nd. Th avrag of th organisms.
stimatd tims thus will onsistntly ovrstimat th tru dat (Figur As pointd out ar
10). Th ovrstimation is otns gratr whn th rat of molular sam asi omponn
 oltt|r uoltitl 119

n a stohastilok th proaility
rt (for ampl, a givn quantity o{
gh day' to rdud y half in
:urs in th atual amount of hang.
hasti lok is quit aurat. Th
lutionary lok lis in th fat that
. ah lok ..tiks'' at a difjrnt
cf a partiular gn or protin-ut s

gnsor protins provids an ind_
y vents.
rnt of vriation osrvd in th vo- z Z3
:han is ptd from a stohasti
rdisprsd''' or somwhat rrati.
ong diffrnt linags ar not s-
in prinipl, to tim phylognti o 2s 50 75 100
-125
d s i r d
, u r m o r g n so r p r o t i n s illions f ersof volutin
xamind than would requird
Th avragrats otaind for sv- Figur 9. Rat lf nuiotid sustitution ovr palontologial tim. ah lf th
rly pris lok' partiularly whn ]5 dots marks th tim t whih a pair f spis divrgd from Ommon
anstor (horizotl scle) and th numr f nulotid sustitutions, or
) n a r y v n t si n v t r l v l o n g t i m p r i -
protin hangs' that hav ourrd sin th divrgn (uertil sl).Th
ngr).
solid lin drwn from th origin to th ()utrmost dot givs th avrag rat of
Lr 9, whih plots th umulativ sustitution.(Aftr Fith 1976.)
rrotins against th dats of divr_
ings) as dtrmindfrom th fossil
rstitutionis fairly uniform. Som
s low th lin at th lowr lft of
owr rat than th avrag for th volutionis slowr, th squnsusd ar shortr, and th tim oms in-
austh mor rnt th diver- rasingiyrmot.
it is that th hangsosrvd will
.A s t h l n g t ho f t i m i n r a s s ,
ny linagar likly to anl on Conlusion

, that molular tim stimats tnd
i asd on othr mthods and, in-
ris is a onsqunof th statistial
ar asymmtrially distriutd. B-
ar diffrnsbtwn two spis
But ovrstimation rrors ar un.
; ar boundd' sin thy annot
h of a typial distriution of sti-
d, gathrdfrom a numr of dif-
ll shap,with a larg numr of
hr at on nd and a long tail of
th othr nd. Th avrag of th
)vrstimatth tru dat (Figur
ratr whn th rat of molular
olular iology mrgdas a disiplin 100 yars ftr Darwin' following
th 1953 disovry of th doul-hli strutur of DNA' th hrditary
hmial.Molular iology provids th strongst vidn of iologial
volutionand maks it possil to ronstrut volutionary history with as
muh dtail and prision as anyon might want.
Molular iology provs volution in two ways: first, y showing th
unity of lif in th natur of DNA and th workings of organisms at th lvl
of nzymsand othr protin moluls;sond,and most importnt for vo-
lutionists' y making it possil to ronstrut volutionary rlationships
that wr prviously unknown, and to onfirm, rfin, and tim all volu-
tionary rlationships from th univrsal ommon anstor up to all living
organisms.
As pintd out arlir in this haptr, all organisms ar mad up of th
samasi omponnts,whih, morovr' are similarly assmldand usd.
150 oleulr uoltiln

't8 organisms, from batr

now ronstrutd w
't6
iology. Darwin ould r

-14
I
o B|LIoGRAPHY
!12
Avis,J. .2004. olu
10
o A B
Sundrland,A: Sin
A y a l ,F . J . , d .1 9 7 6 . o l
*B B a k r ,A . J . ' d .2 0 0 0 . o
o
Lr- arroll, S. B. 206.h
Rlrd of ullutil
h

D i k r s o nR , . . 1 9 7 1 .T h
volution.lourl of 1
Flsnstin, l. 204.lfer
Fnhl,. 2002. Th ri1
UnivrsityPrss.
Fith,.W.x4. 1976.Mol
15 5 55 75 95 1.l5 .15 .155 175 195 215 uolutio,160-.l78. l
Fith, \V. N4.,and . arg
Divergenetime(x100MY) I55 279-284.
Grur, D., nd \7.-. Li. 2
Figur 10. Skwingand ag ias in stimatingmolulardats.h instshows Sundrland,MA: Sin
th tr topology for linagsA, B, and ; t.. and t rprsnt'rsptivly' Hll' B. G.2004. Phlog
alirationand targttims.Th min panl shows a fqunydistriutionof Assiats.
1,000 stimatsof th divrgntim twnlinags and AB in th inst' Hazn, R. . 2005.ge.e
st to hav ourrd3 billion yars ago and taindwith th us of a short (7.5 D: Josphnry Prt
rsidus),slow.vlving(onrplmntpr sit pr 1010yars)protin and Lynh.M. 2007. h orig
with th us of th split twnA and B, st to 300 million yarsago' as a Assoiats.
aliratilnpoint. T and M rprsntatual (3 illion yars)and stimtdman Ni, \4.,and S.Kumar.2
(4'084 illion yars)tims.(odifid from Rdriguz-Trlls t a|.2002.| Oxford UnivrsityPr
Rdriguz_Trlls' F., R. T
toward vrstimatio
Ntir.lldmof
.Wos,
C. R. 1998.Th ur
SisUS 95: 68.5
In all organisms' from atria to humans, th instrutionsthat guid th d-
vlopmnt and funtioning of organisms ar nasd in th sam hrditary
matrial, DNA, whih provids th instrutions for th synthsis of protins.
Th thousands of divrs protins that xist in organisms onsist of th sam
20 amino aids in all organisms, from atria to plants and to animals. Th
gnti od, y whih th information ontaind in th DNA of th ll nu-
lus is passd on to protins, is shard y all sorts of organisms. All organ-
isms us similar mtaoli pathways-squns of iohmial rations-to
produ nrgy and to mak up th ll omponnts.Th unity of lif rvals
th gnti ontinuity and ommon anstry of all organisms.
Th dgr of similarity in th squnof nulotids in th DNA (and
thus th squnof amino aids in th nodd protins) maks it possil
to ronstrtvolutionary history. ah of th thousands of gnsand thou-
sands of protins prsnt in n organism provids an indpndnttst of th
organism's evolutionary history. Th volutinary lationships among all
 oIculr uolutio i ),

organisms'fror batria and protozo to plnts, anirnals, and humans an

-+ now b rollstrutdwith as muh dtail as wantd y mans of molular
t iology.Darwin ould not hav hopd for mor.

t BILIoGRAPHY

. 2004.ollrrkrs,Nturl istor,d uolutil'2nd ed.

,l ,l
115 15 55 175 95 215
ime(xl00MY)
ng mllulardats.Th instshows
.. and t, rPrsnt'rsptivlv'
:Ishows frqunydistriutionof
,nlin8s and AB irr th inst,
obtaindwith th us of short (75
r sitpr 10l0yars)protin and
t to 300 million yarsago' as a
(3 illionyars)and stimatdmn
Rodriguz-Trlls t al. 202'|
s' th instrutionsthat guid th d-
ar nasd in th sam hrditary
ltions for th synthsis of protins.
ist i organismsonsist of the sam
tiato plirntsand to animals' Th
ontindin th DNA of th ll nu-
ry all sorts of organisrns.All organ-
unsof iohmial rrtons-ro
omponnts.Th unity of lif rvals
;tyof all organisms.
of nulotidsin th DNA (and
noddpotins)mks it possil
of th thousandsof gnsnd tho-
providsan indpndnttst of th
olutionary rlationships among all
Avis,.|.
Sundrland' N{A:SinurAssoi21ts.
Ayala,}-.l., d.|976, ollruolt,ttirl, Sundrland'MA: SinurAssoiats.
Bakr,A. J., d.2000.olulrtrldsi llog.Oford: Blakwll.
arroll,S. B. 2006. Th hig of the Fittst:DN d th UltimtForesl
.!?...W.
Rordof ullutil.Nw York: Norton.
Dikrsn, R. L. 1971.Th struturof ytohrom and th ratsof molular
volution. Jtlullof olulruolutio|: 2645.
Flsnstin,J.2004, IfrrigPhlognis, Sundrland, MA: SinaurAssoiats.
Fnhl,.2002, T origitl nd rl L,lllutitlof Lif, oford: Oford
Univrsit,v Prss.
Fith,W. I\4.1976.olulrvolutiorrary llks.In F. J. Ayla.d.,olulr
uolutkln,|60_178, Sundrland,]r,1A:SinaurAssoiats.
Fith,W. ., and . Margoliash.1967.onstrutionof phylogneti trs.Sine
155:279-284.
Graur,D.' nd \W.-H.Li. 2000.Fudmetlls of oleulruolutit.2ndd.
Sundrlnd, }vIA:SinaurAssoiats.
Hll,B. G. 2004.Phllogtireesd ils.2ndd. Sundrland, A: Sinaur
Assoi.rts.
Hazn,R. N{.2005. goosis:h Siti|icQuest for Li|'s rigi. Washington'
D: Josphl{nryPrss.
Lynh,M. 27,Th rigis of Com rchittur.Sundrland,MA: Sinaur
Assoiats.
Ni, ., and S. Krrmar.2000. oleulr uoltltiond Phlogris.oxford:
oford UnivrsityPrss.
Rodriguz-Tlls, F', R, Tarrio,and F. .].Avala.2002.A mthodologial ias
towardovrstimatin of molulrvolutionary tim sals.Prodigs of the
NtioIdmof SisUSA 99: 8112-8l15.
.!os,
. R. l998. Th univrsalanstor.Prodingsof the Nti-llAdem oi
Sins US 95: 6854_6859.
 intplay of diffrnt
ras in gnom stz
As with othr typ
th typs of h:rrrg
within individuals (r
volution of th Gnom matrial of volution
an sprad through t[
of th spis.Thr l
Brin h rlesluorth nd lutionary prosss.I
may ausindividual
Dborh hrlesrth ag; as w shall s,s
als is not th only d
is th random pros
nnts of gntlms thl
tions r finit in siz
hangs in th gnti information stord in th gnom ar th ultimat a- dtrministi flrs r
sis of volution. uh of voltionary iology involvs studis of th o- ovr gnrations, sor
srval haratristis of organisms and th hrital hangs on whih talishd (fixd)in al
thir volution dpnds. But gnoms thmslvsalso volv. Th gnoms trolld y th ffti
of ontmporary spis rflt illions of yars of volution, and thy ar th numr of rd
still volving, oftn surprisingly rapidly. Data on gnom organization hav rious variantsan b
n aumulating sin th ra of lassial gntis, starting with th dis- ordr of th ripro
ovry that gns are arrid in th hromosoms of lls and an Th rlativ nragnitu
mappd gntiallyand physially to spifi loations in th hromosoms. dirtion and spd
Th latr disovry that hromosoms ar mad of nuli aids has ld to n strongly mph
inrasingly fin.sal physial maps of gnoms' most rntly th omplt (2007).
squnsof (parts of th) gnoms of (singl individuals from) inrasing Bfor dsriing r
numrs of spis, now inluding svral spis of animals (e.g', . le- important faturso
gs Genom Squning Projt 1'9991International Human Gnom S- tionary fators that r
quning onsortium 2001; ous Gnom Squning Projet 2002;
Drosopbil 12 Gnoms onsortium 2007) and plants (.g.,Th Araidop.
sis Gnom Initiative 2001), as wll as ovr 80 atrial spis (Sharp t al. Th Compositio
2005). Gnoms of many mor spis will rtainly sqund in th
H GN|
nt fw yars, givn th rapid advans ing mad in squning th-
nology. h hmial sis
This information has gnratd rnwd inrrst in gnom volution, is a long strand tlf
whih an now b dsrid in unprdntd dtail, and two major oks linkd y sugar and
hav rntly n dvotd to it (Burt and Trivrs 206;Ln 2007). !ith 2008). Irr all llul
suh xtnsiv data in hand, W an hop to idntify som of th main pro. lotid is doul-st
sss involvd in gnom voltion. Modls of many of ths prosss - may doul-stran
ist, and som gnral thms ar lar. Gnom volution oftn involvs strandd DNA (su
various opposing fors, So that th organization of th gnom of a givn flunza virus). Th
spis, or of a givn gnomi rgion, may dpnd n situations that wakn orplmrrtary:wht
or strngthn on for or anothr. Partiularly intrsting situations aris nin, th othr stra
whn th volutionary intrsts of individual gnom faturs onflit with th omplmntary l
sltion that ts on th organisms thmslvs. Baus of th ompl thus dsrid ir
IJZ

Gnom
d lutionaryprosss.First, dtrnrinistifors, partiularly natural sltion,
1 may aus individuals with a givn trit to lav mor dsndantsthan avr-
in th gnomar th ultimat a-
iology involvs studis of th o-
l th hrital hangs on whih
ms]vsalso volv. Th gnoms
f yars of volution, and thy ar
)ata on gnom organization hav
al gntis' starting with th dis-
lromosoms of lls and an
ifi loations in th hromosoms.
mad of nuli aids has ld to
noms' most ntly th omplt
i n g l i n d i v i d u a l sf r o m ) i n r a s i n g
rl spisof animals (e'g., . le-
.ntrnational uman Gnom S-
:nom Squning Projt 2002;
7) and plants (.g.,Th Araidop-
:r 80 atrialspis(Sharp t al.
ill rtainly sqund in th
; bing mad in squning th'
.d intrst in gnom volution'
:ntddtail, and two major ooks
Trivrs 2006;Lh 2007). \lith
t o i d n t i f ys o m o f t h m a i n p r o -
Llsof mny of thsprosss-
Gnom volution oftn involvs
nization of th gnom of a givn
dpnd on situations that wakn
ularly intrsting situations aris
lual gnom faturs onflit with
mslvs.Baus of th omplx
uoltion of th Cenom 15
intrplayof diffrnt fors' hangs oftn our in on dirtion (.g.,n in-
rasin ]nomsiz) and thn latr rvrse.
As with othr typs of volutionary hang, w nd to undrstand oth
th typs of hangs in gnom rgnization and omposition tht aris
within individuals (mutations and othr molular hangs that ar th raw
matrialof volution) and alsl how a trait arrid initially y on individul
an spradthrough th whol population, lading to volution of th gnom
of th spis.Thr ar tWO main aussof this sprd, just as for othr vo-
g;as w shll s,suh Dawinian natual sltionat th lvl of individu-
als is not th only dtrministifor that ats on gnoms.A sond fator
is th random pross of gti drift, wic is spially rlvant for ompo-
nnts of gn()msthat do not afft orgnismal funtions. Baus popula-
tions ar finit in siz, gnti variants that ar sujt to suffiintlywak
dtrministifors will prin random flutuations in thir frqunis
ovr gnrations,somtims ausing an initially rar vriant to om s-
talishd(fid)in all individuals of a spis.Th rat of gntidrift is on-
trolid y th fftiu popliltio siee, whih is usually muh smallr than
th nunrrof rding individuals in a spis(Kimura 1983). vn dlt.
rious varints an om fid y drift if th intnsity of sltion is of th
ordr of th ri;lrlalof t fftivpopulation siz or lss (Kimura 1983).
h rlativmagnitudsof drift and dtrministifors thrforontrol th
dirtion and spd of volutionary hangs irr gnoms; this hs rntly
n strongly mphasizd as ausal fator in gnom volution y l,ynh
(2007).
Bfor dsriing ampls of hypothssnd tsts,w first outlin som
important faturstlf gnol orgnization nd siz and thn disuss volu-
tionary fators that may hav sapd thsproprtis.
Th omposition and Siz of th Gnom
H GN| MAR|AL
Th hmial asis <lfth gnti matrial-a gnom'smost asi fatur-
is a long strand of polynulotid' in whih four diffrnt nulotid bases
linkd y sugar and phlsphatgroups stor th gnti information (Lwin
200s). In all lluiar organisms, from atria to mammls, th poiynu-
lotidis doul-stranddDNA (th famous doul hlix), ut in viruss it
my doul-strnddDNA (.g.,th T4 virus tht infts atria),singl-
strnddDNA (suh as anothr atrial virus, tp174), or RNA (.g.,in-
fllnza virus). Th twl strnds of doul-strandd DNA moluls ar
orplmntary:whr th as prsnt in th squnof on strand is ad.
nin' th othr strand has thyrnin, and whr th first strnd has guanin,
th omplmntary strand has ytlsin.Th omposition of on strand an
thus dsrid in trms of th frqunis of th four nulotid ass
154 uolutio of th Geom

(A, T' G, and ), whil that of th doul strfur is spifid y th fr. vlus gnoms ar usuz
qunis of the as pairs A and G. RNA is gnrally single strandd, with flunza) pakag svr
a diffrnt sugar rsidu from that in DNA and with urail ass instad of into tlr sam vius pal
thymins' have diffrnt RNAs pa
Although th origins of th gnti od ar unknown' slf-rpliatirrg thus omin diffrnt
RNA moluls may hav n th arlist form of ..lif,'' ut th transition ing multipl inftions,
to gnoms that usd th muh mor stabl DNA ourrd long ago (Joy F{urns and otlrr
1989). Dol-stranddDNA also allows damag to ortdy opying isms long to th uki
from th undamagd omplmntry strand. Sin DNA largly laks RNAs DNA organizd into l
ability to at as a atalyst, this hang must hav gon along with th volu- lls (Figur I). h
tion of a mans of transriing on stand of a gn's DNA into RNA. In dal with th prolm
prsnt-dayorganisms th squnof bass in th ..mssngr''RNA tan- uls, and tromrs
srid from a gn dtrminsth squnof th amino aids in th protin sion to th daugtr
hain nodd y th gn (Figur 1). Thr ar also many RNA molulsof ukaryots, suh as th
dirt funtional signifian, whih r transrid (opid) frm DNA ut gno1,with just on
not translatd into an mino-aid squn. mat' th is rif d
ah mating partrrr.Tl
oRGANlzA|oN oF H GN| MAR|AL

any prokaryots' gnoms (atia and viruss) onsist of just a singl

DNA or RNA n-rolul,oftn irtrlar (Tal 1). Som atri (suh as T l l . G n m s i z s
Borelli, th aus of Lym disas)lrav svral sParat moluls; although orgnism

Viruss
Plnt viroids
hromosome Genes messenger spIid Protein Influlzavirus
RNA MRNA Rovirus
Phag tpX174

telomer
.
intron
+l trans|tion
-+
SV4O
Vaitriavirus
Batria
intron
f" . oli
'i',, i o1llsm gitlium
I Nulurrgnomsf trk
enromere Y east (Scchro|l,t))L:s
i ntron
Nmatod (orhbd
Fruit fly (Drosophil m
F Ftsh (Fugu rubrips)
FI
+# Slmandr

t
Mat-lrmal (Homo s|li
telomere ress (Arbido1l sis thlit
Miz (Ze ms)
Fritill11'p ritillri,lss
Ntlr: Arvitions:p
Figur 1' Chromosoms,showingthir major ftursand th andingpttrns kiloss;, mgass
that an snwith stiningmthods.Th figurlso shows a shmatiof a ? mns tht th gnnu
rgion of th hromosom ontaining thr gns'th mssngrRNA produd ? ? t h . r ri t i s b s do l i n .
y thm, and th individuaIprotinsnoddby th mssngrRNA. lr <lrgnisrs.
 uolutilof tb Gome ].i.i

l stutur is spifid y th fr- virus gnoms a trsually singl nroluls,som RNA viruss (sulr as in-
,IA is gnrallysirrglstrandd,with
flunza)pakag svrl diffrrrtRNA rrrolulsthat r, diffIltgns
'trAand with urail ass instad of
into th sam virus partil. Otlrr RNA viruss (.g.,alfalfa rr-rosaivirus)
havdiffrntRNAs pakagd into sparatvirus partils.Ths virussan
]od ar unknown, slf-rpliating thus omin diffrnt portions of thir gnom from diffrnt parnts dur-
:st form of ..lif,'' ut th transition ing multipl inftions,a primitiv typ of sxualitY.
l DNA ourrd long ago (Joy Humans and othr nimals, plants, fungi, and man,v singl_lldor!]an-
i damagto orrtdy oying isms long to h ukaryot division of lif' whih is dfind v hving th
nd. Sin DNA largely laks RNA's DNA organizd into linar hromosoms arrid within th rrLrliinsid
Jst hav gon along with th volu- lls (F.igur1). Th hromos<rtnshav spializd rgions (tlomeres)to
nd of a gn's DNA into RNA. In dalwith th prolm of rpliating th nds of dul-stranddDNA mol-
lss in fh ..mssngr''RNA tran- uls,and entromresthat ar quird for ort hromosom ti1nsmis-
rof th amino aids in th protin sion to tlr daughtr lls dr"rringl1divisiorr (Lwin 2008). N,{arrypinririv
r ar also many RNA moluls of karyots'sLthas th grn a|ga hlmdons, hav a prirllarily hploid
ransribd(opied)from DNA ut gnom'with ;ust on opy of h of th diffrrrthrmosorns.Whn lls
. mat' thr is a rif diploid stag, With a hrmosom st ontriutd y
ahmating partnr. This undrgosa spial forrrof ll divisirr (miosis)
l ARIAL

nd viruss)onsist of just a singl

(Tal 1). Som batria (suh as Tal1. Gnorrrsizs
lvralspratmoluls; although Organism Gnom Gn numr ('nom siz

Viruss
Plant viroils ssRNA 0 00p
spIied Protein Influnzavirus ssRNA 12 l 3 . . 5k
MRNA Rovirus dsRNA 22 2kb
Phag<p174 ssDNA 11 5 . 4k
translation sv40 dsDNA 6 ) l]

.+| + Vainiavius dsDNA .100 187k

Batria
, oli dsDNA 4,400 4 . 6
oplsmgitIium dsDNA 500 0 . 6
Nulrgnomsof ukaryots
Y ast(Sbro1)| s cruisi) 6,000 12b
Nmtod1lrhhditislg,s) 19,000 100
Fruit fly (Drosrlph il mellgstr) 14,000 1 4 0
Fis (Fug rubripes) 22,000(?) 400M

+# Salamandr
Mammal (Homl spins)
r ess(rb i dopsi s th li)
3 0 , 0 0 0( ? ? )
2 0 , 0 0 0( ? )
27,000
90,000
3,000
1 0 0 b
aiz (Z lttvs) .r0,000(??) .i,000
Fritillary (F ritil lri s sric) 2 7 , 0 0 0( ? ? l L20,000 M
Nofe: Arviations:bp, baspirs;ss, singlestrndd;ds, doul strndd;kb,
lr fatursand th anding pattrns kilbass; ' mgass.
figuralso shows a shmtiof a ? mnstht th gn numbr is lnll' pproximatlyknlvn fror th gnolrrsqlln'
gns'the mssngr RNA produd ?? that it is asdorinompletvidrror anallgrvit squnl glltlt-t-ts
<lfsimi-
d y th mssr-rgrRNA. lr rsanisrns.
1.'6 uolutir of tb Gom

to rstor th haploid stat.In this division' gl1sfrom th Parntal lls r

rOmindthrough riproal xhangsof potions of thir hromosoms
(crossig our: SeLwin 2008 nd Figur 2). In animls and highr plants
th diploid stagprdominats,nd th haploid sti.ltis oftn just a rif part

f
\
oI rh rprldutivpross'
Chromosom numrs in th haploid st vary widly among diffrnt
spis:16 in th as of hlmdomos,5 in Drosophil,2 in humans,
but somtims just a singl hromosom (in a spisof ant and an stris
roundworm). harrgs twn spis in homosomal stutur ar om-
mon, involvtng dtlplition.snd dItit.lltsof gneti matrial, inursiosof
parts of hromosoms (Figur 3), and trsloctils(hangsof matrial
twn diffrnt hromosoms). hromosom rarrangmnts our infr-
quntly in volution (roughly on pr gnom pr 10 million yars in mammals,
although som goups, suh s rodnts, hav rts 10 tims highr: ihlr
and Sr-rkoff2003). Ovr volutionary tim) tlr rlrdr of gnswithin a hro-
mosom gradually oms srambld y sussiv invsions. Svrl
human hromosotns sm to hav rmaind unhand in grr ontnt

rossingover Figur 3. A lrromlsomt

ABD hromosom, and th a
n rrrangdso that
othr way rttund from

PrentaI
hromosome
(though not ordr) ov
m a d u p < l fs p a r a r l

oD|NG AND N

or rpeatedgns
U n e q u a| ro s s ingo v r bet wendup|iated Gnom siz is dfin
loid gnom. A gnom
r t r n s r i di n r o m t
quns (Lwin 2008
qes ar'd itros.h
I
transrid but ar rr
Y
latd; in ukryots,ir
as th spli<lsot. |
that ar rsponsil fo
and timing of mssng
Figur 2. op: rossing ovr in a rgion of a homosom ontaining fiv gns,
oding u ftlnrion
A to , shlwing th rsulting rominant ombinations of th gns (indiatd
vn with omplt
y uppr. and lowras lttrs nd lso y lk rrd gry lins for tlr twl
diffrnr parlrtal romosonrs). Bottom: rlssing ovr in a gion rrf
g r l s i r s o d i n g
hromosom ontaining two dupliatd gns that r similar nor.rghthat thy q u n s .T h t a s k i s
may mispair, lading o unqual rossing ovr. portion of oding sq
 uolutio of tbe Genom 157

)n' gllsfrom th pantal lls ar

I
s of portions of thir hromosms
r e 2 ) . I n a n i m a l sa n d h i g h r p l a n t s
aploid statis oftn just a rif part
invertd rgion
l-
I
l st vary widly among diffrnt
s, .5in Drosopbil,23 in humans,
(in a spisof ant and an sris

!
n hromosomal strutur ar om-
ls of gnti matril, iursils of

I
lslotios(hangsof matria]
osom farrangmnts our infr-
n pr 10 miiliorr as in mamrnals,
lr non-invertdregion

I
hav ts 10 tirs highr: ihlr
l' th ordr of gnswithin a hro-
y sussivinvrsions. Svral -
raind unhangd in ]enontnt l.
ffi
!
,l
Figur 3. A hrtlnlosom invrsion. Th diagram shorvs th nding pattrn of a
hromosom, and t apparan whn th top prt of th hromosom has
\ lrarrangd so that part Of th gtror in th ight-hand hromosom is th
othr Way around from th orlr in th lft ln.
\
ff,H,i;:il' ParentI
hromosome
(thoughnot ordr) ov th ours of som 70 million yars, ut othrs ar
_/
t:i I mad up of sparatloks drivd from diffrnt atrstralhromosoms.

oDING AND NoNoDING sauNs

up|iated
or repatedgenes
Gnom siz is dfind as th numr of ass in a singl strand of th hap-
- loid gnom.A gnom has two main omponnts:oding squns(whih
ar transridinto mssngrRNA and nod protins) and nonoding s-
W3- quns (Lwin 2008). Nonodirrg squns inlud oth intrgllic s.
qunes and itrls' Introns ar sgmnts of squnswithin gns that ar
transrid ut a rmovd spliig for tlr mssngr RNA is trans-
latd;in ukaryots,intron spliing involvs a protin.RNA mahin known
WE
as te splicosor' Intrgni sqllns and som introns inlud rgions
tht a rsponsiblfor ontrolling g11 xpssionby dtrmining th lvls
nd timing of rrrssngr RNA prodution fronr rrarbygns.Ths ar non-
lrromosom ontaining fiv gns,
lmbinations of th gns (indiatd o d i n g u t f u r l t i o n a lp a r r s o [ g n m s .
llak and gray lins f<lr th two vn with omplt gnom squns'it is diffiult to ount a gnom's
rossing vr in a rgion f gnsausoding squnsar almost indistinguishalfrom othr s-
,s that ar similar nouglr quns.Th task is a ndl-in-t-haystak prolr ausof th low pro-
that thy
tr. potin lf oding squnsin th gnomsof man1.nrultillr"rlr orgnisms
158 uolutilltrlf th olltt,

(stimtd to aout 1.5% of hunran DNA; s Int11i1tional Human th fruit fLDroslphil mL

Gnom Squning onsortium 2001). Th triplt gnti od usd trmintion) has aout 20 r
mssngrRNA, ilr whi th ad jant nulrtids(odos)orrspond to thm omposd of a blol.
ah amino rrid of th prltirr nodd' and rtain triplts-rh sfor sparsr in htrohromatin
odls-stgnify t rrd of traIlslation,an sllggstandidtsfor squtls Th amount and squn
tht nldamino id squrrs.Th rrrrlotidsin th third plsitionsof among diffrnt spis.Fo
many odons do not fft th arino aid nodd, sitrsvral diffrnt mad up of fou short sa1
odons <lrrsptlndtl ir givn amino id (th tld is dgurcrt) (l-1.h thos f its distant rltiv,
2007; I-win 2008). If th sqnof e givn gntlmrgion is ompard . gnom siz (John and Mi
twrrdifft-ttspis,ir prdomirranof diffrnsat vrv thir1position 2007 ). Diffrnt kangaroo
an rval oding squns;in nnoding squnsno suh pattrn rvill fator of 1.6 |a rang simil
appar. Nvthlss,stiratsof gn nunrrs rmin unrtain for th of diffrns in th alnoun
gnoms of mr"rltillular organisms' and thsStin1ats1r ollstntlv ing V a r i a t i o n a m o n g s p i si n
updatd. larg diffrns in gnom(
Gnonr sizs o[ diffrnt typs of <lrgnismsdiffr rvidly otlr in th nttmrs (:rvalir-Smitlr1
numrs and lngtlrslf oding squnsand in th arount of ah typ of
nonoding squns(Tal 1). Totl gn ntlnlsgnrall1,orrlt with Trasposbl elemnts
th numrs nd sizs of intrlnsand th amount of intrgni DNA (Lynh Sstantial proportions of
ar-rdor.rry200; Lynh 2007). Tlr smll gnomsoi prokar,votslrav vry p a t s ( 2 0 0 b s s - 1 0k i l o a
littl spa twn adjant gns, and thy lak spliosomirlintrons, al- whol gnom than stlli
though thir oding squnsm1' intrruptd y mobil squnsthat ttble lemers (Ts)' whih
an spli thmslvsor-rtof tlr mssag(Lu,in 2008). Unillulr ukry- ations in th gnom (/r
ots witlr ompat gnoms'suh as yast' hv gnswith fw, small introns worm norbbditis le5
and littl itgniDNA' In ()ntrllst)th largstln()ms
of ukavotstrrd .s-10% of tlr total gnom
to hav larg introns and muh intrgni DNA (l,ynh and ony 200; humans th figur is about
Lynh 2007). onsortium 2001)' and st
aundans (Lynh and
Sval nrajor typs of T
REPE||VsQUNs lN NoNoDING REG|oNs
of organisms (Tal 2);
In addltion to fur-rtionalsqunssuh s pro()trs and nhanrs in- Within ah of th nlajor
volvd in th r:gulirtiontlf gn tivity ([,win 2008), nln<ldirrgrgins lf lassifid into diffrnt far
gnoms also inlud squnsthat sm lss likly to hv imptlrtant fun- ilar. Unlik th othr typ
tiorrs for th lls, spiallv sqllrrtyps that oLlr rpatdly in th last som mmrs with (
genom. W r,ill()nntrathr on th two rnajor <lmponntsof gntlms for thir transposition' u
of multillulr organisms arr]will not .lsrib othr typs of rpats,suh mutations in thir squn
as nrinistllitsand mirostllits(sB. hrlsw()rtht a|. |994|. n of omplt lmnt
othrs ar frgtnnts who
StIIit DNA human gnom is mad u;
Satllit DNA onsists of long tandm rrys of highly rpatd,n()ntran- Just as with satllits
sid squnslth r.rnitsin sulr arvs vary in lrrgtfrom fiv to s\.al latd spis an du t
h u n d r d a s p a i r s ( p ) n d a r u s u l l y r i h i n t l r a s sA a n d T . S t l l i t without major diffrn
DNA forms th n-rjor()mp()nnt>ihatroromti'tlr loks ot unrtsu- split from its rlativ D.
all dply staining matril visil in dividing lls' hromosomst usually aout thr tims as man
onntratd nar thir ntl:ollrsnl (in som spis)tloms (Jtlhn amounts of ths squn
and iklos 1988; lgin and Grwl 200). Tlry ltn rnak up astonishirrg nd on diloid rispi
amounts of th DNA o[ som <lrganisms.For anrpl,on hrorosom of th nrultipliation of a fv

n DNA; s Intrrrtional trtnan
. Th triplt gnti od usd y
nulotids(ods) orrsponlto
d, aIrl rfin triplts-th srop
]n suggstndidtsfor squns
ruiotidsin th third plsitions of
id nodd,sin svral diffrnt
id (th od is dcgrte)(Lvnlr
tvngnomrgion is ompard _
lf diftrnsat vry third position
ng squnsno suh pattrn will
nutnrsrmain unrtain for tlr
hsstimatsar onstantly bing
.ganislsdiffr widly
oth in th
l and in th amottnt of h ty-pof
t t-tt-lntrs grrrllllyorrIatwith
amollnt of intrgrriDNA (L1,nh
l gnomsof prokaryotshav vy
thy lak spliosomirlintrons' l-
rrttptdy rnlilsqunstht
( L w i n2 0 0 8 ) .U n i l l u l r u k r y -
' hav gnswith fw, small introns
largstgnomsof ukaryotstnd
i DNA (L1.nhnl onrr' ?00;
!oDING RGIoNs
h as prlmotrsand nhanrs in_
Lwin 2008), nn<;dingrgions of
lsslikly to hav important fun-
yps tht our rp:1tdlyin t
lwo 11ll()omponllts of grrlt-trs
sriothr typs f rpats,suh
' harlswortht l. |994).
rraYs of highly rpatd,nontran-
; vay in lngth frorn fiv to svral
. i hi n t a s s
A and T. Satllit
rohromti,tlr l<lksof unusu_
liding lls' hrotrrosms,usr::rlly
(in som spis)tloms (John
3).Thv an mak up astonishin
Fl nrpl,ln hrlt.tloslmof
uolution of the cenome I5e
th fruit f| Drosrryhil melogster (th X chromosottt, inolrvedin sx d.
trmination)has aout 20 million ass (M) of satllitDNA, aout half of
thm omposd of a blok of rpatd 359 p units. Funtional gns ar
sparsrin htroromatinthan in th rst of th gnonre(te echromti)'
Th amount and squnompositiorr of satllit DNA may vary widly
amongdiffrntspis.For instan,aout 4" of th DNA of D. uirilis is
md up of fotrr short satllit squns'whih ar quit diffrnt from
thosof its distant rIativD. mlogsterzlndontriut to its muh largr
gnomsiz (Jolrn and iklls 1988; Dros,rphil12 Gn<rmsonsotim
2007).Diffrnt kangaroo rat spis(Dipodmys) vary in gnom siz y a
fatorof 1.6 (a rang similar to that in mammals as a whl), mainly bus
of diffrnsin th amounts of thr short pats(John and Miklos 1988).
Variation among spisin tlr amount of stliit squnsn hus aus
larg diffrns in gnom siz, whih irr unrlatd to diffrns in gn
n u m r s( a v l i r - S m i t h1 9 8 5 ) .
Trsposb I lerts
Sustantialproportions of th gnoms of many spisonsist of largr r-
pats(200 ass-10 kilobass),with mor sattrddistriution aross th
whol gnom tlrirn satllit squns.Ths rpats ar largl trspos-
bl lnrcts(s), whih aIr insrt nw opis of thmslvsinto novl lo-
tions in th gnom (trsposition). | D. mlogster, the rrmatod
worm eorhbditis lgtts, and th pIant rbidopsis thli, aout
5-10% of th total gnomi DNA ontnt is mad up of Ts (Tabl 2). In
humansth figur is aout 45% (IntnationalHuman Gnom Squning
nsortium 2001)' and slntplants, suh as matze' hav vn highr T
andans(Lynh and onry 2003; Lynh 2007).
Svralmajor typs of T ar rognizalin th gnoms of a wid rang
of organisms (Tabl 2); ths diffr in thir transposition mhanisms.
Within ah of th major atgoris,th lmnts in givn spis an e
lassifidinto diffrnt familis, within whih th T squns ar vry sim-
.
ilar. Unlik th othr typs of rptitiv DNA' most familis inlud at
lastsom mmrs with on or mo grrsthat od for protins rquird
for thir trarrsposition,but ntany also h:rvmmrs with dlrions or othr
mutationsin tlrir squns.Of ths, some an tansposdin th prs-
n of omplt lmnts tlrat provid th transposition mahinry' whil
othrsar frgmr-rts whos tansposition is disald ntirly. uh of th
human gnom is mad up of suh fragrrrntsof Ts.
Just as with stllit squns'larg gnom-sizdiffrnstwn r-
latd spisan du to diffrnsin th amount of T-drivd DNA,
without majo diffrnsin gn numrs. For anrpl, D. melogster
split from its reltiv D. siltlls only out 3 million yrs ]out has
aout thr tinrs as many s (Viira et a|.202). Maiz has nruh largr
amountsof thssqunsthan its losstrlativs(Bruggmann t al. 2006),
and on diploid ri spishas doul th DNA ontnt of othrs ausof
th rrrultipliationof a fw T typs (Pigtlt l. 2006).
1 6 0 uoltil of t|l Ge

Tal 2. T ontnts f human and Drlso1lhlia gr.ro.rs (lu-grnalga)and pro|

mitohondria volvd mu.
Ts in th huran and D. mcllgstr uhromati gnOms
protoatria (Gra 1999
Numrof Nu-rr Latr ndosymiont v
Typ" opls of familis 7. of th gnom zoa (ma|aria parasits and
Hurns drivd from rd a.lga;prt
l-R lmnts 443x10r 100 8 mrs of th hrmop}
StNs 1 , 5 . 5xU 1 0 r 3 l-) hlroplast, whih has r
LINs 8 6 8x 1 0 ' 3 z0 ll. As wll as thir plasti
DNA lmnts 294x10t 60 -) still hav a vstigial nula
D. mlogstr ovr 40O gns)and thus
LR lmnts 682 49 /-.b J and th tru nular gno
LINs 486 27 0.97
DNA lmnts 404 20 0..'
GEN DUPLlAl
Slurcs:Humn dt:rfrln.r [trntilnirl
Humn GnlnrSequningonstlrtium
(200l); Drlslphilir ltirom Kaminkr t l. (2002). N{or frquntly, inras
bbruitils:LIt Irrlnts,[-lngTrmrnirl Rpat |.rrrts (lmntswith rpatd Nw gns aris from d
sqLlIls nd, iri th srllrintatiln,
at a|-r whih rplittl-rrlrrgh
rvrstrnsrip. gnoms (ihlr and Sar
tiln intlDNA lf n RNA lpyof thir DNA squ); SINs, StlrtItrsprsi
an oftn rognizd
Nul:rrlt-nrits (lrrrlrts<.500p 1ong,whih rplitthrotrglrn RNA intrmdit
tios |ose to thir prog
ut lk rpts:1ndrli, n LINs to prvid th t-tzyr nddfor thir rvrstrn-
siptin);I-lN, l-rrglntrsprsdNrrlarlmnts(similrto LR lmnts,ut suh arrays an thn hal
> lkb long r-rdlaking th trnrir-rl rpts);DNA lmrrts(lmntsthat rpliat twn two diffrnt m
withtlr:tus lf n RNA lntrmdiat, rrsullywith shlrtinvrtlrptsat ah nd). opis of th squnan
Som dupliatd array
vord larg amounts of l
Most Ts ar found in intrgniDNA r (to a lssrtnt)in introns. In gnS that nod th Rl
stlm rgions of th gnom' s n vry dnsly pakd, with multipl nism (riosoms) and t}
lmnts insrtd witin on anothr. Th portion of th D. mlogstr gns (Ranson t al. 200
lrtrohromatinthat irdjoinsth uhromatin is largly md up of matrial vantag' so that natural s
drivd fronr Ts (John and iklos 1988); th rgions around th n- pliatd gns. Givn n<
tromrs of . thli.l hrom()soms(!7right t al.200) and many intr- on of th dupliat gn
gni rgions in maiz (SanMigul t al. 1996) ar sirilar. On a dupliatd gn l
it and fixd in th p
protin-oding aility
GNo ADD||oNs
rmnant psudog'.vt
arly gnornsmust hav n vrv small, ut today's organisms hav larg or part of it. Nwly dt
nunrrs of gns with divrs funtions. An important' though infrqunt, quikly lost from gno
pross is addition lf gnomsor ]nsfrom othr organisms. Bati gu- ats mans that' aftr a
larly quir novl gns fr<lmothr atrial spis ()hmant al. 2000). tinguish from Partial du
ukaryot lls ontain orgarrllswith thir own gnoms,as wll s th nu- gnom is rognizaly d
lar gnoms.Grn plants and grn alga hav hloroplasts,and rd alga s q u n d t a t h a t p r o v i
hav similar plastids;plant and animal lls oth ontain rnitohondria(I,ynh tnt, dupliat rgions
2007). Th gnoms lf ths lrganllsr of prokaryot origin and aros arly th strths hav rturr
in th volutilnof ukaryots,whn a trialll was ngulfdintla ukary- mnts will, howvr' v
ot ll nd took up rsidn thr s an ndosynriont. Squnsof gns Somtims, howvr, t
from hl<lroplastgr-roms ar si-rilrto thOsof photosynthti yanotria for inativating mut
 uolutio of the Gome 161

cphiI gnons (blu.grnalgzr)and proaly volvd around 800 million yrs ago, whil
.-rhronrati!]n()ls mitohondria volvd muh arlir from a group of atria known as -
protoatria(Gay 1'999).
Numbr
,%
Ltr ndosymiont vnts hav also ourrd. Th apiomplxan roto-
of fmilis of te gnom
zoa (malaria parasitsand thir rltivs)and dinflgllatsontain plastids
drivdfrom rd alga;prsumly a protozon ll ngulfd a rd algal ll.
r00 8 mrs of th hromophyt group of alga also ontain a rd-algal-drivd
13 hloroplast, whih has rntly n shown to rprsl.lta whol rd algal
z0 ll.As wll as thir plastid gnom, mmrs of th ryptomond sugroup
60 .)
still hav a vstigial nular gnom drivd from th rd algal anstor (with
ovr 400 gns)arrd thus possssfour gnoms. inluding th mitohondri
49 2.65 and th tru nular gnom (Douglas t l. 2001).
27 0.97
20 0..'
GENE DUPL|AIoN AND GN Loss
umn (llt.t-t
S1uI-ring
nslrti um
(2002).
or frquntly, inrassin gn numrs om fronr gn dupliations.
l l R p t I m n t s ( l r l r t sw i t h r t c d
Nw gns aris from dupliations of parts of th gnom or of whol
. w h i h | i l r t h r r ' u h ( v r \ ( .t r ; l I t s r i -
gnoms(ihlr and Sankoff 2003; Lynh 2007). Singl gn dupliations
s q u n ) ;S I N s ' S h t l r t I t r s p r s l
: h r p l i t t h r l t r g h n R N A i n t r r r l d i t an oftn rognizd aus thy fquntly our as tndm duplic-
th nz1'm ndd for thtr vrs tan- tios |ose to thir prognitor gn (Baumgartn t al. 2003). Th sizs of
l m n t s ( s i r n i I rt L T R l r - n n t sl,l u t suh arrays n thn hang furthr y unqual rossing ovr. An hang
) N A l m n t s ( | l n t st l r a t p l i t twntwo diffrnt rnmrs of an array rts on dsndant with thr
vith short invrtd rpats t ah nd). opisf th squnand on with a singl opy (sFigur 2).
Som dupliatd arrays of gns probly volvd undr sltion that fa-
vord larg amounts of th gn prodttt' for ampl' th larg arrays of
A or (to a lssrtnt)in introns. In gnsthat nod th RNA omponnts of th protin translation mha-
vr,vdnslvpkd, with rnultipl nism (riosoms) and th xpandd aays of sonr instiid-rsistn
rh portion of th D. mlogster gns (Ranson t al' 2002). Oftn, howver, highr grr dosag has no ad-
lmatin is iarglv md up of matrial vantag'so that natural sltionwill not at to prsrvth funtions of du-
1 9 8 8 ) ;t h r g i o n s a r o u n d t h n - pliatdgns.Givn nough volutionary tim, an inativ mutant opy of
\X/rightt al. 2003) and many intr- on of th dupliat gns may om fixd in th spis y gnti drift.
1 9 9 6 ) a r s i m i l . on a dupliatd gn has lost its funtion, fLrrthrmutations may ou ln
it and fixd in th population y drift. vntully th gn may los its
protin-odingaility y mutations rating odons' and it thn oms a
rmnant psedlge;vntually deltions f gnti nltrial my rnrov all
rll, but today's orgnisms hav larg or part of it. Nwly dupliatd gns ar thrfor oftn ptd to
s. An irrrportant'tough infrqttnt, quikly lost fronr gnoms (Walsh 1995; Lynh 207). Loss of gn dupli-
rom oth orgnisms.Batriirrgu- atSmans that' aftr a tim' whol gnom dupliations ar diffiult to dis-
: t r i a sl p i s( o h m n t a l . 2 0 0 0 ) . tinguish from partial dupliations; in ithr sitution only a fration of th
i-rirown gnols'as w]l as th nu- gnomis rognizly dupliatd (IJ7onget a|.2002). \/ithornplt gnom
lgahv hloroplasts,and d lga squndata that provid larg gnom strthsof larly similar gn on.
lls oth ontin mitohlndri (Lvnh tnt,dupliat rgions lr rognizd,vn whn most of th gnswithin
of prokaryortlrigin :rnd aros arly th strths hv returnd to th singl-opy stat. Chromosom arrang-
tril ll was ngulfd into a ukary- mntswill, howvr, vntuallyras all tras of dupliation.
n r-rdosymiont. sqtll1sof gns Somtims' howvr' dupliatd gns may gain nrv funtions y mr-rtation
tosof photosynthtiyanoatria for inativating mutations our. For instn, an nzym that atalyzes a
162 uoltillt tlf th (}llll

givn rti<lnstp in a iohmial pthr,aymy gain nw irility tO at on Although it is not kn

th produt of th rtion and gnrata nw produt. S()mgns nod trolld, svral gnom fi
protins with two or mor funti<lns,and irrativ:rtingtrrutirti<lrrstht fft ination rat. O suh a
only on funtion may lad to survival of oth dupliats,ah with just ln of th frquny of G v,
of rh initil funtilns(Lvnh :rnd lnry 2003; Lvnh 2007|. This inrass man gnom th man Gl
th numr of funtional gns in th gnom. Familis of gns that hav ginally from aout 30o%
arisn by dupliation ar importtlt Omp()nntsf gnoms,and somtims onsortium 2001).Th (
hav many mmrsr,ithrlirtdkinds of funtions, suh as mammals' odo- o n o r d n t l y t w nr g
nt rptors (Momarts 2001) or plan diss.rsistangns (Baum- mor AT rih than odin;
gartn t al. 2003). Nw gr-rs ma1,lso form y fusion rlf parts of tw()gns tions (whr m()st mutatl
if part of th mssngrRNA of on gn is transrid ak into DNA nd lats to som tnt with tl
instd into th hotrrosorwithin or adiant to anothr gn (Lorlg squns'suh as pstrd
2001). Suh rvrstransription and transposition into th grrl-t an also tiularly intrstingau
rt nonftlntional lupliats wihorrt intrlns (1lrlssdpsldoglls) aus of th input of nrt
(Lynh 2007; l,win 2008). rh fat that thr is oftn
(Lynh 2007\. In th hum
tnt is highst for suh sitt
RoBINAloN RAs AND RG|oNAL DIFFRNS
high (lntrnational Hurnz
WIHlN GNoMs
and Pigana202),
An important influn On sm of th pattrns just dsrid is variation Anothr orrlat of r
in th rat of gnti romination (rossing ovr) aross th gnom' in high-omination r1
Thr is ssntilly no rossing lvr in htrohromatin. Romination Human Gnom Squn
frqunis alstl var,v withi uhromtin. Rorl.lirrationis fquntlv . thli (ltltrntitln
almost asnt naf th juntion with th ntromri htrohromatin ut \/rightt al. 2003) gn .
is ruh mr fqun irr oth parts f th horrslnr (B' harlsworth 8s t|7eopposir is foulr
e t a | . 1 9 9 4 ) . I n s o m s p i s ,s u h a s D r o s o p h i l a n d m a n y p l a n t s , r o m . Transposal-lmnt.
i n a t i o n i s s u p p r s s dn a r t h t i p s o f h r o n r o s l n r (sB . h a r l s w o t h t [ . D r o sop h il low-romi
1994), ut rats ris sharply nar th tlomrs in hurnn mals (Intrna- srtions than high-rom
tionl Hun-ranGnom Squning tlnsortium 2001). In . lgns,lvhih hav a mor ompl pat
has no wll-dfind ntromrs' romination mainly ours nar th low-romination AT-ril
hromosotn nds and rarlv in th ntal rgions of th hromosom romintion) rgions (I
a r m s ( B r n s t a l . 1 9 9 5 ) . I n m a n r m l i a n g n o m s ,t l r r i s l s o f i n - s a l tium 2001).
variation in rossovr rats, and vry lozrlizd hot spots, with unusually
h i g h r o m i n t i o n r t s ' a r s a t t r da r ( ) s st h g n r l m ( K a u p p l t a l '
2003; yrs t al. 2005). volutionary Fatorl
Sx hromsolns hav ptiularly striking rol.ninationdiifIrs.In and Organization
spiswith sparatsxsand highlv volvd mhnismsf s dtrmina-
tion (animirlslik humrrsand fruit t1is,and som plants), th s of l-rin-
FAoRs A A
IN A GENo
dividual is dtrrnindy gns on pair of s romosoms. Most oftn
fmirlshv two X hromsonrs,wlril mls hr,an frorn thir rothr Among elatd spis,u
and morphologially distintiv Y hromlsominhritd from thir mal lif (parasitism or intra
parnt (in ontrast, in irds arrd Lpidoptra, fmals ar th sx with th quird for survival and rl
norrmathinglrromosom pair). All or mlst of t Y hromosom dlsnot batri that nrak thir
ross ovr with th irnd is thrfor fftivlv inhitd as an asxual nrurhsmallr gn numl
.W
blok of DNA (D. harlsworth t al. 200.'). dsri th volutionary trm rdution lras
onsunsof this latr. smallr thn ithr at

hw1.mygi nW aility to t on
t a nW prldut. Sorrrgns nod
rnd inativatingnrutations that affet
of oth duplits,lr lvith just tln
rv 200; Lnh 2007). This inass
gnom.F.arnilisof gns that hav
nponntsof Pnoms,and solntims
of funtions,suh as nrammls' odor-
rlant disastrsistan gns (Baum-
l forrn y fusion of parts of two gns
:n is transridak into DNA and
or adjant to anothr gn (Long
ransositionir-rtoth gnom n lso
)Llt introns (prosstl psedogs)
REGloNAL DIFFRNs
pttrs just dsriLrdis vzrritiorr
(rossing ovr) aross th grrome.
in htrohromatin.Romintiln
ntin. Rorrrinatilnis frqulrtiy
.h ntromri htrohromarin ut
lf th hromosom (B. harlsworth
Droslphilnd manv plnts, rom-
ilromos<l-ts (B. hrlsworth t al.
tlomrsin hurn rnals (Intrna-
n s o t i u m2 0 0 1 ) . I . l c g
orinatiorrmainly ours nr th
ntrai rgins of th hromosom
lial gnoms,thr is also fin_sl
loalizd lrot spots, with unusually
ld aross th gnom (Kuppi t al.
;trikingrombintiondiffrrrs.In
volvd mhnismsof s dtrmina-
s, nd som plants)' t s of r.rin-
i of s hro-tosonts.Nzlost oftn
Lmals hv an from thir rothr
Olosominhritd fom thir mal
optr' fnrirlsr th s r,vith th
most of th Y hromosom dos not
fftivlylnhritd as an asul
2005). W dsith voltrti<lnry
t,llution of th Geome 16
Although it is rrot knowlr how romintion-rat diffrns ar on-
trolld' svralgnom fturshav n found to orrlat with th rom-
ination rat. on suh assoiation is as ompositiln (prssdin trms
of th frquny of G vrsus AT irspirs, mntiond rlir). In th hu-
man gnom tlr man G ontrrtis aout 41",ut th ontnt varis r-
gionally from about " to 657" (IntrnationlHumn Gnom Squning
onsotium 200l). Th G ontnts of gns and intrgni squnsvry
onordantly twn rgions. Altlrough nonoding DNA trrdsto nrh
mor AT rih than oding squns'th GC ontnt of third oding posi-
tins (whr most mutations do not afft protin squn)usually or-
latsto som tnt with that of adjnt nonoding squns.Nonfuntional
squns'suh as psudognsand dftivtransposallmnts,ar par-
tiularly intrstingusthy rval th G ontnt that will ist purly
ausf th ir-rputof mutations (without rraturalsltion),nd thy rtlt
th fat that thr is oftn a mutation prssur toward AT and away from G
(Lynh 2007)' I th human, . lgs, and Drosopbil genoms, GC on-
tent is highst for suh sits in hromosom rgions whr rossovrrats a
high (IntrnationalHuman Gnonr Squning onsortium 2001; Marais
and Piganau 2002).
Anothr orrlat of romirrarior-ris irrtron lrrgrh;this tnds to low
in high-rominationrgions (omron ar-rdKritmn 2000; Intrntionl
Human Gnom Squningonsortium 2001). In addition, in humans and
. tblitz (lntrnational uman Gnom Squning onsotium 2001;
!riglit t al. 2003) gn dnsitis ar highst in ths rgions, ut tn . l-
gns te opposit is found (. lgs Gnom Squning Projt 1999).
Transposal.lmntdnsitis also var rgionally within gnoms. In
Drosophil low_rominationrgions hv muh highr numrs of T in.
srtionsthan high-romination gions (Kaminkr et aL.202|. Humans
s 'w h i h hav a mor ompl pattn: LIN lmnts (Tal 2) ar assoiatdwith
iow-rominationA.rih rgions, nd SIN lmntswith G-rih (high-
romination)rgions (Intrntionaluman Gnom Squning onsor-
t i u m2 0 0 1 ) .
volutionary Fators that Contrl Gnom Siz
and Organization
FAoRs HA AFF E NUMER oF GNs
lN A GNo
Among rlatd spis'unusually small gnomsar assoiatdwith ways of
lif (parasitism or intrllular symiosis) whr most of th sours r.
quird for survivl arrd rprodution r providd y th host. Fo ampl,
atria that nrk their living as intrallular symionts or parasits hav
muh smallr 8n numrs than fr-living rltivs (oran 2003). Mor
trm rdution hs ourrd in organll gnorrrs, whih ar muh
smllrthan ithr atrialor nular gnoms;thy arry only gnsndd
164 uolution rfth Gom

for thir own pliation and prtltin synthsisand flr a fw spializdor- th mutation rat pr nul
ganllar funtions. any gns ssntial for organll fr-rntilns hav n atria. For ampl, in }
trnsfrrd to th nulus (Gray 1999). ongoing gn trnsfr hs n d- drashov 2003). Givn th n
ttd frorn plant organllgnoms,s wll as transfrsthi1tratnonfun- man gnom, this translat
tional psudognsof orgnllgnsin th nular gnomsof many spis protin squnof aout
( G r a y 1 9 9 9 ;B n s a s s o n t a l ' 2 0 0 1 ) . tion, and around 80% of tl
Gnome-siz volution involvs svrl prssurs.Thr may advan- nt mutation rat to dlt
tags lf smallr gnom siz, ausir rdud gnom n rpliat fastr, largr numr of dltrro
allowing fastr ll division; this arr oLlr through dltions of gns that th nonoding portions of 1
ar no longr ndd, or of intrgni and introni squns.Th finding th stimat of th human
that ompat gnoms oftn hav ovrlapping gns, in whih th sam H i g h d l t r i o u sm u t t i
strth of DNA is transridin two diffrnt rading frms (Douglas t al. (Crow 199) ma limit p.
2001; l,win 2008), suggststhat sltion fvord siz rdutiorr in ths Thortial modls of th
ass.But loss of som squnsmay sltivlynutral, so that dltions at that mutatinal load
an fixd y gnti drift, as outlind arlir for rdundant gn dupli- 2 0 0 0 ) . H o w v r , t h r i s l
ats. In nonrptitiv squnsthat ar unonstraind y sltion, th mhanisms, whih slow
amount of DNA tnds to dras ovr tim aus rpliation rnistaks sours. Th ourrno
sm to aus small dltions rnor oftn thn insrtions of ases (Grgory rats to zro, ut in sxua
2004; Lynh 2007).If sltivonstraints on gns r rmovd, thy v'.ill twn gns' alllsthat
thn gradually dlin in siz ovr tim s mor dltionsthan insrtions - atd with favoral mtat
om fid y gnti drift. Prtlaly oth sltiv and nonsltiv pr- l i t t l d v a n t a g . u t t i o r
s s sh v no p r a t i n g . ar thus likly to rflt tl
rduing th mutation rat
load (Snigowski t al. 20t
MUA|oN RAES AND H NUMBRS oF GNs
I n o n t r a s t ,i n r g n i s m
ost mutations with ffts on an organism's funtions rdu survival or fftiv gnti rom
frtility (fitnss,for short). Unlss sltilnis vry wak, thr will a al- s P i s . n a l l l t h t i n
an twnth mutational input of nw dltriousvariants into a popul- soiatd with any favora
tion and thir liminatiln y sltion. Th prsn of ths muttions qny as th mutation s1
rdus avragfitnss,lmpardwith a mutation-frpopulatin; this mu- m u t i o n a t st h a n i n s }
ttiol lod depedson th totl numr lf nw dltriousmutations that ig ad is obsrvd in al
aris pr lndividual ah gnration,suh that th n!]ativof th natural log- sltion prssurs(Snig
rithm of th man fitnssof a poprrlatilnis qual to th lad (row 199|.
For a givn mutation rat pr gn, thr will thus a gratr load if th GNoM s|z |N
numr of gns in a gnom is larg. RNA gnoms lak rror-orrting oF NoNoD|NG t
mehnisms and hav trmly high mutatilnrts pr gnom pr rplia-
tion, dspit thir small gnom siz: around 1 nw mutation pr rpliatiln Gnom-siz diffrns
yl for th influnza virus (Drak t al. 1998). Suh gnomsannot' thr. v o l v l a r g d i f f r n si r
for, hav larg numrs of gns, austhir {itnsswoLrld tlm vry DNA, with only minor
low. DNA-asd gnoms hav rror-orrtingmhanisms that giv thm Lynh 2007). In som a
muh lowr, ut not zro' muttion rats; the. oli !]nomis stimtdto in lif history and olo
hav a mutation rat of around 10_r0pr nulotidpr rpliation and a mu- hav mor satllit and ot
tation rt pr gnom of only 0.0025 (Drak t al. 1998). ons' suggstingthat th
ultillular lrganismshav larg gn numrs nd many ll divisions nutral with rsptto s
ah gnration.Bausrrr Orrtionlrn nvr e prft,and th muta- fors (lik th balan
tion rat pr nulotid pr ll division is quit similar to that in atria, lir) provids a plausibl

ynthsisand for a fw spializd or-
ial for organllfuntions hav n
). ongoing gn trrlsfr has n d.
; rvllirstransfrstlrt at nonfun_
l th nLrlar]nollls of mny spis
ral prssurs.Thr may a.lvan-
r rdrr.^d 8nom an rplit tstr,
lur through dltilnsof grrsthat
and intror-risquns.Th tinding
rlapping gns, in whih th sam
iffrntradingfranrs(Douglas t al.
tiorr fvord siz rdution irr ths
l sltivlyoeutfell,so that dltions
r d a r l i rf o r r d u n d a n tg n d u p l i -
ar unonstraindy sltitln, th
'r tinl bausrpliation
ristaks
n thn insrtionsf ss (Grgory
llnts on gnsr rmovd' thy vyill
as mor dltionsthan insrtions -
otlr sltiv:ttl{ntlnsltir,pro-
UMBRs oF GNs
ganis'sfuntions rdu survivl or
tion is vry wak, thr will a l-
w dltriousvaril-ltsinto a ptlpr'lla-
. n . h p r s n t l f t h s m u t i l t i o n s
1l-ltation-fr population this mu.
r of nw dltrilusmutations that
:h tht th rrgativof th naturl log-
i o n i s q u a lt o t h l o d ( r o w 1 9 9 \ '
r 'ill thus gratr lol if h
. R N g n o m sl a . . k r r o r - o r r
luttion ratspr gnom pr rplia-
.ound 1 nW muttior-r pr rplition
t. 1998).Suh grromsannot, thr-
aus thir fitnsswould bor vry
orrtingmhanismsthat giv thm
s;th . oli gn()lis stimatd to
r nr.r[otid pr rplition :rn{ mu-
Drakt al. 1998).
n numbrs and nrany ll divisions
)n an nvr prft,nd th mllt-
n is quit similr to tht irr irtria,
uoltio of th Gnom 165
th mutation rt pr nulotid sit pr gnration is muh highr than in
batria.For ampl, in humans it is stimatd to e aout 2x10 8 (Kon-
draslrov2003)' Givn th numr of sitsthat od for amino aids in th hu-
man gnom, this translarsirrto a mutation at p 8nom fo hngs in
protin squnof aout 0.82 nw ltlttions pr individual h gnra.
tion, and around 80" of ths ar likly to slightly dltrious,yilding a
nt mutation rat to dltriusamino aid mutations of aout 0.66. An vn
largrnumr of dltriousmuttions is likly to ilusd by muttions in
th nonoding portions of tlr human gnom (Asthna et aI.207),ringing
th stimatof th human dltriousmutation rat to wll aov 1.
igh dltiousmutation ats and th rsulting low population fitnsss
(row 1993) may limit possil gnom sizs for multillular oganisms.
Thortilmodls of th volution of grrsthat orrtrol mutatiol-lrats indi-
at that mutational load favors a rdud mutation rat (Snigowski t al.
2 0 0 0 ) .H o w v r . t h i s a n o p p o s i n g p r s s u r :t h o s t s o f r r o - o r r t i n g
mhanisms,whih slow down DNA rpliation and onsum llular r-
sollrs.Th ourn of irdvantagousmutations lso disfv<lrsrduing
rats to zro' ut in sxually rproduing spis with gnti rombination
twngns, allls that inrs th mr'rttionrat will not rmain assoi-
td with favoral mutations that thy happn to indu' and so thy gain
littl dvarrtag. Mutati<ln rts of sully rproduing lrighr ukaots
ar thus likly t rflt th volutionary lan att:rindwhn th ost of
rduingth mutation rat quals th advantag of rduing th mutational
l o a d ( S n i g o w s ki t a l . 2 0 0 0 ) '
In ontrast,in oganismstht produ largly without any volutiorrarily
fftiv gnti romirrtirn,suh as asual and highly inrding
spis,an alll that inrassth mutation rat will rmain prmanntly as-
soiatdwith any favoralutations it auss,and will thus inrasin fr-
queny as th mutation sprads in th population. This may promot highr
mutation rats than in sual spis.Tlris is an xanrpl of geti hithhik-
ig and is osrvd in atrial populations that ar sujtto intns novl
sletionprssus(Snigowskit al. 2000).
GNoM s|z lN RLA|oN o H AoUN
t i n g
oF NoNoD|NG DNA
Gnom.siz diffnsamong rlatd taa of higher ukryots oftn in-
volv larg diftrns irr amounts lf highly rpatd, nonoding satliit
DNA, with orrly minor diffrns in gn numr (avalir-Smith 1985;
Lynh 2007). In som ass DNA amounts larly orrlt with diffrns
in lif history arrd ology. For mpl, slowly dvloping spis tnd to
havmor satllitand othr typs of nrrodingDNA tl.rrifastr-dvlping
ons, suggsting that th siz of th nonoding portion of th gnm is not
nutralwith rsptto sltion(avalir-Smith1985). A modl of opposing
fos (lik th alan twn proofraling and mutation dsrid ar-
lir) provids a plausibl and tstablxplarrationof ths pttrns. Svrl
l66 ullutilttof th ot

prossstlrt invo|v rr()rsin DNA rplition irrrdrtnqual rossin!]ovr tistial orrtions for diff.
n amplitr, rril\rsb multipls lf rvhllrptdunits, s thr r fors this is not ptd if thr.
tht inras opy numr' somtims adding DNA to th gnonrof an in- from salamarrdrs and n
dividrrl;gntidrift an tlrn aus it to sprad to fixation (B. halswlrth Johnston 1992).
t l. ] 994;' Lnc 2007). Sltiln against high opy nurr will oftn op- an w s th footprir
.
pos this pross' ut suh sltion is last fftiv in spiswith small sizs, as assumd rlir?
fftivpopultiln siz (Kimr.rr198). This suggststht thr should onntration in rgions o
rltion twnth fftivsiz f a spis (whilr an stimatlfrol w a n . T h r g u m n ti l r \ '
its lr.l of DNA sqlln .livsity;11,n2007) and th ontiutirln oi ray undrgos unqual ro
rptitiv squnsttl its gnom siz. Brod omparisons aross diffnt pliations). Unequal hz
taa suPport this prdition (l-ynh 2007). Hor'vvr,spissuh as D' uir- opis in th array. ut it i
ilis and D. mllgstr diifer onsidrably in thir mount of rptitiv r is nithr advantagous
DNA u sm to hav similar fftiv opulation sizs (Drrlsophil 72 spis to om fixd for
Gnoms onstlrtium 207I. his suggsts tht othr fators must plav rquirs multipl opis, a r
a ll. aftr rmain singl opy (t
A likli" disadvantagof a larg anlount of stllitDNA is a slolvr dvl- opis an lost again y
opnlllt tim, sin ll ]ir,isionswill slow lf it taks longr to pliat that afft array sizs, th
gn()mswith larg DNA nltunts(Cavalir-Smith1985). \Whnrapid dvl- a tandm arry thus produ
opmnt is important, th alan is tippd towrd lowr opy numr, but i{ ity ovr tim that th pop
slow dvlopmnt is not disfavord y sltion' sltion against th au- ially if unqual xhang
multin of rptitivDNA ,ill wakr, nd sr.rlr squnsan aLlmLl- individuals with high rra
l t ( F i g t r r 4 ) . T h u s l a r g D N A o n t n t d < l sn o t n s s a r i l yi m p h , t l r r rsulrs. Larg rays aris
sltiotlhs favord nroDNA (Pagland Jllrnsttrn1992'B. hirrlsworth (B. Chalsworth t a|. \99,
t al. 1994). low-romination gnomi
Of ours, it might somtilns advantagotlsto hav mor nonoding for that pushs rpat n
DNA. For instan' mor DNA pr nulus might nal organisms with for that ats in th ppos
larg lls to mov RNA mor rapidly into th vtoplasm (vali-Snrith No funtional advanta
1985). l. an on distingtrislrtwn ths possiilitis? pritrrnts DNA squns is tlrus nt
th altr th i.rrountof htrohrolnatin in D' rclogLlstrhardly rrfft thir distributin within g
viailitr, lr dvlopmnt tilr-r'suggstingtlrat larg hangs in opy num- latd spis oftn diffr
r s ( m i I l i o n s o f a s s )h r ' a t m o s t v r y s l i g t a d v a n t a g so r d i s d v a n - quns' whih also strong
tags (John and Miklos 1988). Vry slight fitnss diffrnsan, howvr, ]v{iklos 1988; Drosophil
importnt in vllution, so this is not onlusiv' In omparativ tsts dos not man that satlli
tht xnrin dvlopmnt tils and gnom sizs irross diffrnt spis, Drosopbil, portions of t}
th bln rrrodlprdits thilt rlationship will flund vn ftr st..t- mosomal hvior at lnlo
volvd in ntromr [un
probaly did not aus t}
insrtion5,
5ma|| in|uding -..--->
ratdditions
st|lit
mor likly, ths arrays v
insrtis,
|rge i|uding
s lir and wr latr o-opt
small lrge
genome genome RANsPosAL

<- prssur
undrmUttiOn
dltions highostsof rpling
DNA ost typs of transposa
<- fvord
s|tivIy deletins slowgrowtfl t h m s l v si n t o n w p o s i t
<--UPU3| rossing
ovrwithgeneti
drifl nal opy. Ths transpos
m n t n u m r so v r t i m u
Figur 4. Opplsing fors atir.rgOn gnom.siz vrlIution. For ampl e, tn D. mI

liationand unqual rosslng ovf
rptdunits, so thr ar fors
tding DNA to th gnom of an in-
sprlto fition (B. hrlswoth
lst high op numr wil| oftn op-
'ast fftivin spiswitlr
small
This suggststht thr should
lis(rvhihn estimatd fom
nh 2007) and th ontriution of
Brod ompisons ilross diffnt
7).owvr, spissulr
as D. uir-
aly i thi amount of rptitiv
population sizs (l)ros-lphil 12
.sts that otlrr fators must pla
t of stllitDNA is a slowr dvl-
siow if it t:rkslongr rl repliat
i r - S m i t h1 9 t l 5 ) W
. hn rpid dvl-
l towird lou,r opy nun-rr,lrut if
ltion,sltionagainst th au-
r, and sulrsqunsI1aulnu-
lt dos not nssarily imply that
rdJlhrrston1992:B. hlswoth
ltltgousto hl, nror nonoding
:lusrnight nal organisms with
n t o t y t o p l a s m( a v l i r - S m i t h
ln ths possiilitis? priments
n in D. mllgstarhrdly afft
g tht larg hangs in opy num-
' r y s l i g h t a d v a n t g so r
disadvn-
lt fitnssdiffrnsan, howvr,
t o n l u s i v .l n o l n p r t i v t s t s
t o m s i z s r o s sd i f f r l r rs p
nship will found vn ftr sta-
dditions--------->
+ lrge
- genome
highosts
ofrpling
DNA
slowgrwth
-siz volutin.
uoltio of the Geom 167
tistialorrtions for diffrns in nular volum and ll siz, whrs
this is not exptd if thr is an advntag of more DNA. orrtd data
from salarnandrs and nWts still show a signifiant rlation (Pagl and
Johnston 1992).
an w s th footprint lf a mutational prssur that inrass rray
sizs,as assumd arli? Th distriution of stllit DNA, with a strong
onntration in rgions of infrqunt gnti romination, suggsts that
w an. Th rgumnt involvs a simpl modl in whih a tandm rPat ar-
ray undrg<lsunqual rossing ovr (plaind arlir for tandm gn du-
pliations).Unqrral xhang dos not hang avrag nurrs of unit
pisin th array, t it inrassth rpat numr rang. If rpt nunr-
r is nithradvantagousno disadvantagous'gnti drift may irusth
spisto bom fid for just a singl opy. Baus unqual rossing ov
rquismultipl opis, a rgion that rahsth singl-opystatwill thr-
aftr remain singl opy (unlssa nw dupliation ours). Howvr, xtra
opisan lst agin y unqual hangs.In th asnof othr fors
that afft array sizs' th omind ffrs of drift and unqual xhang on
a tandm aray thus produ an asymmtry:thr is an inrasing proail.
ity ovr tim that th population finds itslf with just a singl opy' sp_
ially if unqual hang is frqunt. If thr is som sltion gainst
individuals with high aray sizs, an quilirium disrriution of array sizs
rsults. Lag arrays aris only if rossing ovr is xtrmly infrqunt
(B. harlsworth t aI. 7994). Th lsrvdssoiation of satllit rrys with
low-rominationgnomi rgions thrfor suppots th hypothsizd wak
for that pushs rpat numrs upwd; w s its ffts only whn th
for that ats in th opposit dirtion (unqual rossing ovr) is rmovd.
No funtionl dvantg to hving larg loks of tandrnly patd
DNA squns is thus nssary in ordr to plain th main faturs of
thir distriutionwithin gnomsand among diffntspis.Morovr, -
latd spis oftn diffr gratly in th typs and amounts of satllit s-
quns'whih also strongly suggststhat no sltionis involvd (John and
Miklos 1988; Drosophil 12 Gnoms onsortium 2007). Howvr, this
dosnot man that satllitsnvr hv nssaryfntions in organisms' In
i s , DrosophiI, portions of th htrohromatin ar ssntialfor orrt hro-
mosomal havior at miosis, and human o-satllitsqunssm to in-
volvd in ntromr funtin (Sullivan et al.2001). But ths funtions
proaly did not as th vlution of ths squnsin th first pla;
mor likly, thsarrays wr first stalishdy th prosssmodld ar_
lir and wr latr o-ontd for ll funtions.
RANsPosAL LEMNs
Most typs of transposal lmnts (Ts) (Tal 2) can insrt opis of
thmslvsinto nW positions ir-rth grrom' oftn without lss of th origi.
nal opy. Ths transpositions also rat a prssur toward inrasd l-
mnt nums ovr tim unlss som othr fors kep T numrs in hk.
For ampl, in D. mlogstr it is stimatd tht on avrag t last on
168 urlltilnof th Gom

nw T is addd to a haploid uhromati gnom vry 10 gnrationsin

rarrngmnts hav fw
laoratory onditions whr sltionis rlaxd (asid t al. 2000). Thr
baus of gnti drift, l
ar aout 1,000 fairly omplt Ts in th gnom (Kaminkr t al. 2002), so
mosom strutur. BLlt n
th man transpositilnrt pr pltniallyativ lmnt is aout 10 a pr
highly dltrious:many J
gnration.
arngmnrs (glmi
\/hatrgulatsth aundan of Ts in th gnom? Thr
ar two on- 2003). If this is an impo
trasting viws (B. harlsworth t al. 1994). First, Ts ould maintaind
prdits that Ts should
aus thy ar advantaglusfo thir host organisms; for instan' thy
omination rats,sin
might oasionally indu favorl mutations y insrting into gnsor rg-
mnts and will thrfor
ulatory squns.A prolm with this id is that insrtions mostly aus
topi xhangs. As m
harmful, not advantagous' ffts, disrupting oding or rgulatory s-
proportion lf th Y hr
quns. Many known spontanous dltrious mutations' inluding som
quns (B. harlswortl
human disas mutations' r du t insrtions into oding squns(os.
planations for this pttr
trtag and Kazazian 2001). Suh strongly dltriousmutations are usuallv
of th gnom with rdu
rapidly liminatd from populations, and s mntiond arlir, it is indd
s m y s o m t i m 5i n
osrvd that in th gnom squnsof highr organisms Ts ar largly
mnts' and sltionwill
absnt from lding squns'indiating that insrtions thr ar harmful
T-drivd squnsap
(Intrnational Human Gnom Squnirrgonsortium 200l ; Kaminkr
or onfr lthr nf]ts
t a|.202; \/rightt al. 2003). F.urthrvidnthat T movmnt is disad-
satllit squns,hw
vantagous to th host is that marry spis,suh as yast' th mold Ne-
tion pomotd th initia
rosport, maiz, and Drosophil' ae mhnisms that rstrit th rat of
tory or vn oding fun
movmnt of Ts (Slkr2002; Aavin et aI.27).orovr, th viw that
w r i n s r t di n t t l t h i 1
T insrtions ar advntagousfor thir hosts' futur volution rsts on th
ida that th long.trm survival of a spisis nhand y th psnof
Ts, and it dos not plain how Ts om stalishdin th first pla. Asslv E A
Sond, Ts ould maintaind y thir aility to sprad by slf-rpliation sIz DIFFRN
within th gnom, oupld with transmissiln of nw rpisto th offspring
A | t h o u g h t h r a t | :i
during sual rprdution, vn if thy usually hav hrmful fftson thir
D. mllgasler sms l
hosts (th selfish DN hypothsis).on this viw T aundan will dpnd
roughly vry 10 gnra
on th alan twntransposition that aussinrasdopy numr vr-
tionary timslsif no
sus sltion against individuals with larg opy numbrs (B. harlsworth
tansposition stimatd
et a|. 7994; Lynh 2007). This alan implis tht th rdution in fitnss
D r o sop b il gnratiOns
ausd y n lmnt insrtion musr on avragqual th han that an l.
danes among rlatd s
mnt transposs. stimats of this rat ar around 10_apr gnrtion in
arlir, ar thus radily
Drosophil, showing that this sltionprssurmust wak. This hs n
somtims rlad. An
tstd furthr in D. mlogstery dtrmininglmntpositions on th gi-
whih n autosom
ant hromosoms of larval salivary glands. Th frqunisof T insrtions
yrs ago; this has sin
in th population ar narly always vry low at a givn pla on th hromo-
this gnom rgion to st
som, again implying that insrtionsar disadvantagous.Th strngthlf s-
ur in Droso1lhil ma|e
ltion an infrrd from th frqunis,nd it agrswith th stimatd
sity of T insrtions int
transposition rat (B. harlsworth t l. 1994|.
som spis (.g.,hurna
\/hat is th natur of this sltion? \ihav alrady disussd sltion
(Tal 2), and at many
against larg gnom siz, and harmful ffts of transpositions on th host. q u n i s( o r v n i n a l l j
Anothr disdvantag oms from r:ossingovr twn lmnts in diffrnt
r n g ,w i t h t h | o w i n s r t i
gnomi loations (ecto|li hge), This auss hromosom invrsions, .Why
ar thr suh l
transloations,dltions,and dupliations(B. harlswortht al. 1994). Som
lrgr amounts of int
 tlolutio of th Gom 1,6L)

ltl gnomvry 10 gr-rrations in rarrangmnts hav fw or no harmful ffts and n sprad in populations
rlad(asid t al. 2000). h
bausof gnti drifl, thry orrrriuting to volutionary h:rr-rg in lrro-
gnom(Karninkrt al. 2002), so
mosomstrutur. But many rarrangmnts disrupt gn funtions nd r
Llly tiv lmnt is alut 10_apr
highlydltrious:nrany humn gntidisssr ausdy hromosom r-
rrangmnts(glmi disclss),inluding mny ns (Diningr t l.
in th gnom? Thr i1two on.
2003).If this is n importalrt disdvurntg t having high T aundan, it
94). First, Ts ould maintaind
. host orgarrisrs; prditstht Ts sould altmulat mainly in gntrmi gions rvith low -
for instrr. thy
ominrtionrts' sin tlry ar tlr lss likly tl us harnrful rarrang-
rtiorrsy insrtinginto gnsOr rg-
mntsnd will thrfor not rmovd y sltion against th produts o{
ida is tat insrtions f]lostly atls
topixhangs'As mntiltldarli, this is oftn tru' For anrpl, a larg
srllting oding or gulato s_
pportion of tlr Y hromlsms of many spis onsists of -drivd s_
triousmutations' inluding som
quns(B. harlsworth t a|. |994|. Howvr, thr ar othr possil x-
srtionsinto oding squns (os-
planatinsfor this pattrn' srrhas th dud ffiiny of sltirrin rgitlns
|y dltioLrs muttions r usuall1,
of th grromwitlr rdud rorintion(sltr disr"rssiorr).
d as mntiondarlir, it is inded
Ts miry somtimsindu favoralmutations or hromosom rarrang-
f highr organisms Ts ar largly
mnts'and sltionwill thrl sprad thm throughout th plpulation.Son-r
g tt irrsrtionsther r harmful
-drivedsqunspp21rto rgultgn tivity (Diningt l. 200)
ing onsortium 2001; Kamirrkr
or onfr othr nfitson thir hosts (Brookfild 2003). As w argud for
:vidnthat T movmnt is disad-
satllitsquns'howvr, a prsnt-daynfitdos not impl,v that sl-
lis, suh as st' rh nrold Nzl-
tion prlmotdtlr initial sprd of tlrssqtlns.lr is possil tht rgul_
lhanisms th:lt rstrit h rat of
tory or r,nodin8 funtions hav n aquid by suh Ts long aftr tlry
r aL.2007). orovr, t viw tat
wrinsrtdinto thir Prsnt loations and fid in th population.
hosts' futr:rl.o]utionsts on th
is is nhandy th prsn of
:om stalishedin th Iirst pla. MAssIV AUULA|oN AND PoPULA|oN
ir ailit1'to sprerd y slf-rpliation sIz DIFFRNs
ssion of nw opis to th offspring
Although th rat of inrs of Ts y trnsposition in spis suh as
rsuallyhav harmful ff:tson thir
D. melnogsle/smslow (on nw lmnt in th haploid gnom lmnt
h i s v i r , , T a u n d a n w i l l d p n d
oughlvvry 10 gnations)'opl,numbr lvill inasrapidl,vovr volu-
t attssinrasdopy rrumrvr-
tionarytimsalsif no opplsingfo oprts:for xlmpl,with th rat of
g opy numrs (B. lrarlsworth
transpositilnstimatd arlir, a tnfold inras woLlld tak only 2,000
mplis that th rdution in fitnss
Drosopbil gnrirtiotls,lss tlran 2,000 yars. Lrg diffrnsin T Lrrr-
vrgqual tl-rhan tht an l-
dansamong rltd spis,s in th maiz and ri ampls dsrid
r a r o u n d l 0 - a p r g n r a t i o ni n
arlir,a thus radily plial if sltion prssus against thm r
:ssurmust wak. Thls has r
smtimesrlad. Arrothr ampl is provild Droslphil mirnd, itl
:mininglmntpositions on th gi.
whih art autos()nlm attahd to th Y hromosom out 1 million
ls. Th frqunisof T insrtions
yrars ago; this hs sin n transmittd only from fthr to son, ausing
oW at a givn pla olr t hrt>mo-
this gnom gion to stop romining usrrmirrtiondos not o-
lisadvantagos. Th strngthof s-
ur in Drr,,sophil ma|s.h rgion has sin thn aumulatd a high dn-
's,and it agrswith th stimatd
sityof T insrtionsinto nonoding squns(Bahtrog 2003)' Gnoms of
19941.
somspis(.g.,lrumns arrd miz) lrva vry hig ovrIlT aundarr
V lrv alrady disussd sltion
(abl2), and at many hromosomal sits, Ts may b prsllt at high f-
[ftsof transpositions on th host.
qunis(or vn in all individuals) (Ostrtag and Kazazian 2001)' ontrast-
g ovr btwnlmnts irr diffrent
ing lvirh th low instilnfrqunisin Drossphil.
h i s u s s h r < l m s o n tin v r s i o l r s ,
\fhy ar thr suh larg diffrnstwn spis?Prhaps th ruh
(B. harlswortht al. 1994). Som
largr nrounts of intrgni DNA and largr introns in humans than in
1 7 0 uolutio of tb Geom

Drosophil 1nthat insrtions hv lss risk of ausing harmful mutations Gns irr gnonrr.gion
(although this would not tu if intrgnirgions' squnsoftn ontrol similar volutionry pros
gen ativity). A mor intrstingpossiility dpnds on th waknss of s- ls of adaptation. his ma
ltion against ah nW T insrtion. If Ts rarly is from a sit, gnti omirring hrotnosomal rt
drift an somtims lad t a sit oming irvrsilyfid for an lmnt. (ph:rpslso in pIt Lls
In spis with low population sizs, suh as humans, sltion against l- strikingly illustrr th
mnts is lss fftivin opposing gnti drift' and this possily allows th ( D . h r l s w o r t h t a i . 2 0
prssrrrfor lmnt uildup to prdominat (Lynh 2007I. Nw insrtions s o m g n s ' t r ' i n gr h
into lady_upidsits wold flot prvntdy sltion(thrwould pai, ut tlr Y laks n)()st(
e no diret hrmful ftts,just a slightly gratr risk of topi xhang), th gnti informtion on
and so lmnts ould snowall ovr tim. This may aount for th fixd l- nonr()mininghromoso
mnts in mnrmalian gnoms' ut rrot for th vry high T aundan in p d t h i r i l i t yr o m a i n
maize, a spiswhos high DNA squnvalaility indiats a larg pop- fastr than tir -linkd
ulation siz (Tenaillon t al. 2002I. Romination in maiz may largly d i s d v e n g o u sr . e r i e r l t si
onfind to gns' so that Ts in intgni squns ould not rmovd gns thn lik normal gn
y topi hang. Thus lmnts would uild up twn gns' as o. s s o f s . l i n k dg n si n
srvd (Sanigul t a|. 1.996)'owvr, th austion ould th othr ( F i d o l i s s o n n d l l g n.
way round: Ts may simply romi lss than l.lostsquns. gns addd to th Y hro
onfirm hetthis is du tll i
mutations (Bartollrnand
H VoLUIoNARY oNsoUNs oF GNI
Similar ut mildrfft
RoMINA|oN AND PoPULAloN s|zE
gions with diffntroml
SltionagainstTs is only on Way in whih small population siz and lak odons that nd irr G or
of romination an importnt for gnom volution. Romination ontnt of third oding pos
rts play important ols in numous voluionryprosss'spiallyth o th djant r-rontrdi
volutionary advantags and disadvantagsof sual rprodution. Svral gs, and ' thlitl, ter
diffrns among gnom rgions r onntd with romination rat flts th ation of sl
diffrns' nd th ffts of low ronrination rts ar similar to thos of fastr rrd/or mor aurt
small fftivpopultion siz (Grdo arrd Chalsworth 2001). tin squir(Akasi t al
Sx and rombintiorr llow sltion to at indpndntly on diffrnt sits is' horvvr, vr1' small o
within th gnom. This inrassth ffiiny of sltion.For ample,if fa- protin squns.so that
vorbl mutaions aris indpndntly, but vry rrly' at two sits, th popu- prfrd ovr A. In an o
ltion will rarly trntainany individualsthat arry doul mutants'vn if this omPonnt of tlr gllomr
is th fittst omination (unlssth popultion siz is so hug that th sond whr romination is in1
mutation vnt happns for sltion fis the first favoral mutation). G- mal odon usag ight
nti rombination allows th doul-mutant omination to e formd. would lss isdthn
Whn th is littl or no enti romination' or whn rprodution is asx- o f t h o s r r ' r ]r l l t i o n s
ual, sltio on on gn thus impds sltion ating on othr gns' making ontnt of oding squn
fvoral mutations lss likly to sprad and dltrious muttions (suh as and for th wakning of
amino aid hngs in prtins) mo likly to fid y gnti drift, pati- ttd for th gns addd
ularly whn population sizs ar low (Gordo and harlsworth 2001). This and harlswlrth 2006).
Pross proaly ontriuts to gnon-lrdution in symioti atria that Horvvr, th G ()nt
liv within inset lls (romination btwen tria of ths spis is pr- . lgs also <lrrltsv
vntd by isoltion);thsrdudgnomsalso hav fastramino id squn ing sqttnsand rrontr
volution' as prditd givrrth lowrd aility of sltionto liminat muta- m n t i ( ) l ] d l r l i r 1 \ { r l r
tions with wakly disdv:rntagous ffts(oran 2003). rats i1nd dirtiolls of mut

l risk of ausingharmful mutations
ni rgions'squnsoftn ontrol
ity dpndson th waknss of s-
.s
raly xis from a site, gnti
rg irrvrsibly fid for an lmnt.
as humans' sltion against l-
: drift, and this possily allows th
la (Lynh 2007). Nw insrrions
lrvntdby sltion (thr would
r gratrrisk of topi hang),
. his may aOunt for th fixd l-
[or th vry high T aundan in
vaiailityindiatsa larg pop-
bination ir-rrniz may largly
i squnsould not rmovd
ld uild up twn gns' as o-
' th ausationould th othr
;s thurnhost squns.
:Ns oF GN|
.IoN
sIz
lrih smali population siz and lak
]nom volution. Romination
olutionaryprosses,spially th
s of sxul rprodution. Svral
nntd with romination rate
ination rats ar similar to thos of
l hirrlsworlr2001 ).
)at indpndntlyon diffrnt sits
:ny of sltion.For xampl, if fa-
vry rly, at rwo sits, th opu-
at arrdoul mutants' vn if this
rtion siz is so hug that th sond
es th first favrlmutation). G-
.utant omination to formd.
Ltion,or whn prodution is as-
tion ting on othr gns,making
rnd dltriousmutations (suh s
, to fid y gnti drift, parti-
do and harlsrvorth 2001). This
:dution in symioti atria that
'ntriof thsspeisis p-
alsohavfastmino id squn
l i l i t yo f s l t i o n
o l i m i n a t m u t -
oran 2003)'
uolutkl rlf the (}ut 171
Gnsin gnom r]ionstlrat lk gnti rlminatilnshould r.rndrgo
similarvolutionirrvprosssthar irpdslionrrlld tl rdud lv-
ls of daptation.This may also ontiut to th uildup of Ts in nonr-
omininghromosomal rgions and to ths rgions' inrasd intron sizs
(prhapslso irr p2rrtausd y -drivd matrial). Y s lromos<lnts
strikingly illustrat th disastrous onsqttns lf not romining
(D. harlswortht al. 200.5).Th mammlin Y and hromosoms shur
som gns' tr..rvingthir omnlon dstlt fr<lm nlrmal homlso-t
pair,ut th Y lks most lf th mor than 1,000 gnson th -it has lost
thgntiinformatirron shrd with th X' This is jr'rstas prlitdflr :r
nnrorininghrlmosorrr: sltionating tltl hur-r]rds of grrswilI irn-
pdthir aility tl maintai adaptation. Y-linkd gns should thus volv
festrthar.rthir X-linkd ountrparts if th ility of sltion to rmOV
disadvntglus r,ariantsis irnpdi;tlry slrould volv r-rrorlik pstldo-
gnsthan lik normal gns.Fastr Y volutilnis indd dttdin analy-
s so f s . l i n k dg n s i n m m m a l s ( G r r a r d a n d F i l a t o v 2 0 0 5 ) a n d i r d s
(Fidolfsslnarrd llgrn 2000). A sirilr alration is osrvd f<lrtlr
gnsaddd to th Y hrontosom of D. mird, and plymorphism dlrta
onfirmthat this is luto infftivsltionagainst dltriousmino aid
1uttions(Bartollmnd hlsrvorth2006).
Similar ut mildr fftsar osrvdwhn w ollparotr gnom r-
gionswith diffrntromination rts. Gns oftn disproportilnatlyus
odons that nd in G or ratlrr tharr tlros tht nd in A lr ' so the G
ontntof third oding positions tnds to highr tharr that of thi intrtltrs
or th adjant nonoding squn.In atria, D. mllgstr,. l-
gs' and . tlllilt' thr is onr,iniIrgvidrrthilt tlris odrn usag is
rfltsth ation of sltion, proly us G-nding odons allow
fastrand/or mor aurat translation of th mssngRNA ir"rtoth pro-
t i ns q u n ( A k s l r i t l . 1 9 9 8 ; S h p t l . 2 0 0 5 ) . T l " r s l t i o l l r s s L l r
is, howvr' vry small ompard ll,,iththat xting on most hlrngsto th
protin squns'so that thr is orrly a sttistial tndny flr i to
prfrrdor,r AT. In an orgrrism lik D. mllgslr, r'r,ith srrstirntil
omponnt of th gnom in rgions nar th ntromrs and tl<lrns
rvhrromination is infrqunt, t aility of sltioll to maintin ()Pti-
m l o d o n u s g m i g h t l s s n ds r " r f f i i n t l 1 , t h g
t r r si n t s g i t l l l s
w o u l d l s s i a s d t h n l s w h r i n t h g n o m .T l r i s a o u n t s f o r p a r t
of th osrvd rIatilnship twn ror-nination ats nd th G
o n t n to f o d i n g s q l l n sl n D r o s o p h i l a ( i V l a i si r n r l P i g r l i l t r2 0 0 7 ) '
and for th wakning of sltion flr odor-rusag ias that has n d-
ttdfor th gns addd to th Y hromosol inD. mirda (Bartolorn6
nd harlsrvorth2006).
Howvr, th G ontnt lf nonoding squnsin D. mltlgsterttnd
' lgsalso orrlatswith r<lminationrts' nd G ontnts of od-
ing squrrsand nonodirrg squnsar olatd rvith ah Othr, aS
mntiondarlir (arais 2003; Lyrrh 2007\. Nonsltiv flrs,suh as
ratsand dirtions of mutatiln,thus pral lstl varr, with romin:rtion
172 uolutilrlf th Gome

rats. rtainly, not all muttions oLlr qually frquntlv' As lrady mn- latiol.lsizs drsdas
tiond, th rlativly high A ontnt of nonoding DNA is ausd by muta- trons xp()ss gns t() ]
tional bis toward AT. A pross that may oppos this is bisd ge hang th prltin irnd
onursion; aus of th molular mhnism of gnti romination' spliir-rgmust Onsv
whn on gnom has G at a sit and th sam sit in th oth has AT, r. y s l t i o nt l n l y i n s i
omination an podu a vry slight ssof G in th daughtr gnom prssur inrss intron
(arais 2003). This will inasG ontnt in oth oding and nonoding pokllry<ltsand prs.ll
squns' ausing Gs to fid mo frquntly than ATs' just as though 2003; Lynh 2007). Sl
Gs wr favord by sltion.Baus it is pat of th romination pro- o r r h r e s o n s .h u i n
ss, isd gn onvrsion is xptd to at mainly in rgions with high Again, lr<lrr.vr, it s
rats of romination, lading to a highr G frquny in suh rgions. Prsn o l l i n t o n sl l l
Thid positions of odons will hav an dditionl nhanmntof G from aus intron siz and r-
th fftsof sltion for odorr ias' Statistialanalyssof th G ontnt h l p p r l . i , f t l i t l s t l n .i n
.Whatvr
of th gnom of D' mlnogstel"supPort this hypothsis, as do data from t l. 2002). tlr
th highl romining parts of sx hromosoms of nrammls (arais to lm domstiatd
2 0 0 ). suggstdfor tltlrr gn<l
ltiv onstritints on intr
kryot gnoms (Hallig
nlusions F.urtlrr progrss in unl
volution is to ptd
Gnom volution, lik othr volutionry hangs, is opportunisti. hangs trm possil. Th Tw
in gnom faturs hzrvonsqunsof two kinds. First, hang may lad to gnoms <lf 12 rnmrs.
futhr hangs. For instan, hr{rmosomrarangmntsafft romina- stP in this ]irtion(Drt
tion. A gn an mov into a rgion of lowr or highr romination, afft. th rtnsntlrrlgsin p.
ing th ln twn iasd gn onvrsion that pushs G ontnt up partitllar vnts in gnorl
vsus mutation Prssur in th othr dirtion, So that GC ontrrt will start ing spissoirld allorv s
hanging. If a rarrngmntrdus romintion, as happns whn a hro- auss Of many hangs.-
mosom arr is joind to a Y hromosom, th drasdaility of natrrrl s- possil to onrpar indp
ltion to liminat wakl disadvntagoushangs may ultinratlylad to mon frors, sirrrihr to th
th loss of funtionality' or vn th physial prsn'of gns from this hro- volr.ltionayhngs(.g
mosom' as indd has hppnd in D. tird (D. harlsworth t al. 2005). ous tsts of hypothssthl
Sond, vn whn a gnonl fatur r,olvswitlrout ing favord y in-
dividual sltion,th an oasionlly ptntially advantagouseffts BILIoGRAPY
for organismal funtions. For ampl, Ts that ontain squnsthat on-
trol th prssion of thir own gns n lso fft th ativity of gns A k s h i 'H . , R . l \ { .K l i m n ,a
s | t i l r rl r n d h r ' t > | t
los to thir insrtion sits.Initially parasiti slfishDNA (spradingpurly
49-60.
as a onsqunof sltion for th squns'own aility t rpliat)an Th Aridopsis Cnlm In
..domstiatd''
bom as a part of th gnom of a spis,prsntin all in- flllvring pltt rbidl,
dividuals nd funtioning in its host, rathr lik th organlls'whih wr Arvin, A. A.' G. J. Hantln
on indpndntorganisms (Brokfild 2003). This possiility is often ovr- rovids an alirtivd
lookd, ut it is important to avoid th naiv attitud that prsnt.davn. Asthl-r,S., \. s. Nol, G.
St:rnll-ttovnnlgl<rul2
fits an plain th origin of irll gnom fatus.
tl.rhuma gt.ltlm.Prr
Genom vlution is far mor ompl than straightforward adaptiv vo- 12410-1241 5.
lution y natural sltion,and many fatursmay hav hd a long history in Bahtrg' D. 2003. AunrL
gnoms for ny nfis t<l th organism volvd. It has vn n lIrtlr
trtrl.lsp()s<rns,
Bitll,lgl, llt] L u,lltt!i,'tt
suggstdthat introns may initially hav sprad within gnomswhn popu-
 uolutio lf th col 17.1

qullyfrquntly.As alrdy mn- ltionsizsdsds rrruIrillrrl orgnisrsvolvd.Th prslrof in-

nonodingDNA is usd y mut- trons poss gns to tr hrmful mutations in addition to thos that
: my oppos this is bisd ge hangth potir-rand lrrtollirrgsquns (gu5gsis rquird for
:hanismof gnti romination, spliingmust onsrvd).It is ptlssiltht thir prsnan rsistd
1 sam sit in th othr has AT, r- ,vsltiononly irr spisrvith vry l:rrgplpulations.I som rlrutational
:ssof G in th daughtr gnom prssurinrirssintron nunlbrs. this uld aOllnt for introns' rarity in
ltnt in oth ding and nonoding rokaryotsand prsrrin multilltrlar uk:rryots(Lynh and onry
tiquntly than A, just s though 2003;Lynh 2007). SItionm lstl firvo small rokarvot gllom slz
it is part f th romination pro- for othrrasons' llut in prinipl all rrpposingfors must Play som rol.
to at mainly ir-rrgions with high Again, horvvr,it srs unlikly tlrt n.rutirtill prssllr plains th
hr G frequny in suh rgions. prsnof ll intons in gns of living multillular ukarylts,spially
dditionalnhanmntof G 1rom ausintron siz and nunlrs an drs,s wll as itrras(this has
.ltistilanalyssof th G ontnt happnd.frrr instn,in th smll gnom of th plant . tbli;.Wright
'rt this hypothsis,as do data from tal.2002). \/htvrth aussof thir initial origins, introns hav volvd
lromosoms of mammals (arais to lt-l-t
drnstitdnd l-rvqrrird llulr funtirns' as w lrev
suggstd for othr gn()mfaturs,and thr is inrasing vidn for s-
ltivonstrail-ts tltr inttln rrd <ltlrrkirrds of nonoling squnesiIl u-
karyot]noms(Hallign and Klghtlv 20061Asthna et al' 207).
Furthrprogrssin undrstandirrgth ausaIprosssinvolvd in gnlm
volutilnis to ptds mo ilnd mor dtaillomparisonsof grroms
, hngs'is opportunisti. hangs ompossil.Th Twlv Gnoms Projt, irr whih larg parts of th
wo kinds. First, lrang may lad t gnomsof 1) irrrrs<lfth gtrtls Dros;pbil have n squld,is a
l rarangmntsafft romina- stpin this dirti<ln(Droso1lil l2 Grroms onsortium 2007). Although
vr or highr romination, afft- th onstnt hngs in gntnlsmk it ditfiult t undrstand th arrsslf
vrsion tht pushs GC ontnt up partiulirrvnts il-tgnOmr,olution' suffiintly dtaild omparisons of liv.
tion' so that G ontnt will start ingspissh<luldlIow us tl disrnpattnsin gnmsnd tl inf th main
lination, as happnswhn a ho- ussof mnv h;rngs.Bausgnom hangs ar frqutt,it should
l, th drasdability of naural s- possiltO ompar indpndntassot simi[rhangsand t<ldisovrom-
)us hangs may ultimatly lad t mon ftors,similar to t produswi]lvusd for sttrdyingoth typs of
al prsn,of gnsfrom this hro- volutionary hangs(.g.,rv and Pagl 1991I.Ths provid mor rigor-
,d(D, harlsworth ousssf hypothsstlrn r<lal
t l. 2005). oInparislnsamolrg distntly rltdaa.
,.olvswithout ing firvord y in-
potntially advantgous ffts
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F i n a l l y .s p i e t i o ni 5 .
Zang, x. H.-F., and L. A. hasin 2006. omparison of multipl vrtrat
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oms a ompltly spa
volutionary hang witlr
tion into nw linags(FiE
ontinuum tlf spietion
tionry iologists wh w
this hurdl, t study lf l
ing aras of volution:rry
 esin the size of the
: the C-valueparadox.
19 119-124.
m, K. Collura, D. S.
,ublinggenome size
rn-driven genomic
;. GentnneResearch\6: The Patternand Process
4way.M. V.
sen.2002. Evolution of
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rkharov,A.
1.Avramova,and J. L.
Margaret B. Ptacek-dnd ShalaJ. Hank"ison
;enicregionsof the maize

Adtancesin Genetics 46: Tl-reessentialbit of evolutionary theor\.which is concernedrvjth the ori-

gin and nirture of spet:iesremainsr.rtterlyrnyster',.rut.
*. Batcso. (192r)
,. Sockett.2005.
.mongbacteria.Nwcleic

2000.The evolution of
BioEssays22: Speciation,the evolutionary processby which new and distinct lineagesarise
and mirintain their independenttraiectories,has been a perplexingphenome-
'terminingcentfomere
non ever since Dirrwin (18-59)first proposed that all living organisms have
uiewsGenetics2:
diver:sified frorn sharedancestors.There are severalreasonswhy unclerstand-
ck, J. F. Doebley,and i n g h o w s p e c i a t i o no c e u r si s c h e l l c n g i n g .F i r s t . m o s t s p e c i r r i o nt v c n t s h r v c
ion alongchromosome 1 occurredin the past, leaving only the end products, species,:rs a signattrreof
- 14 1 3 . their occurrence.Species,l.r<tth exrant and extinct tornts, [re,rrthe cher,rctcris-
rt. 2002. Evolution of tics rlssociatedwith their divergencefrom conlrlon ancestors'but in addi-
ing invaded by
tion, they also carry unique phenotypic and genotypic changes that have
ion 19: 1154-1161.
occurreclsince speciation.Disentanglingthe characteristicsassociateclrvith
ew fnnctions? Genetics
the speciationprocessfrclm those thzlt have evolved since divergellcecan be
,olution and particularlydif6cult.
4sof tbe National Second,new speciesarise through r'rvariety of evolutionary mechrrnisms
(e.g.,natural selecticln,sexual selection,genetic drift, anci mutatioll), and
of recombination rateand is
therefore,finding a unifying definition that describesand clefinesall sf-'ecies
Arabidopsisthaliana.
an impossiblechallenge.Thus much clebatehas occurredal'routthe nature of
a n d p a t t e r n so f m o l e c u l a r speciescharacteristicsand how scientistscan distinguish speciesfor taxo-
lar Biologyand Euolution nomic,systelratic,and conservatiorrpurposes.
Finaily, speciationis a seriesof processes, with :r beginning stageof initial
nultiple vertebrate a middle stagewherein species-specific characteristicsare refined
divergence,
tns. Proceedingsof the
bv vari<,rus forces of evolutiotr,and an end point at which ir new speciesbe-
)L.

comesa completely separaCeevolutionary lineage on its 1;wn trajectqry of

evolutionarychange with the potential for extinction or further diversifica-
rion into new lileages (Figure 1). Knorving the locirtion of urspeciesalor-rgthis
continuum oi speciation has proved to be a perplexing ;lroblem ftlr evolu-
tionary biologists who wish to understand how new species:rrise. Despite
t h i s h u r d l e , t h e s t u d y o f s p e c i a t i o ni s o n e o f t h e m o s t i m p o r t a r r ta n c le x c i t -
ing areasof el,olutionrrr,vbiologv. A resursenceof interest in speci:rtionby
178 The ttr d Pross of Spitio

slrion.gnti drrft. a
Speies Speies
gnotypi diffrnsbt
how nd why nw spi
in whih nw spis r
primary mans y whih n
dnt voIr-rtionarypathwa
hv gul.rto rsllv th
xllo

Th Natur of Spi

Spis onpts, :r st ot
a o u n d i n t h l i t r a t u r ( }
llng-standingdat enro
lm is ntrd arlund ir
llll. pttrn-orintd <lrPr()
Dlvr frnsof opinion tln wht
plial (Figur 2). P.rtt
spis. For ampl, th
Common y Thodositrs Dozhrs
( | 9 4 2 , \ 9 6 ) , d f i n ss p
Anestor
nisms. Rprodutiv islla
arrirs to sussfulrp
Figur 1. Th prossof speiationis a ntinuum.InitiaIdivrgntwn
Poss-orintd dfinit
populatinsis followd y priod of angnihangwithin ah nw linag
m h a n i s m s p, h n t y p i
whrin spis.spifihartristisar rfind y various fors of volution.
In som spislong pridswith littl volutionaryhang(stasis)may ()ur. ion of som lvl of div
one formd,spismay furtherdivrsifyinto nw linags(ladognsis) or s p e i s i n t t l d i s t i n t n t i t
may om tint. Th hallngin th study of spiation is unovring ( v a n V a l n 1 9 7 6 ) d f i n ss
whre along this ontinuuma spisatuallyxists. l q u i l n t s n d l d n '
from othr spis thr<ru
llogial hartristlst
volutionary iologists ovr th past 15 yars has ld to mny iting ad- grphi xhangaility'' 1
vans. Ths inlud th disovry of svral spitio gs that onfr r- Al:rn Tmplton (1989)
produtiv isolation, a ttr undrstandingof th importan of sympatri uniqu llogiirlprr)prt
spiation (spiationwithut gographiisolation);and a surrtionof in- volutior-rarySpis <
trstin how natural sltionan lad to th formatilnof nw speisas y- s p ar e t v o lu t i o n 1 r' l j
produts of adaptiv diffrntiation. on pross' that is, voluti
In this ssayw outlin som of th major hallngsin undrstandingsp. l u r i o n a r y l i n a g d f i n i t i
iation and showasxampls that illustratth major advansin th study sin n formlizd with
of th origin and divrsifiation of spis. w start with th prolm of and ollagus in th Phy
dfining spis, and how an undrstanding of th undrlying mehanisms Spis ar d{inds sP
that lad to th formation of nw spis may the st approah in at- that rsLlltrorn divrgn
tmpting to dfin what a spisrally is. Nt, w outlin th various ryps s p i sd o n o t o m d i n
o f i s o I a t i n g r r i st h a n n i n i t i t 'p r o m o t .a n d o m p l t t h s p i a i o n ompltly sd (FigLrr
pross.Finally' w disussth volutionary mhanismstht driv th spi- Wht do w do with all
ation pross. Outlining th rlativ rols tht natural sltion, sxual dfin spis irnywy? T

peIes
Ct-oosls
}Ssls
Arsts
)mmon
nestor
inuum.Initialdivrgntwn
;nihirngwithin eah nw ling
'findby variousfosof volr.rtion.
utionaryhang(stasis)may ou.
into nw linags(ladognsis)
, u d yo f s p i t i o ins u n o v r i n g
lly xists.
yars lras ld to m:rny xiting ad-
.era|spittio|1
gnrs that onfer r-
ing of th importan of sympatri
: islation);and a rsurrtionof in-
th formation of nw spiss y-
jor hllngsin undstandingsp-
atth major advirnsin th study
:is.\7 start with th prolm of
ling of th undrlying mhanisms
]s may th st approah in at-
. N ,W o u t l i n t h v i o u s t y p s
)mot' and omplt th spiation
try mhanismsthat driv th spi-
lls tht ttural sltion. sxual
hc Pttr d Prlssof Spitil 1 7 9
sltion,gnti lrift, and muttion plv in prom()titlg phnotypi and
Speies
gnot1,pidiffrnstwn populations is ritial to our undrstanding of
D how ar-rdwhy nw spisaris. Our glal is not t() ovr vry potntial way
in whiIrlrw spis r form.lt rathr tl illustrat with mpls th
primarymans y whih nw s1-lis ar f<lrrndirnd rrr:rintinthir indpn-
dntvolutionry pthwys. Through suh studis, vllutionary iologists
havgunto rsolr,th ..mystrisof spiation'' (Batson \922)'
Th Natur of Spis
SpisOnpts'a st of ruls tlr hartistisusd to dfin a sP1'igs'
abotrndin th litratu(N[ir,vdn|997;aLl|e l)nd hilr,n a srrjtof
long.stndingdtamong vllr"rtionary illogists.Tlr natur of rhis pro-
lm is ntrdarund a disagrnrnttwn v<llutionaryiologists on
pattrn.orintd or pross-orir-rtd dfinitiolrs,whih, in turn, lads to dif-
frns of opini<lnltl whr in tlr spiation pross dfinition is most ap-
plial(Figur 2). Pattrn-orintddfinitilnsus ruls or ritria to dfin
spis.F.orampl, th Biologial Spis onpt (Bs), first artiulatd
y T h o d o s i u sD t l l l z h a n s k 1 9 7 ) n d l a t r h a m p i o n d y r n s t a y
(1942, 196.3),lfinsspis ln th asis tlf prrldutivisolting mha-
nisms.Rprodutiv isolatiorr 1ay Our thr<lughprmting or posrmating
arrirsto sussfulrprodution.
Pross.orintddfiniti<lnsdo not us slifihartristis (isolating
mhisms,phntltv1.li harats)to dfin i1 spisut athr rrs rit-
rion of som lr,lof divrgrr(vllutionryor ollgial)that spaats
lr spisinto distint ntitis. }.<lr-ampl,th ollgial Spis onpt
(vanValn 1976) dfinsspisy thir uniqu nih, th sust f ologi-
l rquirmntsatld adaptations plssssiv a spis that hs divrgd
from othr spis through irrtrspifiotnptitin, prdatiorr,lrrrd othr
.,dmo-
olgialhar:rtrististhat <rrganiztlnrtrrunitisof spis.Th
graphixhangaility''part of th ohsiln Spis(ilnptpropsd y
Aln Tmplton (1989) provi1sa si.rilr]{initionof spis as hving
uniquoitlgiirlpoprtistht sPartthrn tronr tltlrrspis.
r.llutior-rary Spis orrpts dfin spis on th asis of uniqu and
sparatvolutilnarytrajtoris (.g.,Simps 1'961);gain, an mphasis
on pross'that is, volutionirrvt1ivrgn, is usd to dfir-r spis.Atr vo-
lutionrylinag dfirritionwirs first proplslb1, d Wil1, 0978) zrrrdlras
sinnformalizd with tlr us of phylgntiinfrny Jl araft
a n d o l l a g u si n t h P h y l o g t i S p i s o n p t ( r a r a f t 1 9 8 , 1 9 8 9 ) .
Spisar dfinjetssparetvolutitlnary linagsor ronophylti lads
tht rsult frrlrl livgnfrtll-tlll ()mmOn l]Ilstor.Tlrtrs vtllutitltrry
spisdo not ()mdignlsluntil glr flow witlr all lthrlinl-rgs has
omltlyasd(Figur 2).
What do w dl rvith all ths diffrnt dfinitions,and why do w nd to
dfinspisan'ur.v? Th lglrismsthat lr-rkup t..lisrt ntitis that
1 8 0 h Ptter nd Procss of Spitil

Tabl 1. Spisonpts'thir dfinitions'and rfrns

Spis
onpt Dfinition Rfrns

Patternorintd Biologial Groups lf intrrding D o z h a n s k y1 9 3 7 ;

natural populationstht ar 1942,196
r produtivlyisolatd
from othr suh groups.
ohsion', Th most inlusivgroup T m p l t o n1 9 8 9
of organismshvingth
potntialfor gnti
and/ordmogrphi
xhangility.
Phnti A st of organismstht Sokal and rovllo
look similar to h othr 1970
and ar distintfom othr
suh sts.
Rognition A st f populationsthat P a t r s o n1 9 8 5
shra Ommon
frtilization systm
(spifi mJt-rOgnitit-ln
systm).
Prossointd ohsion'. Th most inlusiv group Tmplton 1989
of organisms having th
potntial for gnti
Figur 2. An illustrtionof }
d i f f n tp o i n t s i n t h s p i
and/or dmographi .Whn
into two. divrgntlir
hangaility.
Spisonpt (S) will
ologil A st lf organismsdptd ana|en 1'976
rprdutivisolation (ISM1
to a singlologilnih,
sparatspisundr th B
volving sparatlyfrom
rPrsntmonphyltilad
lingsoutsidits rng.
ommon anstor)for
volutionary A singllrnag S i m p s o n1 9 61 ; Spisonpt (PS).
(an anstral-dsndant W i l y - l 9 7 8
squnof ppulations)
volvingsparatlyfrom
othrsand with uniqu sintists all spis sm
volutionarytndnis annot sm to agr.Th
and its own historialfat. having som form of dfi
P h y l o g n t i T h s m a l l s t d i a g n s a l r a r a f t1 9 8 3 , vn if diffrnt spis rt
lustrof individual t989
aris. First? dfining iolog
organismswithin whih
nr in whih to lassify.
this parntlpttrn
f anstrynd dsnt. Sond, dfining diagnosa
ntitis that xist in natur
".Prts f dfinition onsidr oth pttrn and prss lf spiation.
spis ais without havir
pla. Fourth, spis,as
th volutionary history O
surd y th numr and
usuallymad at th sp
atgorizing and onsrvl
 T'he Pttct rld Pross of SDition 181
and rfrns

Rfrns Lr'

intrbrling D o z h a n s k y1 9 3 7 ; BS
lulationsthat ar 1942,1963
ltivl1,
isolatd
srrhgroups.
-rIusivgroup T m p l t o n1 9 8 9
ls havingth
lr gnti
rgrphi
rility.
;anismstht Sokal and rovllo
r to hoth 1970
tint fom oth

pultinsrhat Patrson-l985
nmon
l systm
rt-rogniion

nlusivgroup Tmplton1989
ns havingth
)I gntl Figur2. An illustr:rtionof lrorv diffr:ntspisonpts dfi.rspisat
rographi diffrntpoints in th spiationprossaftr th splittingof a sil ling
rility. into two. lfhrr divrgntlinagsdisprsinto diffrntaitats'ih tgial
adptd van Vln 1976 Spislnpt(S) will dfin tlrrns sprtspis.Th volutin of
;arrisms
ologilnih, rprodutiv isolation(ISM1 or ISM2) twnth two lincswill lfin
paratlyfrom sarat spisundr th BiologialSpisorrpt(BS). iinags must
tsidits rang. rrsnt monlphyltiiads(all p<rpultilr.rs irdt-snd.i frlm singl
o m m o na n s t rl)r f r l
i n gr t l g r l i z ds s p i su n t l t h P h y I o g n r i
tag Simpsn 196 1;
Sis onpt (PS).
l-lsnd nt Wily 197tl
f populations)
paratlyfrom
with r"rniqr-r sintistsall spissm to know thir own oundaris' vn if billgists
.y tndnis
nnotsm to agr.Thr ar, howvr' som vry imprtant sons ftrr
r lristorialfat.
h a v i n gs o m f o m o f d f i n i t i o nt h a t s i n t i s t s n u s o o g n i z s p i s ,
sr digr.rosai a r f t1 9 8 ,
ven if diffrnt speis rquir different ritria for dfining thir ound-
rdividual 1989
aris.First, dfining iologial ntitis s spisprovids a svstmatiman-
withinwhih
arntalPattrn nr in whih to lssify divrsity and is th sis of modn taonomy.
and dsnt. Sond,dfining diagnosal groups as spis rrsponds to th disrt
ntitisthat ist in natur. Third' on annot study th pross of how nw
l r s f s i l l i n
spisaris without having som dfinition rf what a spis is in th fist
pla. Forrth, spis, s th last inlusiv tgory of livrsity, rprsnt
thvolutionaryhistory of organisnrs'Finlly, iodivrsity is gnrally ma-
surdy th numr nd kinds of living organisrs,and ths stimtsar
usuallymad at th spis lvl. Thus for a varit of asons, inluding
atgtlrizingand onsrving divrsity, as wlI s undrstanding its oigitls'
182 h Pttr d Pr-lss
of S1litk

dfining spisas disrt iologial ntitis is ssntialto our undrstand- l 2. Isolatingarri

irrg of th pr<lssof spiatilnand th pattrns of iologial divrsity. No
yp.
on spis onpt will aomplish all ths purplss.volutionry ilo-
gists should think aout th natur of thir qustion with rgard to th Prmating ol
..spisproim,'' as wll as th natural history of thir organism of intrst,
and thn hrosth onpt that applis st ttl th solving lf that prtiular
proinr. Tm

olli
Th Natur of IsolatingBarrirs

Th importan of isolating arrirs is narly univrsal rgardlss of th

Bha.
spisonpt ing applid. Although th link twnrprodutiv isola-
tion and tlr BS, whih dfinsspison th asis of rrirs to intrrd.
Postmating, h
ing, is lvilus,isoiatirrg arrirs also provid th mans through whih przygotl
plpulatilrrsgin to divrg through drasdgn flow, th first stp in
spiation. Thus vn m()r Pross-orintdonpts may linkd indi-
rtly tO th fftsof isolating arrirs ausths arrirs may at som
plint play tr rol in ssationlf gn flow twnvolutionry linags' gg-s
Thr r nulrous Ways to lassify isolating rirs;htlwvr' on of th
rnost widly usd divisions is twn prmting and postmating brrirs.
Ths rrirs inlud thos that ar du to trinsi fators, suh as olog-
ial lr haviral rhanisms, and thos that ar du to intrinsi fators,
onsp
suh as gg/sprminompatiilitis lr gnti inmpatiilitis twn hy-
pr
rid gntlms(Tal 2).
Prnlating errisinlud thos tht prvnt or drsth liklihood of
mating twn taa. In fft, prmating arrirs prvnt taa (or th ga-
Postmating, Hyri
mts from thos taa) from mting.In ologil islation potntial mts postzygoti
dtl ntlt met ausof diffrnsin haitat us. For ampl, many insts
Hyri
ar lirnitd to fding, ourting, mating' and ovipsitioning on spifi plant
hosts. Thus diffrrltspiser prvntdfrom intrrdingy th rstri-
Dr
tion of ah ttl spifihost plant (Futuyma et a|. 1994). Altrnativly,indi. fitnss
viduals may tmporally isolatd' diffring in thi timing of rprodution.
For ampl, th sympatri oral spisotstr tlulris, . fullt,
and . frksi ar tmpoally isolatd through diffrnsin pak spawning
tims, and diffrns lf only 1 to 2 hours twn spawning vnts provid othr pre- or postmatil
nough tim to dilut gamtsand prvnt intrspifihyridization (Knowl- rding) may avoid int
ton t al. |997).In othr assindividuals of diffrntspismay prsnt ing signals that rstrit
at th sm tim and pla arrd still avoid intrrding.Pollinator isolation is laorat mal ornam
tln suh mthod. haratrs suh as flowr shap,rwd (polln vrsusn- rltd spis r oft
tar), and olor attrat diffrnt typs of pollinators (.g.,moths' bs, hum- (Andrsson 1994),
mingirds) that rrray prfrntiallvvisit only rtain flowr tps, nsuring Dspit th inhrnt
tht plln is trarrsfrrdonly within spis. mating at th outst' m.
A {inl typ of trinsi isllatingarrir rlis Ol-lhavioral diffrnsas pn oth pr- and postz
prmting rrirs.losly rlirtdspisoftn diffr in partiular traits tht in taxa with postmatinl
funtion as signirls in th mating pross. Thr-rsvrrspisthat hiit no pla. For xampl, in
 Tb Pttr nd roess of Speition 18

:isis ssntilto our undrstand- Tal2. Isolatingarrirs(othpr- and postmting)

ttrnsof iologial divrsity. No
Typ of arrir Dfinition
lspurposs.volutionry iolo-
hir qustion with rgard to th Prmating ologialisolation Spisar sparatdy
istoryof thir organism of intrst, diffrnt hosts or
mirohaitats.
stto th sllvingof that partiulr
Tmporal isolation Rprdution ours at
diffrnt tims.
Pollinator isolation Di ffrnspisttrt
diffrntpollinators,or polln
of diffrnt spisis arrid
on diffrnrody parts.
narly univrsal rgardlss of th Bhavioralisolation Bhaviorsusd as mting
r link twnrprodutiv isola. signalsdiff twnspis.
' th sisof arrirsto intrrd- hanial isoltion
Postmating, Sprm transfrannot tak
lrovid th mans through whih przygotl pla ausgnitaliaar not
:rsdgn flow, th first stp in ompatil(or pollen dos not
rtd onpts may linkd indi- adhr)twndifferent
lausthsarrirs may at som spis.
t w n v o l u t i t l n yl i n a g s . gg-sprmrognition Proprtis of th gamts
lating arrirs;howvr, on of th (oftnsurfaproteins)are
rmatingand postmating arrirs. not ompatiltwn
spisand prvnt
o trinsifators' suh as olog-
frtilization.
that ar du to intrinsi fators,
onspifi sprm Htrospifisprm ar
nti inomptiilitistwn hy- prdn outomptd y onspeifi
s p r m 'r g a r d l sosf m a t i n g
v n ro d r a s t h l i k L i h t l o do ordr.
g arrirsprvnt taa (Or th ga. Hyrid inviability
Postmating, H y r i d sd i f o r h i n g
ologialisolation potntial mats postzy8oti sxual maturity.
i t a t u s . o r x a m p l .m e n y i n s t s Hyrid strility Hyrids ar inapa1of
rnd ovipositioningon spifi plant rproduing.
d from intrbrding y th rstri- Drasdhyrid Hyrids hav lower fitness
,ma t al. 1994).Altrnativly,indi- fitnss t h a n i t h rp a r n t sl p i s
ing in thi tirning of rprodution. and sltdginst.
\otstrnulris, . fueolt,
rough diffrnsin pak spawning
.s twnspawning vnts provid othr pr- or postmating isolating arrirs (e.g.,sympatri or synhronous
t intrspifi hyridiztion (Knowl- brding)may avoid intrrding aus of prfrns for partiular mat-
s lf diffrnt spismay prsnt ing signalsthat rstrit thir mating options to mmrs of a singl spis.
intrrding.Pollinator isolation is laoratmal ornamnts nd ourtship displays that diffr twnlosly
'r shp,rward (polln vrsus n. rlatd spis ar oftn thought to rsult in hvioral isolating arrirs
pollintors(.g.,moths, s' hum- (Andrsson1994\.
o n l r t a i nf l t t w rt y p s ' n s u r i n g Dspit th inhrnt appal of arrirs that prvent diffrnt taa from
ls. matingat th outst' many isolating arrirs our postmating and may hap-
: rlison havioraldiffrnsas pnoth pr- and postzygotially. As implid y th nm' mating may Our
s oftn diffr in partiula traits that in taa with postmating, przygoti arrirs, but frtilization dos not tak
;. Thus vn spistht hiit no pla.For ampl, in oth plants and animls onspifi sprm (polln)
184 The Ptter d Pross of Spitio

prdnis ommonly fund, whr, dspit multipl mtings, th fmal

(or fmal strutur of plnts) only irpts sprm (or pllln) from on.
spifis, or, altrnativly, onspifi sprm outompts htrospifisprm
for frtiliztion. Anita Diaz arrd ark Manair (1999| studid onspifi
polln prdnin th monkyflowrs imlus guttltL|sand . sutus. n
/
. gutttus onspifi polln-tu growth was fastr than htrospifi
polln-tu growth' nsuring orrspififrtilization nd onfrring rpro- (speies2) AAbI
dutiv isolation twnths spis.
Postmating' postzygoti arrirs rprsnt finiri atgory of isolating r-
rirs. Although this is a fairly O1monmhanism of rprodutiv isoltion,
Drwin Ws onrnd with this isolating irrrir ausit is th asis for
I
th prodution lf strilhyrids. His dilmm, simply statd,was this: if two AABT
ta volvd from a ommon anstor' how ould th omination of ths
.Would \
taa form a stril hyrid? that not imply strility t som tim in th
volution of ta anstralto prsnt-dayspis?Today w know, howvr,
that although hyrid strility is oftn an important islltingarrir, it is far
frlm th only form of postzygotiisolation, and a ttrundrstandinglf g-
ntis has yildd a solution to Darwin's prolm of stril hyrids.
Figur . Th Dozhansky
As with th study of many volutionary prosssin iology, Drosophil
dmonstrts how spiat
spis hav provd xtrmly usful in undrstandingpostzygoti rrirs.
intrmdiat hyrid stag.
In a lassi study of postzygoti rrirs, Alfrd sturtvnt (1920) showd that possess th variant allr
that rossing D. mloglsterfmals with mals of thir sisr spis taon rsults in spiatron
D. simulns yildd only fmal hyrids, witlr mals dying at th larval to pu. inompatil' mating tw
pal transition. orovr' th riproal ross gav ris to only hyrid mls invial, or othrwis lss ]
bausth fmals did as mryos. This study providd not onlY vidn gn intrtion oms fro

of postzygoti barrirs to hyridiztion ut also vidnthat ths arrirs

an appar at diffrnt dvlopmntalstags,that th dirtion of th hyrid
ross an b influntial, and that hyrid inviaility an span th rng from
partial to omplet (oyn and Orr 2004). fid) and th AA gn
Although th postzygoti arrirs just disussd provid vidn of th with th anstral alll (
varity of intrinsi postzygotiinompatiilitis,thy giv us only a rudimn- spis nd pass througl
tary undrstandingof ths arrirs nd thir rlationship to spiation;that and ..', allls ar inom
is' thy tll us what might happn' ut not why or how ths rrirs volv. d i f f n tl i n a g sm t 'p r
For that, W must turn to gnti modls of postzygoti isolation. rwis lss fit) aus tl
Darwin Was unawar of gns or gn intrtions,ut his onrn aout bakground, ar inomp
stril hyrids was on that trould vlutilnaryiologists for som tir. thugh this pross may s
Th qustion of how two spis an volv from ommon anstor yt' mating barrirs among
whn matd, form strilhyrids without at last on of thlsspispassing multilous gnti inom
through a maladaptiv form was quit prpling.A solution to this dilmma Anothr important ar
is providd y th Dozhansky-ullr n-rodl(although, s notd y orr obsrvation y J. B. S. H
11996]' Batson ssntillysolvd this sam prllm 25 yars arlir). Th with two diffrnt sx hr
solution dpnds upon undrstandingthat gnti inompatiilitisthat lad mals, ZW, in irds) is tht
to unfit offspring my amlioratd y mr.rltilor.rs intrations.ln th sim- aility. alld Haldan's rl
plst modl, onsidr two loi, AABB (Figur ). In ln spis a mutation in a varity of taxa (oy
aussth ..'' alll to appear and lmfid, rsr-rlting in th aBB gno. rogamti. Flaldan's ru
typ. In th sond spisa mutation lads to a ,.'' lll (again oming postzygoti isolation (oy

it multipl mtings, th frrrl
)ts sprm (o polln) from on-
outomptslrtrospifisprm
rnair (1999) studid onspifi
lus glttttusand . nsuts.
h was fstr than hterospifi
. r t i l i z n t i ll n d o n r r i l ] gr p r ( ) .
t a finl atgoof isolirtingar-
hnism of rprodutiv isolation,
zrirausit is th asis fr
, simplv statd,was rhis: if two
.,ould th omination of ths
irrrlystriIityt som tim in th
pis?Today w know, howvr,
lportnt isolating barir, it is far
' and ttrundstandingof g-
.oIrnof strilhyrids.
prosssin iology, Drosl|lhil
ldrstandigp()stzygotiarrirs.
Alfrd Sturtvarrt(1920) showd
l i r h m e | so f r h t ' i r s i s r s p
itlr malsdying t th l:rrvalto pu-
Ossgav ris to only hyrid mals
study providd rrot only vidn
rt lso viinthat ths arrirs
s' that th dirtilnof th h,vrid
tviailityan spn th rang from
disussdprovid vidn of th
ilitis,th1,givtrsrlnly a rudilrrn-
rir rlati<lnship ttt spiation;that
wh,vor how thsarrirs vr>lv.
lf postzygotiis<llation.
ntations,ut lris onrn aout
u i o n a 1 1'i o I o g l s t f r . l s t r m r i m .
llv fom a ommofl anstor yt'
rt lstlnlf tlrlsspispassing
pling.A solutilnto this dilmma
nldl(ltlrough,s notd y orr
prolm 2.5 yars rlir). Th
: gntiinompatiilitisthat lad
nultilous intrations.In th sirn-
g u ) . l n o n s p i sa m u t : r t i o n
fid,rsultingin th aBB gno-
..'' alll (agin tlming
ds to
Th Pttrnd Processof Specitiott 1 8 5
(inviable/steri
le hybrid)
AaBb
/ \
(speies
2) AAbb A A B B (speies1)
tI
AABb AaBB
\ /
AABB
(nestr|
spies)
Figur3. Th Dozhansky-ullrmodlof irrtrinsi
postmatingisolation
dmonstratshow spitionould produnhindrdy a dirimental
intmdiat hyrid stage.he anstralspeisdivrgsinto two sParattaxa
thatpossssth vriant allls..a',nd ...''Fiation of th nvl lls in ah
i s taonrsultsin spition(spis1 and spies2). Sine..a'' and ..'' ar
inompatil, mating twnth two spisrsultsin a hybrid tlrat is stril'
invial,o othrwislssfit thn th parntalspis.Support for this modl of
geneintrationoms from str"rdis >[Drosophil (oyn and orr zo04).
fid) nd th AA gnotyp. In ithr s th nw alll is omptibl
with th anstralalll (.g.,oth AaBB and AAB ar vial). Thus nithr
speisnd pass through a sltivlydtrimntal stg' Howvr, th..a''
and ..'' allles ar inompatil whn thy o-our. If individuals from th
diffrntlinagsmat, produing an AaB hvrid, it ould stril (or oth.
rwislss fit) baus th..'' and..'' allls, whn in th sam gnti
bakground' ar inompatil and form a postmating isolating arrir. Al-
thoughthis prossmy sm unlikly, multipl xampls,spiallyin post.
mating barris among Drosophil spis, show that ths typs of
multilusgntiinomptibilitisrny in ft quit ommo (orr 1995).
Anothr important ara of study in postzygoti isolation has fousd on an
osrvtiony J. B. S. aldan (1922) that th htrogametisx (th on
with two diffrnt s hromosoms, suh as mls, Y, in mammals, and f-
mals,ZW, in irds) is th on most Oftnafftdy hyrid stility or invi-
aility.alld Haldan's rul, this osrvation has onsistntly n supportd
in a varity of taa (oyn and orr 204| regardlssof whih s is ht-
rogamti.Haldan's rul appars to funtion arly in th volution of
postzygotiisoltion (oyn and orr 2004), sin rlativly rntly divrsd
1 8 6 h Ptter d ross rlf Speitiott

taa ar mor likly to xhiit aldan's rul; lldr taa oftn show svm-
omplt th spiatron
mtri hrid prolrns that fft orh ss.
ovrlap in gographi rar
Although Haldn's rul has primarilv n osrvd to rsult in hybrid
srv 5 strong ai
strility or inviaility of th htrogamtisx, sol vidn suggststhat
whos ranges ovrlap (s
htrogamtihyrids ray suff draslfitnsstlrrough a varity of othr
or whn prviou.slyisoll
mhanismsas wll. For arpl,Nal Davis anl lllagusshowd tlrt in
o n d r y o n t a t ) 'i n t r i n s
th Notropil uttrflies Arti ftim and . mtb strong assrrtlrtiv
rding om ns
mating urrd within ah spis'ur whn thy wr rossd artifiilly,
importan of rprodut
fmal hyrids showd dud trtiiity,onsistrrtwith ldan's rr-rl(t-
of spiation' and muh
mal uttrflis ar htrognlti;Dvis et aI' 1997). In addition to this
n t r do n r h m h a n i
mor typil rsult,howvr,hyrid fmalsfrom th . mtb (fmal)x.
twn spis'
ftit (mal) ross showd rdr'rdtnlnyto milt (Dvis et a|. 1997),
lassi modls of th s
his study provids vidnthat Haldarr'srul mzryxtnd yond th typ-
bution of populations un
ial osrvation of hyrid strility or inviaility, and that hyrid {itnssmay
gographi isolation to t}
drasdthrough th prmating and postmting arrirs'
d g o f g r t g r p i s
A omplt gnti undrstndingof postzygoti islating isis n.rh
an important ontriutol
mr omplx than aldan's ul aln implis. Of partiular intrstis th
of spition hav fous
onpt of speitir gns. Speiation gns ar gns that diffr tr-l,n
(natural sltion,sual
ta and us rprodutiv isolation trvnthn-r.A gn lld d1,55g'',
d i v r s if i r i o n e n d s p i
for ampl, shows vidnof rapid volution in srmfruit flis,dspitrl-
ativly slow volution in other organisms. This gn appars tl rsponsi-
l for strility in hyid mals trvenD. simllusnd D. mouritia (Ting H RoLE oF NA
t al. 1998). Lik olyssesJ]nsthat prinrpid voltion(t lilst in
Th imortan of adapt
som taa) may likly to us gnti inompatiilitistwntaa and
llgicl spcitio) Was fil
thus good andidats for spiation gns.u of th wrrk ln sPla-
in th 1940s in dsriin
tion gns rmains thortiurl; howvr, volutirr:ryilogists r addrss-
livd that spititlnin I
ing iting qustiorrssuh as whthr partiula gns ar gnrally rnor
divrglrt physiologia1r
likly to lad to rprodutiv isolation, or whthr gnsassoiatdwith r-
m n t s . n s M y r \ | 9 4 2
tain funtions ar ttrndidatsfor spi:rtingnsthn tltlrrs.
latd gnti diffrns
a n d | o g i e la d e p t a i o
rprodutiv isolating mt
Th Prossof spiation
dutiv isoltion is th dri
allopatri spiation (FigL
volutionary mharrismssuh as natural sltion,sual sltion,and g-
Nl[or rntly th imp
nti drift an promot diffrrrstwn populations in phnotyps and
d u t i v i s o l i o n i t h ri n
thir undlying gnotyps'vl1to th trrttht diffrnt ;lopuiatiorrs-
n important rl fo natr
om fixd for ontpltly diffrnt ltrnativs. Why, thn, is not vry
om lar. Laoator1
larly diffrntiatdpopulatiln nsidrd a diffnt spis?Th ru of
hav ofirmd tlrat o
th prolm sts on two issus.First' htl.,vmuh gn flow lurs tr-l,n
thr rzrisdfruit flis (D. l
diffrntiingpopultions? Gn flow :rtsto honrgnizpopulatiorrs both
vironmnts (varying tml
phnotypially and gntially;hn it works against slti<lnand drift in
s o m l v l o f p r n t a t i n gl
promoting daptiv or randrt diffrntiation' rsptivly.Sond, lrow
in diffrr-rtnvironmnts
muh diffrntiation is noLrgh?This issu is thlrIrirand points tO th im-
dpndnt lins raisd in
portan f rprodutiv isolation in th spitionpross.Although divrg-
s r t h o w t h p r r l s sm i
ing populations do not hv to volv intrinsi arrirsto rprodution, som
In th simplst modl
brrir to rprdutiv harrg mllst prsent to sto gIr florv and
volvs twrr popuiati

ul;oldr taa oftn show sym-
)nosrvdto rslllt in hyrid
s' som vidn suggststhat
l fitnssthrough a varity of othr
lis and ollagusshowd that in
n d . m t h s t r o n g a s s o r t t i v
hn thy wr rossd artifiiurlly,
nsistntwith Haldan's rul (f-
t a|. 1997). ln addition to this
from th . mth(fmal)x .
ny to mat (Davis et a|. 1997).
s rul may tnd ylnd th typ-
ility,and that hyrid fitnssmy
ostmtingbarrirs.
tzygotiisolatingarrirs is muh
nplis.of partiular intrstis th
ns ar gns that diffr twn
nthm.A gnalld odysses,
tion in sm fruit flis, dspit rl-
h i s g n a p p r st o r s p o n s i -
simulns and D. muriti (tng
rinrapid volution (at last in
nompatiilitistwntaa and
rs.Muh of th work on spi-
rolutionaryillogistsar ddrss-
rrtiulargns ar gnrally mor
whthrgnsassoiatdwith r-
:iationgnsthan othrs.
sltion,sxual sltion,and g-
n populations in phnotyps and
tnt that diffrntpopultions -
nativs.Why, thn, is not vry
: d d i f f r n ts p i s ? h r u o f
w muh gn flo,ours twn
:s to homognizpopulations oth
lorks against sltion and drift in
;iation, rsptivly.Sond' how
is thornir and points to th im-
iatiopross.Although divrg-
nsi barrirsto rprodution, som
prsnt to stop gn flow and
h Pttrn d roess of Spcitio 187
ompltth spiation proess. For allopatri populations' whih do not
ovrlapin gographi rang' th physial distan imposd y gography an
srvas strong arrir to gn flow. owevr, for divrging populations
whosrangsovrlap (sympatry)or ar adjnt to on anothr (parapatry),
or whn prviously isolated allopatri ppulations om into ontat (s-
ondaryontat),intrinsi or trinsi arrirsto gn flow that prvnt intr-
rding om nssary to omplt th spiation pross. Thus th
importanof rprodutiv isolating arrirs has n paramount in studis
o{ spiation,and muh of our undrstandingof th pross of spiation is
ntrdon th mhanisms that favor th volution of rprodutiv isolation
btwnspis.
lassi modls of th spiation pross fousd on th gographi distri-
utionof ppulations undrgoing divrgnnd th rlativ ontriution of
gographiisolation to th formation of nw spis(Figur 4). Although the
dgreof geographi sparation twn divrging popultions is rtainly
an important ontriutor to th disruption of gn flow, mor rnt studis
of spiationhav fousd on th importan of volutionary mhanisms
(naturalsltion,sxual sltion,gntidrift, and mutation) in promoting
divrsifiationand spiationithr in allopatry or in symptry.
H RoL oF NAURAL sLIoN lN sPE|AloN
Th imprtan of adaptiv divrgn in th spiation pross (trmd eo-
logilspitio)was fist notd y th gntiistThodosius Dozhansky
in th 1940s in dsriing spiation in Drosophil. Dozansky (1946) -
livdthat spiation ln Drosophila ourrd minly through th volution of
divrgntphysiologial ompls that wr sussful in diffrnt nviron-
mnts.rnst ar (1942) Was th first to point out that many of th umu-
latd gnti diffrns twn populations' partiularly in physiologial
and ologialadaptations to partiular nvironmnts' Wr also potntially
rprodutivisolating mhanisms. Suh ..y-produt'' volution of rpo-
dutivisolation is th driving for hind ayr's lassi mod f viarian
allopatrispiation (Figur 4).
Mor rntlyth importan of ologial divrgnin promoting rpro-
dutivisolation ithr in allopatry or in sympatry has n rognizd, and
an importantrol for natural sltionin a varity of routs to spiationhas
omlar. Laoratory primnts (rviwdin Ri nd Hostrt 1993)
hav onfirmd that ologial spiation an our; primntal studis
that raisd fruit flis (D. melnogstr or D. psudoobsur) in diffrnt n-
vironmnts(varying tmpratursor food sours) hav dmonstratd that
som lvl of prmating rprodutiv isolation volvd twn lins raisd
in diffrntnvironmnts,whil no prmating isolation volvd twn in-
dpndntlins raisd in th sam nvironmnt. Ths primnts dmon-
strathow th pross might also work in natur.
In th simplst modls of ologial spiation, rprodutiv isolation
volvstwn populations inidntally as a y-produt of adaptation to
188 h ttr tld Prlssrlf Spitkl

altrnativ sltin rgr

tiv isolation has n r1
Viriane AlIopatri
Thes irds ar on of th
and hav evolvd an im1
funtion, in ordan v
hav om to oupy. Lo
,a\
(A)
(Grant et aL. 1976) ar'd tl
Darwin's finhs has vo
\ ",." ."/ spondn to hanging
\i/ i n t r s p 6i o m p t i r i o n
tions twn ak morp
n n o f m a t i n g s i g n a l si n
bhavioral rprodutiv i
PeripheraIlsoItesor D a r w i n . s f i n h s .T h p
Founder ffet (pith) and tmporl pr
h n g s i n a k g a p ;h .
vary as a funtion of
th strutur of song a

-*/B/
\
finhs on Santa uz Is
aks adaptd fo rushi
((\/-
\7 I o w a t s o f s y l l l r
sis with smallr ak
aus song plays a ntra
this linkag twn ak
its onomitant influ
Sympatri pls of y-produt volut
portane in spition of

PARALLL sP|l
oF oLoG|AL l

ParallIvolution of sim
ronmnts strongly impli
s p i a t i o n n r s u I tf o n
produtiv isolation that
4. lassi modls (lr rods) tlf spiatitln r asd on th dgr of
I-.igr-rr rlatd populations as a t
gographi spration s t driving fo of spiation. In viarian alloptry,
nstrl ppulations of a on-ontinuous, larg rang of a spis r
sparatd y virirr vnt (.g.,mountains, riv, oan). Diffrns in
nvironmntal onditions on ithr sid of th arrir to disprsl pronrot loal
adaptation and y-prldut rprtldutiv istlliltion, 1adingto th formtion lf would rapid in suh fou
nw spis (ayr 1942). Th prihral-isolats or foundr-fft modl of ould rsult in gntiror
spiatiln p()stultsgnti drift as th prirry mhnism that prrmots i s o l a t i o na n d r s u l ri n r a p i
divrgn n1spition. Priphral isolats or foundr vnts rlur whrt prrpulationsthat ovrlprn
pirrrd Outward from th dgs of thir distriutions to OuPy
lpr.rlatilr.rs spiationwhr rprodu
nw l-raitats,rr lvhn ir s.ral]numr of ir-rdividLrlsdisprs to nW loatior-r not geographiallysparat
nd stlish a nw population. ayr (1954) prtlplsithat gnti hang aompanid y assortativ
 Th ttrld Prlcssof Spition 189

altrnativsltiongims.A rnt ampl of suh y-produt rprodu-

tivisolation has n rportd irr Darwin's finhs of th Galdpagos Islands.
Thsirds ar on of th most lbratedillustrations of adaptiv radiation
and hav volvd an imprssiv 21rraof spializations in k form and
funtion, in aordan with th divrs fding nihs that diffrnt spis
havom to oupy. Long-trm fild studis y Ptr and Rosmary Grant
(Grrtt aI. 1'976)and thir ollagus hav showrr tht ak morphology in
Drwin's finhs hs volvd y nrans of natural sltion in pris orr-
spondnto hanging ologial onditions, inluding food availaility and
intrspifiomptition. Jffry Podos (2001) tstd for potntial orrla-
tions btwn ak morphology ar-rdsong strutur. Song is a major ompo-
nntof mating sigrralsin irds, and diffrnsin irdsong lad to prmting
bhvioralrprodutiv isolation in a varity of spis of irds, inluding
Darwin's finhs. Th produtin of irdsongs with variation in frquny
(pith) and tmporal proprtis (.g., sylll rptitions) rquirs rapid
hangesin ak gap; hn voal prforman apaitis ar prditd to
vay as a funtion of ak morphology. Podos tstdthis ida y ompring
th stutur lf song nd ak Inopholog1'in ight spis of Darwin
finhson santa ruz Island (Figur 5). H found ht spis with larg
aksadaptdfor rushing hard sdshd volvd songs with omparativly
lowr rts of syllal rpetition and narrowr frquny rangs than thos
speiswith smallr aks volvd for fding on smallr, softer sds. B-
aussong plays ir Irtralrol in rPodutiv isolation in Darwin's finhs,
this linkag twnak morphlogy sltdfor fding prforman and
its onomitant influnon song strutur illustrts on of the st xam-
pls of y.produt volution of rprodutiv isolation and its potntial im-
portanin spiatinof Darwin's finhs.

PARALLL sPIA|oN As A s|GNAURE

oF oLoGIAL sP|AIoN

Paralllvolution of similar traits in populations that inhait similar nvi-

tonmntsstrongly impliats natural sltion s i] ausativ agnt. Plll
speiationan rsult from th paalll volution of trits that dtrn-linr-
produtivisolzrtionthat volv patdlyin indpndntlydrivd, losly
ation r asd on tlr dgr lf rlatdpopulations as a y-produt of adaptation to diffrnt nvironmnts
spiation.In viarian allopatry,
lar rang of a spis ar
rins, rivr, oan). Diffns rn
n blri to disprsal pronlrt loal
. r i t l l' l d i n g t o t h [ o r m t i n o would rapid irr suh founding ptlpulatilnsauslf gntidrift. This
)lirtsor frlundr-tftmodl f ouldrsultin gr-rti
roganizti<lsthat ould inidntllyyild rprodutiv
r.rrymhnism that romots islationand rsultin rapid spiatin.Sympatrispiationours btwn
's or foundr vnts our wht-t
populationsthat ovrlapin gogrphidistriution.It is th only modl of
; rlf thir distriutirins to oupy spiationwhr rprodutivisolationvolvstwnpopulationsthat ar
dividulrlsdisprsto i'1llw lotion notgogrphiallv It is usull,vassoiatdwith host o haittshift
spirratd.
1) p<lpsdtht grrtihang a<lmpanid y assortativnratirrgon th nw lrostor in th nw haitat.
190 he Pttrn d Prlssof Spitio

a->\ 9
o
(Shlutr and Nagl 1995
from symptri spiation

t\*.***
4- te^ ulr nvironmnts have l
2l
.**,*-..'
gn in daptiv traits ;
uildup f assortativ m
ns in thos adaptivtra
h tlrrspin stiklbt
systm for th study of p;
9
o ak is a marin fish fol
4- Alaska and British Columl
2r_ **#& #, This spis is anadromot
t h j u v n i l S t g i n m a r i
6a adults tl brd. Indpnd
=---t

'-
4|_
2L-
*
4' ** &** to multipl stram-rsid
th oan. In addition,in
nizations y marin stikl
ir ttt&L b th rrt of th glairs

-t
2L pndnt volution of lak
r I a k s .S p i a t i o ni n s t i

LLL
nw envionmnts and th
;F
-a\
L ody-siz diffnsarrd

;| \ Wb plts usd to filtr food 1

a h n v i r o n m n t .H e v t
o twn diffrnt stikl
o '|- In a rnt study Jffry
-- 2t- t } } B.,Bf,n found stong vidn of a

+
prfrns for mal ody
8

%-*.
6l-
4-
kHz 2l

0.5s

Figur 5. An illustrtionof v-produtrprodutivisolirtionas a sultof

adptivdivrgnin ak lnlphologyours in Drlvill.sfinhs.Bak
morphologyand rprsnttiv sound sptogrmsof solrgsfror.rr
ight
Dilrwirr'sfih spisotl Santarrzlslnd (frm ry trl btltttlllt:Gospiz
tltgirostris,G. iortis' (}. Iuligis, . st]lts'rhhus prulus'
. psittul, . pllid' . tlliu)ar shown. Sgltrogrmfquny
rsolution,98l{ sal.lr,0.5s.Intrspifivrrilrlis llpprntin oth
rr.r<lphologyand song stutur.Spiswith largks(.g.,G' mgirostris\
hav lowr ratsof syllalrptitionand nrrowrfrqunyrangsthan thos
Figur 6. Paralll spiatio
spiswith smallraks (.g.,. oliue).(From Polos2001.)
d t r m i n r p r o d u t i v is o
adptation to diffrntnv
twn otypes from th
(width inrasswith inr
etwndiffrntotyp
 The Ptter d Pross of Spition 191

(Shlutand Nagl 1'995).Th st ampls of paralll spiation om

from sympatri spiation vnts whr indpndntolonizations of parti-
ular nvironmnts hav ld to similar, rpatd pattrns of haratr divr-
gn in adaptiv traits assoiatd with diffrnt nihs and th paralll
uildup of ssortativ mating y seltivnvironmnt asd upon diffr-
nsin thos adptiv trits (Figur 6).

h*frn* Th threspin stiklack (Gsterostus uletus) as om a modl

systmfor th study of pralll spiation. Th anstral thrspinstikl-
ak is a marin fish found aross a wid gographi distriution from
Alask and British olumbia through lland, Sotland, Norway, and Japan.
tflbs This spisis nadromous, ing spawnd in frshwatr strams, spnding
th juvnilstag in mrin nvironmnts,nnd migating ak to strals as
adults to rd. lndpndnt olonizations of stram nvironmnts hav ld
&e*' s*-o to multiple stram-rsidntstiklak populations that no longr migrat to
th oan. In addition, in British olumia and Alaska indpndnt olo_
niztionsy mrin stiklaksof postglaial frshwatr laks formd aftr
Fbtehb th rtrat of th glairs at th nd of th Pliston hav ld to th ind-
pndntvolution of lak forms (nthisnd limntis)in multipl fshwa-
tr laks.Spiationin stiklakshas involvd th rptdolonization of
nw nvironmntsand th aomparryinghangs in adaptations, primarily
L L * tr* ody.sizdiffrrrsand numr and siz of gill rakrs (spins on th gill
h tr \ p|tsusd to 6ltr food patils),ssiatdwith ologilrl diffrnsin
ahnvironmnt.Hav thsadaptiv hangs ld to rprodutiv isolation
twndiffrnt stiklak otyps?
In a rntstudy Jffry Kinnon and ollagus(Kinnon t al. 2004)
t}' t&.$'?& found strng vidn of assortativ mating y siz through diffrntial fmal
prfrnsfor ral ody siz in altrnativ stram and mrin nvironmnts.

t"
.Tfl

lutiv isolatitln as a rsult of

rs in Dlvin's finhs. Bak
]granls li songs flrn ight
,from
top tl bottottl: (}ospiztt
lds, dmdrh b u,s1lruulu s'
rvn. Sptrogram frquny
varilrtion is aprnt in otlr
larg aks (.g.,G. mgnirostris)
rIowr frqur-ryfangs thn thos
Figur6. Paralllspiationtrnrsultfrom th parlllvolutionof traits that
(From Pr-lls2001.)
dtmin reprodutivisoltiontht volvreptdlyas a y-podutof
adapttionto diffrnt nvir<lnmnts.
Solid lines are usd to show matings
btwenotypsfrom th sam habitat with a gratrdegrof ompatiility
(widthinrasswith inrsdompatiility). Dshd lies tndiatematings
etwendiffrntOtypswith a lorv dgrof mting omptibility.
192 Th Pttern d Proess of Spitio

Anadromous marin populations ar lagr in ody siz than stram-rsidnt

populations, prsumaly an advantag in th long-distanmigrations that
marin populatins undrgo. Mating omptiility Was mor than tr,l,otims
highr whn fmals W paird with mals from th sm otyp (from
British olmia r Japan) than whn thy wr paird with mals from a
diffrnt otyp. To tst th importan of ody siz as th asis for ssor-
tativ mating y otyp,Kinnon and ollagus(MKinnon t al. 2004)
rard stiklaksfrom stram-rsidntand marin populations in th lao-
ratory and manipulatd thir ody sizs y hnging th lngth of th grow-
ing sason. This produd ithr larg-odid or small-odid fish of oth
sxsin oth mrin and stram-rsidntotyps.In mating trials with ths
primntally manipulatd fish, thy found that mals and fmals had
highr mating ompatiilitis whn thy wr mathd for siz, rgardlsso{
their eotyp of origin. Ths rsults dmonstratth importan of a singl
Ass<
phnotypi trait, ody siz, in adaptiv divrsifiation in stiklks and ph
suggst that divrgnt natural sltion on this trait in diffrnt rrvirontnnts
also maks a primary ontriution to rprodutiv isolation that lads to
spition.
{
H RoL oF NAURAL sLloN lN Hos
sHIFs AND sYMPAR| sPElA|oN

Th ourrnof sympatri spiation,spiationwith<lutgographiisola-

tion (igur 4), has bn ontntious aus th onditions undr whih Figur 7. Sympatrisp
sympatri populations an sap th homgnizing ffts of gn flow wr failitatdy host or ha
thought to rar and rquir eptional irumstans.Mor rntiy th divrging phnotyps.Tl
rprodutivisolationv
rognition of disruptiv natural sltionaompanid y strong assotativ
ir-rdividuals
tln th nw hr
mating of divrgntlysltdphnotyps (Figur 7) as a likly modl for th spiationprosswithtl
origin and divrsifitionof sympatri spishs onvind many volution-
a r y i o l o g r s t st h t s y m p t r i s p i e t i o ni s n i m p o t n t n d v n p r i m a r y
typ of spiation in rtin groups of organisms. Impliit in th pross of
sympatri spition is rapid volution of rprodutiv isolatilntwndi- primary haratristit
vrging forms ausgography annot srvas th prirrraryarrir to gn host shifts in Rhgoltis
flow, as it dos for divrgnt allopatri popultions. On onrmon way in rd on th sam host
whih suh rprodutiv isolation n rapidly volv is thrtlugh host shifts y spis is low. For xam
parasitesor phytophagous inststhat spializin partiular host spis. ing ggs on appl trs
Guy Bush providd th first potntial ampl of a sympatri spition oviposition sits.This st
vnt initiatd y a host shift with his studis (|966,1969) of th host ras twn hawthon and
of hwthorn nd appl maggot flies (Rhgoltis |lomollrl). Hawtorn trs u m u l a t ds i n t h i n i t
ar nativ to North Amria, and th hawthorn ra of R. pomolla is th nology, suh as timing
anstor of th appl host r. Appl maggot flis wr first dislvrdon a d u l t s ( . g . ,F d r 1 9 9 8
introdud, domsti appl trs in th Hudson Vally rgilnof Nw York in natural sltion as th
th mid-1800s. Bush and his ollagus hav n studying diffrns- fruiting phnology t
twn th ntiv hawthorn ra and th appl ra sin th mid-1960s and diffrns y ah hos
hav doumntd a numr of phnotypi diffnsin lif history and r- th host ras and ug
prodution that hav failitatd spiatilntwnth two host ras. on nes. Thus th Rhgol
 Th Pttr d ross of Specitio 19

in ody siz than stram.rsidnt Disruptive

seletion
h long-distanmigrations that
t i i l i yW e sm r t h a n t w o t i m s
ls from th sam otyp (from
y wr paird with mals from a
f body siz as th asis for assor.
ollagus(Kinnon t al. 2004)
.d marinpopulations in th lao-
hanging th lngth of th grow-
Jid or small-odidfish of oth
otyps.In mating trials with ths
ind tht mals and fmals had
r mathdfor siz, rgardlssof
nstratth importan of a singl
Assortativmating within
ivrsifiationin stiklaks and phnotypsor habitats
. h i st r i ri n d i f f r nt n v i r o n m n t s
lrodutiv isoltion that lads to

oN IN Hos
'toN

iationwithout gographi isola-

aus th onditions undr whih
gnizingfftsof gn flow wr
irumstans.or rntly th
r o m p a n i dh s r r o n g s s O r t t i V
(Figur7) as a likly modl for th
is has onvindmany volution-
, a n i m p o r t a n t n d v n p r i m a r y
1anisms.lmpliit in th pross of
produtivislation twn di.
rv as th primary arrir tO gn
p u l r i o n so. n o m m o l l w a y i n
dly volv is through host shifts y
i a l i z i n p e r r i u l rh o s s p i s .
. m p l o f a s m p t r i i t i o n
l i s ( 1 9 6 6 ,\ 9 6 9 \ o f t h h o s t r a
oltis pomonll).Hawtorn ts
rthorn r of R. pomlll is te
tggot flis wr first disovrd on
dson Valiy gilnof Nw York in
rav n studying diffrns -
ppl ra sin th mid-1960s and
; diffrnsin lif history and r-
r twnth two host rs. on
Figur 7. Sympatri spiation an Our whn disruptiv natual sltton
failitatd y host or hitat shifts is aompanid y assortativ mating y th
divrging phnotyps. Th symptri modl lf spiation rquirs that
..lik''
rprodutiv isolatiln vllvss a rsult of ssortativ mating f
individuls on th nw host or in th nw hitat in odr to initiat th
spiatilnpr()sswithtlut gogrphi isolation.
primary haratristithat promots sympatri divrgn assoiatd with
host shifts t Rbgolti.sand othr phytophagous insts is that thy fd and
rdon th sam host spis;thus disprsal twn diffrnt host plant
s p i iss l o w . F r a m p l .f m a l so f h a w t h o r no r i g i n r a v r s t o o v i p o s i t -
ing ggs on appl trs, whras appl-origin fmals prfr apple trs for
ovipositionsits.This strong host fidlity limits intrhost movmnt of adlts
s twnhawthrn and appl trs. In addition, gnti diffrns hav a-
umulatdsin th initial host swith that promot hangs in rdingph-
nology, suh as timing tlf mating, pupation and diapaus' and losion of
adults(.g.,Fdr 1998). Ths hangs ar thought to hv n favord y
natural sltionas th diffrnt host ras of flis adaptd to diffrnsin
fruiting phnology twnhawthorn nd appl hosts. Suh loally adaptd
diffrnsy ah host ra ontriut to rprodutiv isolation btwn
th host ras nd augmnt host fidlity through fmal oviposition prfr-
ns.Thus t Rhgolris flis provid fundamntal ampl of ologial
194 Th Ptter nd Pross of Speitio

divrgnwhr omplt gogrphi isolation is not lways rquird to ini- prfrnsfor onspi
tiat spiation. of D rosolt h iltt psendolts
Although phytophgous insts offr svral wll-doumntd ampls of lutlne to mat with ht
sympatri spiation though host shifts, a rnt study provids at last on popultions (Noor 199.5
mpl of th potntial for sympati spiation via a host shift in a vrt. frtil, allowing flr gn
rat' Afrian indigoirds (gnus idu) ar host-spifirood parasits, on of th st-dotr
laying thir ggs in th nsts of firfinhs (gnus Lgonostit)' and diffrnt t h i a n d l i m n t i o t y p
indigobird spis spializ in pzrrasitizingdiffrnt spis of firfinhs.In- ak. oward Rr"rrrdl:
digoird nstlings ar rard along with th host young and mimi th mouth lor21toryrosss anl ir
markings of thir rsptiv hosts, thus lowring th proaility of ing r- rids suffr a foraging d
jtdy thir host fostr parnts.In addition to this uniqu mimiry adapt- prntal nvironmnts.It
tion y young indigobids, as adults, mal indigoirds mimi th song that is vironnrnt with rmpl
sung y thir host spissir, and fmlsrard by a spili host spisof parrrtal spis.Horvv
firfinh prfrntially mat with mal indigoirds that sing thir natl host- parr-rtlnvironmnt, lit
spifi song. This failitats host-assortativ mating and lads to rprodu. rats :rnd wr smallr at
tiv isolation among diffrnt spis of indigoirds that parasitiz diffrnt Wr poor at thing
host spis of firfirrhs.A rnt phylognti study y ihal Sornson fild nvironmnts iru
and ollagus(Sorensont al. 2003) that ompard n-litohondrialDNA s- In adlitiol-l' nthi f
quns for a numr of diffrnt spisof indigoirds and thir rsptiv likly to mat witlr irrtrr
firfinh hosts dnronstrtd that diffrnt indigoird spis that parasitiz mals from al|opatri p
diffrnt firfinh hosts ar gntially diffrntiatd from on anothr, a sults support a modl of
finding onsistntwith assortativmating of indigoirds rard y a partiu- t h r o u g l l n t r l r ls | t i o
lar firfinh host spis.Thse rsults suggstthat divrgnthrough loal fitnss in ithr parntlll t
adaptations in nstling mouth markings, adult nrl song opying, and f- How impotant h.rs r
mal prfrns for host spis song typ has ld to positiv assortativ h p t l | mi s t h l t r i n
mating twn diffrnt host-spifi spis of indigoirds. This pattrn of pattrns of strongr div
host-spifi adaptations linkd with positiv assortativ mating within a tions. Imagin that two s]
host typ is onsistnt with sympatri spiation irr Afrian indigoirds gr of prldutivisrll
through host swiths in th firfinh spieswhos nsts thy parsitiz. I i k l v o r l t i n u t ( ) ( ) i s
ls of produtiv isolat
or th tintion oi th
RPRoDU|V lsolAloN MAY B RE|NFoRD
spis pairs) that w ol
Y NAURAL sL|oN
volvd a high dgrof
Natural sltionan at to dirtly inasth mount of prmating isola- following sondrv o
tion btwn two divrging populations by slting against hyrids formd v o l r ' di n m a t i l l g l i s i o
twn thm. This pross' first proposd Thodosius Dobzhansky drrrrinsth prssof rt
(197) and trmd riforcemelby Frank Blair (1955 ), an our in arasof romot divrgnirr rn
sondary ontat whn prviously allopatri populations om into ontat must tightlv linkd wi
and intrrd. Whn hyrids twn formrl allopatri populations ar ity. Tlrs quirmnts
partially infrtil, rinformnt n omplt th spiation pross. Rin- Ths argumnts do n
formnt was popular onpt ftr th modrn synthsis aus it allows plting th spiatilnpr
natural sltion to driv spiatin dirtly, ut its mpirial dmonstration that th vidn for tn '
has n diffiult, and its istn as an important omponnt of ologial siv. of th two pross
spiationhas n ontrovrsil(.g',Noor 1999; arshall t al. 2002). rr'ith ologiel s1rgi1
Most of th vidn for rinformntoms flm omparisons twn rinfo:tnt-th first i
allopatri and sympatri popultions in th strngth of fnral mating spiation; th irnpotn

lation is not always rquird to ini_
',rlwll-doumntdxampls of
rnt study provids at last on
iation via a host shift in a vft-
r host-spifirood parasits,
(genusL gostit), and diffrnt
3 diffrntspis of firfinhs. In.
rhost young and nrimi th mouth
,wringth proaility of ing r-
:ion to this uniq mimiry adapta-
indigoirdsmimi th song that is
; rard by a spifi host spies of
ligobirds that sing thir natal host.
tiv mating and lads to rprodu-
ndigobirds that parasitiz diffrnt
gntistudy y ihal Sornson
.onrpardmitohondril DNA s-
o f i n d i g o b i r d sa n d t h i r s p t i v
t i n d i g o i r ds p i st a t p a r a s i t i z
liffrntiatd from on anothr, a
o f i n d i g i r d sr a r d y p t i u -
ggstthat divrgn through loal
adult mal song opying, and f-
yp hs ld t positiv assortativ
:isof indigoirds.This pattrn of
'sitiv assortativmating within a
spiation in Afrian indigoirds
:is.hsnstsrhy prsiiz.
r R|NFoRED
as th amount of prmating isola-
ly sltingagainst hybrids formd
'osd y Thodosis Dobzhansky
k Blair (1955),an our in aras of
rtri popultionsom into ontat
rmrly allopatri populations ar
rpltth spiatin pross. Rin-
: modrn synthsisbaus it allows
tly, ut its mpirial dmonstration
important omponnt of ologial
> o r1 9 9 9 ; a r s h a l l e t a | . 2 0 2 ) .
t oms from omparisons twn
n th strngth of fmal mating
h Pttrn d ross o| Spcitit 1 9 . t
prfrnsfor olspifis.For irmpl, fmals drivd from populations
of Droslphil 1lseudotlbst,lrasvrrrpatriwith D . persimilis lisplay grat r-
lutanto nlat with htrospifimls than do fmals 1rorn aIlopati
populations(Nolr 1995)' F1 hyrid mals ar stil,ut hyrid fmals ar
frtil,allowing flr gn hang and sltionagainst hyrids.
on of th st-doumntdirmplsof rinformntours in th n-
thi and limnti OtYpsfound in lak populations of th thspinstikl_
b a k . H o w r d R u n d l a n d D o l p h S h l u t r ( 1 9 9 8 ) d m o n s t r t d o t h i n
laoratoryrosss nd in F1 lrr.bridsdttd in wild populations that hy-
rids suffr a foraging disadvntagrlativ to thir parntal spis in th
parntalnvirlnmnts.In th laoratory F1 hybrids rard in a ommon n-
vionmntwith ar-rrpl assto food showd growth rats similar to thos of
pantalspis.Howvr, in fild studis F1 h,vidsrard in itlrtyp of
parntalnvirlnmnt,littoral :rnd limnti zons of laks' had lowr growth
ratsand wr smller at maturity than ithr pur parntal spies.Hyrids
Wrpoorr at athing nthi invrtratsand plankton in th diffrnt
fild nvironmtsausof intmdiatgill-rakr numr and ody siz.
In dditiln, rrtlrifmls from svmpatri poptrltions wr rrruh lss
likly to mt rvith intrmdi:rt-sizd F1 hyrid mls than rvr nthi f-
mals from allopatri populatilns whr limntis did not our. Ths r-
sultssupport a modl of rinformntof siz-assortativmating prfrns
through natural sltilnagainst intr-rdiat-sizd hyrids that hav lower
fitnssin ithr prntl environmnt.
How irrrpotanthas rinf<lrtnnt n in ss of ologil spiation?
Th prolm is that rinforrnntis not th orrly plnation for osrvd
pattrnsof strongr divrgn in mting prfrnsin sympatri popula-
tions.Imaginthat two sympatri populations had alrady volvd a high d-
gof rprodutivisolation wlril still in allopatrv. Ths populations will
likl,vontinu tO Oist,whil formrlv alloptri populations with low lv-
ls of rprodutiv isolation n121y lost ithr tlrr<rugh gn Ilow and fusion
or th tinti<lnof th rarr population. Tlrus th only poprrlatins (or
spispairs) that w lsrvliving in sympatry in natur ar thos.that
voivda high lgrof isolation in allopatry, not usof rinformnt
follwing sondv ontat. In addition, ronrir-rationtwn loi in-
volvdin mrtingdisions rrd th hratr(s)ir.rfluning hyrid fitnssurr-
drminsth prss of rinformnt.In ordr f rinformntto proprly
promotdivrgnin rntpr[rns,gnsthat od for mating disions
must tightlv linkd with gns that od for hyrid infrtility or inviail.
ity. Ths rquimntsmay not oftn mt in ntural populatiolls.
Thsargumntsdo not shw that rinformntl.rsnot oprtd irr om_
pltingth spiti<-ln pross during ologial spiation.Tlry only suggst
that th vidrrfor th widsglradimportan <lfrinformnr is inonlu-
siv.of th twl prosssthat an promot rprodutivisolation assoiatd
with logial spiation-divgn with isoltion as a y_produt nd
informnt-thfirst is wll doumntdnd almost rtainly irnportant in
spiatiorr; tlr importan lf th sond still rrdsfurthr strst:rrrtiation.
196 The Pttern nd Prrl.,sslf SDeitil

akgrlund radian i
sxUAL sLE|oN AND oLoG|AL sPlA|oN
ovrall rightnss of tJ
Sxual sltion' a for that ats on traits that inrasmting suss'an m n t h i r v v t l l v d r i
inflund y th nvironmnt and' as suh' miry ontriut to ologial nd tallsmissiun t |
spiation.Sxul sltionthat lads to spitionirs a rsult of divrgnts- hitats'whr th a
ltion twnnvironmnts iln rsuit fronr sptial viation in sonday i n d w l . r ps i g n a l sw l l u
sxual traits, suh as linal variation in mal plumag or oloration in r- wirlr tlrWrongdw|.r
spons to a vaying nvironmntal gradint. For ampl, linal variation in and hrrlss likly t
th availaility of foods high in arotnoids that ld to right rd and or-
ang oloration in mal irds has n suggstdas th primary aus for di-
H RoL oF s
vrgn in plumag among popultions in mal lrous finhs, rpodus
NoNoLoG|
mius (Hill 1994). A rr-ro{lthat dmonstratsth importan of praill
linal variation in fml mting prfrnsas a sult of nvironmntal o d l s t r f s u a Is |
variation in mal signling traits was first proposd y Russll Land (1982), mating prfrnsfor
ut good ampls of linal variation that lads to spiation via sxual s- u [ r i o n s n d i v g l l
ltion ar laking. and spiation. Suprf
Natural sltion through variation in fatursof th nvionmnt whr s p i e t i l l l . p t t h i t
mating signals ar propagtdn sult in divrgntsxual sltionof mal m l n l t i n g ,s i g n l r l sr h
mating signls' suh as ourtship displays or nuptial oloration' lading to p r o p r i st o t h s i g n l I
prmating isolation twn populations from divrgnt signaling nviron- p r o m < l d h r t ' u g hd i v t
mnts (Ryan and Rand 199; Boughman 22).Natural sltion an also on modl of fma
at upon th snsory systm of fralsas wll, lding to snsory iassfor runWa pross (or ..
rtain proprtisof ml signals (.g',brightr oloration, largr siz, loudr F i s h r i n p o s s sa r
and longr durtion of volizations). Partiular faturs of lal nviron- tlr tttlstxrm rtr.tl
mntal onditions ould altr sensory ias or mk som mal traits ttrat p r J i s o r t h i g h r
ploiting biass in rtain nvironmnts than in othrs. In addition, fmal tion of ths traits ris
mating prfrns for mal traits that ar indiators of suprior quality an that nod th attrat
also inflund y th signaling nvironmnt' For xampl, if partiular fo that teit. Fisr.s
ml traits ar mor rlil indiatos of quality in som nvironmntsthan a n d t h i s u n d a n tt
in othrs (Shlutrand Pri I99), divrgnirr prfrnsbtwn nvi- promoting th volr.rti
ronmnts for thsmal traits ould lad to prmating rprodutive isolation 1994); howvr, studi
twn thm. Thus diffrnt ovolutionary paths of signals nd prfrns ult to pinpoint. On
in diffrnt nvironmnts ould ]nratspiatiorl. span length in stalk-
Th st xampls of an importirnt rol for sual sltionin prolnoting Grald Wilkinson and
spiation as rsult of varying signaling nvironrIrtsom from spis wild flis' a slrort-y-
that liv in environmnts that vary in light irrtnsiryor transmission propr- o s l i o n .t h y f o L r n
tis that influnth sptral quality of signals (i..'whih olors irr st r- a I l y s I t dm a | t r e i t
fltd y partiular nvironmnts).For xampl, allopatri populations of from th lin sltdf
Aolis cristtlls lizards our in two diffrnt nvironmnts (ri and with short y spans' .
msi) that diffr with rspt to light intnsity and th sptral quality of prfrr1mals with l
olors tht r st fltd.nul Ll and Lo Flishman (2004) hav r n s i o r - s p a nl t
shown tht mal anols frorrr th two nvironrnts diffr in th olor and s t a l k - y df l i s t h e t d i
pattrn of thir dwlaps (folds of skin undr th throat that ar distnddin S x u l s l r i nm l
mal displays usd aS tfritorial nd mating signals)in th dirtion that in- g s \ .U n d t h i s m o d
rasssignl dttailityin ah haitat. als fom th xri nvironmnt fmal lvith informatil
hav darkr and slightly rddr dwlaps that miriz ontrst whr th mk mating dision

oG|AL sP|A|oN
ts that inrasmating suss'an
suh, my ontribr.rtto ologial
piationas a rsult of divrgnt s-
from spatial variation in sondary
m a | p l u m a g o r o I o t i o n i n r -
)nt.For trrpl,linal vaition in
lids that lad to right rd and or-
ggstds th primary aus for di-
in mal hous finhs, rpodus
ionstratsth importan of paralll
ns as a rsult of nvironmntal
proposdby Russll Land (|982)'
at lads to spiation via sxual s-
fatursof th nvironmnt whr
in divgntsual sltionof mal
ys or nuptial oiortion, lading to
s from divrgntsignling nviron-
o 2002\. Natural sltion an also
s wll, lading to snsory iass for
ightroloration,largr siz, loudr
'artiilla fturs of loal nviron-
s or mk som mal traits ttr at
i than in othrs.In addition, fmal
indiatorsof suprior quality an
:onmnt.For ampl' if partiular
f quality irr som nvironmnts than
rgnin prfrnstwn nvi-
to prmating rprodutiv isolation
lary pathsof signalsand prferns
spiation.
I f o r s t r a sl l t i o ni n p r o m o t i n g
tg nvironmntsom from spis
h t i n r n s i r yo r t r a n s t n i s s i o np o p r -
;ignals(i..,whih olors ar bst f-
xampl,allopatri populations of
diffrnt nvironmnts (ri and
ntnsityand th sptral quality of
al and Lo Flishman (2004) hav
nvironmnts diffr in th olor and
rdr th throat that ar distndd in
ing signls)in th dirtion that in-
t . l sf r m t h x r i n v i r o n m t r t
; tht maimiz ontrast whr th
Th ttr d Prrlssrlf S1litiltt 1 9 7
bakgroundradian is rltivl high (i..,thy will app?1darkr than th
ovrllrightnssof th akgrund), whil rals from th msi nvtrtln-
mnthav volvd right dwlaps ausof thir rlativly hig fltan
and transmissiont all wavlrrgths.Suh mls ar mor dttalin msi
hitats,whr th bkgrounl haitat light lvls ar low. Tlris divrgn
in dwlap signalswould minimiz gn fltlw twnhaitats, ausmals
with th Wrong dwla olor pattrn would lss dttl,vonspifis
and hnlss likl t() sura titory or otin nltings.
H RoL oF sxUAL sLE|oN IN
NoNoLoG|AL sP|A|oN
Modls of su;rl sltion, partitrlarly mtldls of th vollttioll lf fml
matingprfrnsf<lrml trits (Mad anl Arnold 2004I, slrlwthat pop-
ultionsan divrg in ths prfrns,prlrrrotingrprodtrtivisoiation
and spiation.Suprfiially,this pross mirrors that osrvd in llogial
spiation't that instd of natural sltion promoting divrgn in
ai mating signirlsthrough sltion against hrids or rathing of signl
proptisto th signalingnr,ir<lnnrl1t' th sltivfor is sual sltion,
prlmotdthror.rghdivrgnt [rlmating pfrs.
on modl of fmal hoi that nray ld to spiation is th Fishrian
runawy pross (or ..sxy sons''), first proposd y Ronald Fishr (190).
Fishrianprosssar oftn itd as th asis for th volutilrrof som of
tlrnrosttrm maI traits, sulr s th loratpltrmagof mal irds of
pradisor th bright olors of ml tropial tishs.Th rrm aggra-
tion of thstraits ariss as a rsult lf th gnti orrltion twn gns
thatnodth ttrativmal trait and grrstt ntldfrrralprfrns
for tht trit. Fislrr's runWy Pr()sshas l-lmodld -rathrnati:rlly,
alrdthris aundarrtmpiril vilnflr th rll of Fishrian sltilnin
promotingtlr volution of ggratdrnal traits within spis(Andrss<lrr
1994);howvr, studisthat support its xlusiv rol in spiirtionar diffi-
ult to pinpoint. On potntial arnpl involvs artifiial sltioll tlf y-
span lngth in stalk-yd flis (rtodiopsis dImlni), | th lortlry
Grald \Wilkinso and Paul Rillo (1994) ratd two linagsof flis from
wild flis,a short-y-sparr lirrirnd a long-v-spanlin. Attr l3 gnratiorls
of sltion,thy found a gntilrrlationtwnnral y spn' a su-
ally sltdmal trait, and frnal prfrnfor y-span lngth. }.mais
from t lin sltdflr short tlal y spas prfrrd to mat with mals
rvith short y spans' and fmals from th lins sltdfor long y spans
prfrdrrr:rlswith iong v spns. Suh iivgnin f-r:rl mting prf-
rnsfor y-span lngth rrrightaount for sition lrlon spis f
stalk-ydflis that diffr in y-spanlngth.
Sual sltin may also at on indiator trits (also known as grlod
gs).Undr this modl of fmal hoi, xaggratdmal traits provid th
frlwith inflrInatiorrltrtrh l]tltiquality of th maI.all<lu.inghr to
mak mating disions asd on informati<lnthat will potrrtilly provid
198 h Pttr d Pross of Spitio

suprior quality to hr offspring. Although this mhanism may potntially Sxual sltion has t
lad to spiation if th indiator traits diffr twnpopulations, lik run- part aus of its dir
away sltion,mpirial xampls ar rar. Som vidn indiats that a 2 0 0 1 ) . S p i so f i r d s r
good.gns mhanism may promot diffrns in ml gnitalia that lad thought to xprin st
to divrgn in som inst taa, although this hypothsis is still undr d- monogamous linags;tJ
t (oskn and Stokle 2004).It should notd that for oth indiatr spios (Mitra et a|. 199
and Fishrian traits' th lak of onlusiv mpirial vidndos not mn of high sPitioI1 r a t si
that divrgnis unlikly to a produt of ths mhanisms;rathr, it is prisonsof dihromri
oftn diffiult to distinguish whih mhanism may hv oprtd in th past. with fmals dull) lads
An additional rol of sual sltionin spiationan trad to undr- usually right) lads (l
standing th mhanisms assoiatd with sual onflit. Sual onflit amountS of fthrornar
ariss whn th st stratgy for ln sx is diffrnt (or vn dtrimntal) hav n osrvd in an
ompard with th st strtgy in th othr s. This antagonism is pr- and othr invrtrats
ditd to lad to a ovolutionary arms ra whr diffring sts of traits - t a l . 2 0 0 1 ) . A r n t x
twn th ss intrat to dtrmin th natur of mal-fmal intrations rsult f sual sltio
(Arnqvist and Row 2005). An ampl of within-spissual onflit is Lupl. Tamr Mndl
th ffts of sminl fluid in Drosopbil melnogstr. Sminal fluid has spis at molular mar
n shown to nfiial to mal fitnss' rduing fmal rmating and in- tiv ags of th diffrnt
rasing ovulation, ut it is toi to fmals; th mor sminal fluid thy r- i n o n l d o f t h s r i
iv, th fastr thy di. Sual onflit apparsto our aross a wid rang spis pr million yars
of taxa (Arnqvist and Row 2005), ut prditaly,som of th st vidn iation in this group app
for a o1of sual onflit in spiation oms from insts (Arnqvist t al. song trill rat, whih f
2000). Baus sual onflit may lad to rapid volutin in th rprodu- appar to any olog
tiv trat, it follows that it may also inrasth pa of rprodutiv isola- rikt spis.Ths a
tion twnpopulations. tion an an importn
Prmating isolating bar
sXUAL sL|oN' lsoLA|oN'
AND RAs oF sPlAloN H RoL oF Ml

Sual sltion,ausof its rol oth in promoting th volution of pr. Muttional hangsass
mating rprodutiv isolating arrirs and in ontinud divrgn twn thought to a primar
spis,may hav onsidra1impat on th rat of spiation (rviwd rWhit (7978, 336) onl
by Panhuis t al. 2001). For xampl, taa with a wid rang of snsory pr- th primary rol in th
ption apailitis may b partiularly pron to spiation through sxual widsprad rol of kay
sltion, sin thy hav n lvatd aility to produ or dtt a road aus th aumulation
rang f signals, thy visul' aousti, or phromonal (Andrsson 1994). t h t t u g h t t h l a r g , l i r
Spis-rih groups f frogs' for instan, an dtt a widr rang of problm with th viw
advrtismnt-all frqunis than thos in lss spios linags, potntially tion ntrs on th diffi
aounting for why suh linags of frogs ar mor spios ompard with lations rhing fiatto
thos with narrowr frquny prption rangs (Ryan 1986). In addition, ltrnativ rarrangm
rinformnt,as dsridarlir, an omplt th spiationpross upon rids, thry rduing
sondary ontat y nhaning divrgn of fmal mating prfrns. mosoml rarrangm
Sxual sltionplays an important rol in sympatri spiation as wll. As- trm hromosomal r
sortativ matin8' th propnsity to mat with lik individuals, hs n pro- in hangs in lrromos
posd s on way to quikly volv rprodutiv isolation (lyn and orr hromosrmal rarrang
2004) and is a nssaryomponnt of th sympatri spiation pross. ar infftiv rrirs1

gh this mhanisnrmay potntially
.ffrtwnpopulations, lik run-
rr. Som vidn indiats that a
frnsin mal gnitlia that lad
gh this hypothsisis still undr d-
rld notd that for oth indiator
mpirialvidndos not man
;t of thsmhanisms:rathr. it is
nisrrrmay hav opratd in tlr past.
r spitionan trad to undr-
th sr'rlonflit. Sual onflit
is diffrnt(or vn dtrimntLrl)
o r h r s . T h i s . r n t g o n i s mi s p r -
l wlrr diffring sts of traits .
n a t u r o f m l - f m l i n t r a t i o n s
of within.spissxual onflit is
l mllgsler.Sminal fluid has
s, rduingfmal rmating and in-
rls;th mor sminal fluid thy r-
p p r st o O u ra r o s sa w i d r a n g
'ditaly, som of th st vidn
orsfrom insts (Arnqvist t al.
to rapid volution in th rprodu-
astI]pa of rpodutiv isola-
{,
in promoting th volution of pr_
rd in ontinud divrgn twn
on th rat of spiation (rviwed
:a with a wid rang of snsory Pr-
pron to spiationthrough sual
lility to produ or dtt a oad
, or phromonal (Andrsson 1'994).
n' an dtt a widr rng of
in lssspioslinags,potntilly
's ar mor spios ompard with
ln rangs(Ryan 1986). In addition,
mpltth spiationpross upon
:n of fmal mating prfrns.
in sympatrispiation as wll. As-
with lik individuls, has n pro-
:odutivisolation (oyn and orr
1symPatispiation pfoss.
Tll ttrn d Process of Spitio 199
Sul sltiorrhas n linkd tl rlativly higlr rts of spiation, in
part aus of its dirt fft on mat rognition traits (Panlruis t al.
200l). Spis of irds with highr lvls of promisuity, for ampl, ar
thouglrt to prin strongr sual s.ltionthan losly rlatd, mor
moogamous linags;ths promisuolls linags gnrally ar also mor
spios(itra et al'.1'996).othr studis of irds oform to th prdition
of high spiation rats in taa with strong sxual sltion,inluding om-
parisons of dilrromati (ss of diffrnt olors; nrals gnrally right,
with fmalsdLrll)lads vrsus monohromati (otlr ss th sam olor'
usually right) lads (Brralough t l. 1995) and lads with vaying
amountsof ftrornamntation (ollr and urvo 1'998).Similar trnds
hav n osrr,din angiosprnrs(nd thir assoiatdpollinatos),insts
nd othr invrtrats,arrd fishs (rviwd in Andrsson |9941 Pnhuis
t a l . 2 0 0 1 ) . A r n t x a m p l h i g h l i g h t i n gt h r a p i d r a t o f s p i a t i o na s
rsrrltof sual sltion hs bn shown in Hawaiin rikts in th gnus
Lupl. Tamra Jv{ndlsonand Krry Shaw (200.5)usd divrgnamong
spisat rnlularmarkrs liratd with gologil stimtsof th rla-
tiv agsof th diffrnt Hawaiian islands to show tht th rat of spiation
in on ld of thsrikts from singl islnd was alulatd at ovr four
spispr million yars,th highst rat vr alultd for arthropods. Sp-
iatio i tlris group appars t<lb promotd y diffrnsin ml outship
songtrill rt, whih fmals us in spisrognitin, austhr do not
apparto arry ologially distinguislringfaturs that spart diffrnt
rikt spis.Ths and many othr ampls highlight how sual sl-
tion rr n irrrportntvolutionary flrtht lads to raid divrgnin
prmatingisllirtingbarrirs' rsulting irr high rats of spiation.
HE RoL oF MUA|oN lN sP|A|oN
uttional lrerngs
assoiatdwith hromosomal rarrangmntswr on
thought to primary nrhanism of spiation. For ampl, . .}. D.
whit (1978, 336) onludd that hromrsomalrarrngmntshv ..playd
th pimary rol in th majritv of spiatiln vnts..'or rntly th
,idsprad rol of karyot,vpihang in spiatiorrlras n qtlstiond,.
usth aurnulation of hromosoml diffrnstwn populations is
thought to largly inidntal to spiation (Sits and oritz 1987). The
prolm with t viw that hromosomal rarrangmntspromot spia-
tilnntrson th diffiulty f ltrnzrtivrrrangmntsin divrging popu-
ltionsrahirrgfiation. All modls of hromosoml spiation assum that
altrnativrarrangmnts impair th frtility or viaility of intrspifi hy-
rids,thrrduing gn flow and promoting fiation of altrntivhro.
mosomal rrrnglrrts.Although tlris nry iikh' tru in rples of
trmhrorlsomalrrratrgmnts(suh as lutational vnts that rsult
in hangsin l.rromosomnumr)' numbr of studis hav shown that
homosomalrarrangmntsOftn hv littl fft on frtility nd hn
ar irrfftivairstO gl1flow (Spiito |998),
200 Tbe Pttr d Pross of Spitio

Lorn Risrg and ollagus hav studid rominational spiation in ontirrs diploid sp
sunflowrs (Helitbus spp.) and th ffts of hromosomal rarrangmnts spis Hl uersiolor
in ontiuting to th spiation pross.Rominational spiation ours lopilty (H. hrsoselis
whn two diploid spis intrbrd and giv ris to a nw linag that is northl.l lralf), ut thy
both frtil and tru rding br.rt is rprodutivly isolatd from both tlrir rarrgsovrlap. Di1
parntal spis.Risrg (2001) argus that hromosomal rarrangmnts produtivly isoltd frt
that ar htrozygous in hyrids hav majr ffts on supprssing romi. nl diffrns) and p
nation twn hromosoms that ay th rarrangd sgmntsand gns intrrding. als pl
linkd to ths sgmntson th hromosoms. Gns unlinkd ro th ho- rats, with th dil<lidh
mosomal rarrangmnt sgmnts,spially those that ar nutral' frly in- r i l t 5t h a n t h o s t l f t h t
trogrss twn the hyridizing parnt,al spis. Rarrangmnts that l Grhardt arrd oll
supprss rombination ut lak a ausal fft on hyrid fitnss ould at ttraploids ar highly s
synrgistially with linkd isolation gns to xtnd thir ffts ovr a largr valr'latingall hart
gnomi rgion. Thus hromosoml rarrallgmntsproaly do play an im. n a t i n g g a i n s t a l l s r v i t h
portant rol in rduing gn flow aross spis rrirs, ut not Irssarily A n ts u l vl l y A I i
through th mhanisms of rdud hybrid fitnss traditionally nvisiond. dmonstratd an unusu
Th hyrid sunflower spis H' omlus, whih rsults from hyridization spis H. ursir>kt.Us
twn H. us and I.I' petiolris, is thought to hav arisn in suh mitohondrial gn rgr
mannr. In ddition, H. omlus has highr fitnss in novl nvironmnts this study showd thirt
whr oth of its parntal spis pognitors do poorly. Suh n adaptiv r i u dt o m t r l r i p l o r i g ,
rol for hyridization in fostring spiation has bn suggstdfor a numr 8). Two of th thr dip
of plant and som animal spis (Arnold 19971Dow|tng and Secot 1997). suggstd y th ourr
not in th iiving dipl
sPlAloN Y PoLYPLolDY t t r l l l i d sa r o \ t h r ( ) u
diffrnt dipllid nsto
Polyploidization ours through th dupliation of ntir sts of hromo- rls from diffrnt dip
soms and is on of th fw mhanismsthat an produ instantanoussp- symptry. Busth dl
iation (polyploid spisar rprodutivly inompatil with thir diploid latd from on antlthr,,
prognitors). Polyploid spis ar formd ithr trough gnom duplia- u r s i tl l t , r . P l p l t ' i di z . r t
tion of a singl diploid spis(autopolyploidy)or through th fusion of two sisntharrgin th dv
or mor diploid gnoms as a rsult of intrspifi hyridization (allopoly- ll siz, whih hs n
ploidy). Both mhanisms of plyploidy ar ommon in frns' mosss,alga, ially prodd utotrip
and virtually all groups of vasula plants, partiularly angiosprms (St- hngs in trill rat s a r
ins 1950;Lvin 2002). til.ltlit twn lirrgs
How polyploid spisonrstlishdis not fully undrstood.Svral rinftlrd tri Il-reltdiffl
onditions, partiulaly ommon in plants, might nal a nw polyploid to gry tr frogs illustrat
inras and form a vial population. Ths inlud slf.frtilization' vg. ais nd furth shows
tativ propagation' highr fitnss than th diploid prognitor spis, or mnt lls, an lirt
nih sparation from th diploid prognitor spis'Indd,many polyploid
tax rprodu y slfing or vgtultivpropagation, nd most diffr from
H RoL oF G
thir diploid prognitors in haitat and distriution. lnrassin ploidy altr
ell siz, watr ontnt rat' rat of dvlopmnt, and many othr physiolog- Irr thory, gnri drift
ial proprtis' so many polyploids may hav sltivadvantagSin nw outward from th dg
ologil nihs imnrdiatlyupon thir origin. whn a small nulrlr tl]
An nusual as of polyploid spiation in animals ours in th gay rr nw popr-rizrtilr-r.
Irrpr:
frog spisomplx found throughout th astrn Unitd Stats.Th omplx fomatirn rf rrw spt

rdidrominationalspiation in
ts of hronrosoml rarrangmnts
R o m i n t i o n slp i a r i o no
giv ris to a nW linag that is
produtivly isolatd frm oth
that hromosomal rarrangmnts
jor fftson supprssing romi-
th rarrarrgdsgmnts nd gns
oms.Gns unlinkd to th ho-
rlly thosthat ar nutral, frly in.
Ltal spis. Rarrangmnts that
l fft on hyrid fitnss ould at
to xtnd thir ffts ovr largr
angmntsproaly do play an im-
spisarrirs'ut not nssariIy
rid fitnsstraditionally rrvisiond.
s, whih rsultsfrom hyridization
thought to hav arisn in suh a
i g h rf i t n s si n n o v l n v i r o n m n t s
ritors do poorly. Sh an daptiv
on has nsuggstdfor numr
1 9 9 7 ;D o w l i n g a n d S o r t 9 9 7 ) '
lliation of ntir sts of hromo.
hat an poduinstantanoussp-
ly inonrparilwith thir diploid
d ith through gnom duplia-
,loidy)o thrugh th fusion of two
rtrspifihyridization (llopoly-
trommon in frns, mosss, alga,
.ts,partiularly angisprms (Stb-
hd is not fully undrstood. Svral
s, might nbl a nw polyploid to
h s i n l u d s l f - f r t i l i z a t i o nv, g .
t h d i p l o i d p r o g n i t o r s p
to spis.Indd,many polyploid
propgation' and most diff from
istribution'Inrassin ploidy altr
)pmnt'and many othr physiolog-
'hav sltivadvantgs in nw
origin.
n in animals ours in th gray tr
) astrnUnitd Stats.Th ompl
Th Pttr nd Procss of Spition 201
ontains a dipllil spis, Hl hr51lslis(2N=24) and a ttraplid
spisl ursicolor (4N=48). Both spis oupy lag rgions of al.
u r s l o p a t r y( I J . h r l s o s e l i s i n t h s o u t h r n h a l fo f t h r a n g ,H . u e r s i o l o r i r r t h
northrn half), ut thy also ist irr Sympatry in multipl loalitis whr
thir rngs ovrlap. Diploids and ttraploids rprodu sually and ar r-
produtivlyisoIatdfrom ln anothr through oth prmating (mting sig-
nal diffrns)nd postmating (tiploid hyrids ar stril) arrirs to
intrrding.Mals produ advtisrntalls tht diffr in thir trill
rats,rvith th liploid H. cr\,slsllrvays produing illls with fastr trill
ratsthan tlrs of the ttraploid H. ursilllr(Grh:rrdt2005). Studis y
arl Grhardt nd ollagushav shown that fmals of oth diploids and
ttraploids ar highly sltiv (sially in sympatri populations) whn
valuatingall haratristisof mal advrtismntalls, strongly disrimi-
natingagainst lls with th trill rate of th ..wrong'' spis.
A rntstudy ,vAliia Hollowy nd ollgus(Hollow t al. 2006) has
dmonstrat] n unusul pattrn of p<llyplidspiation in th ttraploid
spisH. L'rsioklr. Using DNA squns frm oth mtrnally inhritd
mitohondrial gn rgions and iparntally inhritd rrular gn rgiorrs,
this study shlwd that at last thr diffrnt diploid pognitor Spison.
triutdto multipl origins of th ttraploid gray tr frog H. ursiolor (Figur
8). Two of th tlr diploid prognitl spis ar from tint linags' as
suggstdy th ()urrnof uniqrrlllsfound in ttrploid linagsut
not in th livig diploid lirrags amind. Tlrs rsults suggst tht
ttraploids 21roshrough rurrnt allopolyploid spiirtion vnts involvirrg
diffrnt diploid anstors. Intrstingly,diffrnt linags of ttraploids tht
aros fom diffrnt diploid anstors frly intrrd whr thy our rn
symptry. Baus th diffrnt ttraploid linags ar not rprodutivly iso-
latdfrom on irn<rthr, thy ar still onsidrda singl ttaploid spis'H.
rsiolrr.Polypltli1iz;rtilrr in this spis-layhav ld to pditl and on-
sistntlrirngi th dvrtisrrrrrt all, phaps s ..rdirt sult of inrsd
ll siz, whilr hs n shown tl lrlatwith drsd trill rat in artifi-
ially produd uttriploid gray tr frgs (Kllr and Grhardt 2001). Thus
hangsin trill rat as a rsult of plyploid vnts hav ld to rprodutiv ln-
tinuity twn linags rprsnting diffrnt origins of ttraploids ut av
rinfordtrill-rat diffrnstwndiploids and ttploids.Spiationin
gray tr frogs illustatsth pl'xt]annrin whih polvploid spisan
aris and ftrthr shlws how lr:rrrgsin isolating ilrrirs, suh as advrtis-
i s ,o r mnt alls, :ln dirtly afftd y hyridization and polyplidy.
HE RoL oF GN| DR|F |N sP|AIoN
In thory, gnti drift should pronlot spiation as populations pand
outwerd fror-r-r tlr dgs of thir distriutions tO otlpy nrv hitats' or
whn a srrr:rllrrmrof individuirls disprsto a nw loation and stlish
a nw populti<ln.In prati' ho'uvvr, littl mpirial support xists for th
formation of nw spis solly through random gnti hangs promotd
 t:
202 The Pttr d Process of Spition t

aus of random fiat
of anstral oadaptd 1
would lad t th potr-
sts of gns in th nr
ould inidntally yild
for rapid spiation. Al
p o | d t l o u r i n f t t
jor gn rorganization
son (1975) and Alan T
on potntial xamp
drift, howvr, oms f:
tion of shlls) twn
gnus uhdr' Tese :
t w nt h m . h i r l i
with dtral (right-han
handd) oiling. Bau
xtint (ths snails r hrma
|iyla sp. f i d f o r o p p o s i r o i l i n
(2N)N ltd from on anothr
H, avivoa
posit oiling typs ar l
gnti drift is likly t
typs m()ng pllpultr
drift is th most plaus
hirlity (vn Batrru
of at last four indpn
through phylognti
(Ushima and Asami 2

Figur 8. A modl of ttraploidoriginsfrom diploid anstorsillustratsmultipl

Consqunsof s
allopolyploid origins for th gray tr frog Hl uersiolor.Th rlationshipsof
diploid gray tr frogs nd a rltd spis,H. uiuo,ar dpitdat th
otto- of th tr. tint diploids (2N), l-/. sp. A and H. sp. B, wr infrrd Th ulk of this ssayh
from ttraploid lll linags.Ttraploid gray t frogs, H. uersiolor (4N), mhanisms y whih t
wr formd multipltims from xtintdiploid anstors(AxB, A' and sultd in th normous
B x ). Ttrploid gray tr frogs from diffrntlinags(.g.,N, NW) ar unovrd in th foss
intrfrtiland thus form a singl iologial spistht sharsa divrsgn hallnging;hnmot
pool of multiplanstrlprognitorspis.(From Holloway t l. 2006.) pls. Although mrrh d
as illustratd y th f
on spis into anotht
vrsifiation and spt
y gnti drift. Th ida of fouder-effet spitio (Figur 4) was hrst pro- ologists gnrally agr
poso o'y ns yr 1i-}.+i,t6..;retrnu'pz?rijtri "if.tittit1\.ltl "r';. ditzrgntdav amo
asd on his osrvation that in man irds and mammals, populations p- s p o n s i l f o s p i e t i
Th tmpo of sp
riphral to th distribution of a proabl..parnt'' spiswr oftn highly
groups' suh as hors
divrgnt,to th point of ing lassifidas diffrnt spis.Mar proposd
that gnti hang would vry rapid in suh loalizd founding populations for ovr 1.50 million

xtint
yla sp.A
2N)N
diploidnstorsillustratsmLrltipl
|lursiolor.Th rlationshipsof
H. uiuo,ar dpitdat th
' sp.A and H. sp. B' wr infrrd
ay trfrlgs,H. uersiolor (4N),
loid arrstors(A x B, A x , arrd
ntlings(.g.,N, NW) ar
spisthat sharsa divrsgn
. (FromHollowy t al. 2006.)
' spcition(Figur4) was first ro-
ed priptri spitio) and was
rds and mmmals, populations p-
..parnt''
spis wr oftn highly
rs diffntspis.N4ayr proposd
suhlolizdfoundir-rg populations
.l'h
Pttr r] Prlsso|.Spitiltt 20.
bausof rarr]omfixtitrrr<lfallls y grrtidrift' rsulting irr bakup
of nstraloadaptd gn ompls arrd th formtion of nw ons that
would lad to th potntial f<lrnatural sltionto favor diffrnt illtrating
stsof gns in th nw foun]rnvirotlnnts.Suh .nrorglliztions
ould inidlrt:rllyyild rprodutiv is<llationand lrn provil l1 avnu
for rapid spiation. Although gnti lrangsassoiatdwith gnti drift
praldo ottr in foundr populations, littl vidnfor th typs lf m-
jor grrrorganizationsProposd y ay (1954) and ltr Hnlpton r-
s o r r( 1 9 7 5 )a n d A l n T r r r p l t o n( 1 9 8 0 ) x i s t s .
on potntil ampl of spiation pOmotd y hangs du t gnti
dift, howvr' oms from spisdiffrnsin hirality (th liling dir-
tiorl of shlls) twn diff.ntspis of th Japrrsland snil in th
geusuhtlr.r.hs snils liv in srirllpopulatiorrswith liritldisprsal
twnthm. hirality is ltrmindy a singl grrlous with twl allls,
with dxtral (right-hndd)oiling muh mor ommon than sinistrl (ift-
harldd)triling. Bus of th physial diffiulty ill tw<l-waylpr.rlatiorr
(hssails a lrrnlaphr<ldits) twn srr:rilsfroru populatitlrrstht r
fid for opplsitoiling dirtions' ths populatiorlsar rprodutivly iso-
ltd from on anothr and so ppar tO rprsntdistint spis.Sin lp-
positoiling typs ar slt.d againstwhrr thy ar ra witin populations,
gntidrift is likly tl hv plyd a rol ilr th appran of nrv liling
t\'psamong populations. Gnti modls hav drrllnstratdtlrt gnti
drift is th most plausil thanism for swith frlm dtral to sinistral
hirality (van Batrrurgand Gittnrgr 1996) and th pralll spition
of t lstfour il-rlpndntl ]ivdsinistal spislras n dnlostrtd
through phylognti anrly5i5of mitohondrial DNA haplotyp vrietiorr
( U s h i m a n d A s a m i 2 0 0 ) .
nsqunsof Spiation
Th ulk of this ssayhas for"rsd on amplsof living spisand th veioLts
mlranismsy lvlrih th ar thought tl lrav spit.l.Yt spiati<lrr has r.
sultdin tlr normous divrsityof norv-tintanimirls,plants, irrld riros
unovred in tl-rfossil rrd as wll. Dtrmining pross frorrr pattrn is
hallnging;hn most studis tlf th spiation prtlss fous ln living m-
pls.Although n-ruhdathas ntrdOn th mod llnd tmpo t>fspition
s illustratd y th fossil rord (phlri gduirlisnl, tlr sllrv trr-rsitiorr
of
on spisinto anothr' vrsuspuntuatdqr'riliriurrr' rapid volutionarydi-
vrsifiationand spiationfollowd y long priods of stasis),volutionar,vi-
ologistsgnrallvagr that th sam volr'rtionarylhanismsthat prorot
divrgr-retod1,nrong popultitlnsof living spis(nrirovt>lutiolr) r r_
sponsilflr siationvnts(mrovtllution),otlr lngoingar-rdin th past.
Th tmpo of spiation vais dramiltially among diffrnt taa. Som
groups'suh as hrsshoras (gnus l-imulus), hav rmind unhangd
for ovr l 50 million vars, w,hil othrs' sr'ls Hr'r.iianrikts (gnus
Te Pttrn d Pross of Spitio

Lupl), hav spiatd rapidly, with up to 4.17 spis pr million yars )iz, A., and M. R. nrr'
(ndlson and Shaw 2005). Thus spiation an our quikly or slowly, p r z r . g tilt r p r o d u r i v i
undr varity of ologil onditions, and with or without th volution of p . l g r l i t o r . g u t t t u s .i
Dozhnsk,v,T, 1937. Gt
rproiutiv isolation.
LIr-rivsitPrss.
Unravling th mystris of how nw spis aris is on of th nrost xit- l 946. (io.rpltt'pr
ing ras of volutionar iology' and although w hav mad muh progrss spis lf Drostphil.
in undrstanding spiation sin William Batson's arly onrns, futur Dowlir'rg,. ., d . l,. S
studis promis to yild vn mor nw and fasinating disovris. Th hop in th divrsifi:rtiol.ttlf :
28 59_61'9.
of th futur, in prsrving and protting our plat's iodivrsity, lis in an
F d , . | .L , | 9 9 8 . T h a p p l el
r.rndrstanding of th pross of speiation and ommitmnt to saving th Onvntionalwisdom a
produts of this pross: th spis that ist on rth today and thos that ds., lld/ess Flrnts:S
hv yt to voiv in th ftrtr-rr. Urrivrsit,vPrss.
Fishr,R. A. l930. Th ,t
Prss'
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1'96.ill S1lis d t'olutil.arnridg,A: Blknp Pssof brood p rsiti indig'b
avrd Unir,rsiPrss. S p i r i t t - l. . 1 9 9 8 .T h t l l f
innon' J. s.' s. lri, B. K. Blkn.ran,L. Dvid, D. . ingsl1',L..|anris.ln, D. J. Howard nd S. H'
J' 1rou,arrd D. Shlutr' 2004' vidn ftl llt>g's rol in spii]tion. 32o-3z9.Oxford:Of.
Ntttr 429 294_298. Stins, G. L. 195. rit
ad, L. S., nd S. J. Anold. 2t)04. Quntitti\,gntirlls<lfstll sltin. Ul-rivsitPrss.
reds i rlllg1, d t'tllutitl 1'9:164_2-71. Sturtvant' A. H. 1920. Gr
.r.' Hyrids wtth Drosophl
N{ndlson, nd K. L. Shaw. 200.5.Rapil spiition in n irrthrood. Ntur
4:375-76. Tmplton, A. R. 1980. Tht
N{itr, S., . I-ndl'nd S. Prrrtt-Jorrs. 1996. Spisrihnssvlris rvith 94 1|1-1038.
mating systnl it ir.ds./,,uk 11:544_.5.'1. l989. h maning
N4ollr,A. P.' nd J..|. trrvo. 1998. Spiation nd flrthr<lrnrnnti.ttilrr irr irds. n d J . A . n d l r , d s . ,
uolutitlll 52: 859_t]69. A : S i n u rA s s o i t
Noor, M. A. F. 199.'. Spitilndrivn y nturl sltiot-t in Droslphil. Nfutr ing, .-T.' S. . Tsaur, .
3 7 5 : 6 7 4 - 6 75 . homoox at th sit O
1999. Rinformnt and Othr onsqunsf symatry. Hrdit U s h i r n a ,R . . n d . A a m i .
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Orr, . A. 199.'. Th populti<)ngntisf spition:Th vllutionof hyrid van Batnurg, F. H. D., n
impatiilities.Gctis 139: l 80.5_18 l 3' oiling: omputr pt
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1 . ] .1] _ l 3 5 . Whit, .J.D' 1978'od'
Panhuis, T. M., R. K. Butlin, NI. Zuk, nd T. Trgnza.2001' Srtalsltiorrnd ompany.
-l
spiation.rltds i oklgl d t,ltil 16 64_37 |. Wily, . o. 978. Th vo
Patrson' H. . H. 1985. h gnitionOnptof sis.ln . S. V:r' d., Zoolog 27:17-26.
.Wilkirrson,
Spis.rd Sci|tti,21-29.Transv:rIMusut onlgrph rlo. 4. Ptlria, G. S., nd P. R.
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Drwi's finlrs.Ntur 409: lti.5_]88'
Ri, W. R. 1998. Intrgnornionflit' intrlous ntgonistior,olutiot"t, and th
l.llutionof rpodr-rtiv isoltitln.In D..J' Hlward nd S. H. Brlhr,os.,
dlss Flrms: S1lisd Spitio,261_270. Nw Ytlrk: ofrd Urrivsity
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Ri'.!(. R., and . . Hlstt.1993. I,aoratoryprints n spietion:!(/l-rat
hv rv lrlrdin 40 yrs? F|lltklrt47: 167_1653.
Risrg,L. H' 2001' hrrnrtlsmaIrrngmntsand spiatiOl1.Trds i
tfulgd tllutiltz16: .51_j.58.
Rundl. . D.' and D. Shlutr. 1998. Rinflrlentof stiklirkr.nat
prfrns:Syrnptrr.rlsnrmpt. t l l tttio .'2,:200_208.
Ryan, M. J. l986. Nufoanrtomyir.rf1uns spilrtinrilts :]nlOngxnurJns.
Prldigslf th Ntiol rlm tlf SiltsL/S B3: 179_182.
Ryan, M..|., nd A' S. Rnd. 1993. SpisrOgllitiond sul s|tinas
unitary prolrn in ar-rimalmm unitio. lltttitt 47 : 647 _6 57 .
Shlut,D.' nd l,. M. Nagl. l995. I)arlllspiritlrly natural slti<ln.
lritt Nturlist 146: 292_()|.
Shlutr' D.' and T. Pri. l99.j. Holrst-y,PrtiOn,anl p1lLrlirtiorr divrgnin
sxually sltd traits, Prot:digsrlf th Rln,l Sit lf Lldo, Sris B
2 - 5 3 :1 1 7 - I Z Z .
.I'lttlll1'.
Simpson, G. G. 1961 , Pritlipls f il Nw Yrk: lLtlnbi
Univrsitv Prss.

f Spis'Nrv Yok: olumi
t and volution.IrrJ. L{ulv,
t|o|1|ls Pross,1.57-180' I,otrdn:
mbridg,A: Blknp Prss of
David, D. . Kingsi1',[,..|an-ri'son,
-.Illttgr..' .ll ilt siltitllt.
ltiv gntillodlsof sul s|tion.
4-27 1.
spiatiorrin n arthropod. Ntur
)6' SPlsrihssoviswit
l4--s.51.
ion and fthronrmntationin irds.
ttttrIsItionin Drurphil. Ntr
3qunsof sr,rnptry.Herdit
;piation:Tlr volutior.rof lryrid
il3.
gntisf slititln.
Gtis 744:
Trsnz.200l. Sual sltiorrand
tio 16:364-71.
lptof spis.In . S. Vra, d.,
l usurn Mlnograph no. 4. Prtoria,
Itrllt,gyltntJrrt1|sig11.11
5tl'lI-l
in
.|ousntagolisti
ovllutill.t,
nd th
' J. wrd .rndS. . BrlIr,ds.,
61-270. Nr,vYlrk: Oford Univrsity
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WIr:r
7:1637-1653.
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rmntof stiklakmat
tlrllutio 52: 200_208.
ipirionatsmon8 l1urns.
S i c sU S 8 : \ 7 9 _ | 8 2 .
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'n.L.uolution17: 647-657.
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ption'and oultilndivrgllin
Rl Sllit'tlf I'lttd, Sris lJ
|o||orIJ'.
Nrv York: olLtmia
h ttt d Pr.>css
;f Spitiotl 207
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valution.rin N turlist,l04: |27 -7 5 .
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rood parasiti indigobirds. Ntur 424: 928_93|.
Spirito,F. 1998. Th rol of hrmosomal rarrangmntsin spiation.In
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9 4 : 7 1l,l _ 1 0 3 8 '
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-l998.
ing, .-T., S. . Tsaur, .-L. I/u,nd .-I. Yu' A rapidly volvirrg
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oilig: omputr pimntso hirality in snils' Hrdit 76:278-286.
vn Vln, L. 1976. oiogial spis,multispis,and ks. To ) 5:2-29.
.W.
Whit, . .J.D. 1978. odcs of Spitio|1.Sn F-raniso: H. Frmn and
ompany.
!ily,. o.1978. lr vlutilnrvspis()nptorrsidrd'Sstmti
Zoollg 27: 17-26.
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sltion on an aggrrtdmal trait in stalk-d fl. Proedigs of th
Rol! 5u,i,'.r' of Lodo, Sris B 255: l_6.
 i
I
Y

volution and Dvlopmnt

GrgorA. Wr
/Ft
bitt
genorye lnt
Th intlltual histlris of dvloprntaland volutilnay iology hav
Figu 1. Gnotyp to p
n intrtlvind alr.l-rost frolrr th irt of iology as a distiIlt sintifidis. gnti information into
i p l i n ( a v r 1 9 8 2 ; R i h a r d s 1 9 9 2 ) . . h r a s t l n s r n o t d i f f i u ] tt t l 1 i n d . havior,and physiolog
Both v<llutionnd dvllpmntar tmporl prosss,and vy individ. whre thy influnsur'
ul's dvlopmtltirllristory is mddd within, an1rflts,a muh dp volutionarymhanism
voluti<tnaryhistory. This tmporal paralll forrns th sis flr th orrpt sussivgnration.A
<lfrapitultitln' tht n-rryonidvlopmnt litrally rplays volutitlnar1' how prossesopt1
link in our knowldg w
hisroy. Rirpitultitlnis on of th ,ldstand most prsistnrl1,ontrovI.
sil onpts in vtllr'rtionzrry dvlopmntaliology' ut not its mst illumi-
n a t i n g ( G < r r . r1l d9 7 7 1 P . a f f1 9 9 6 ;W i l k i n s 2 0 0 2 ) .
Th rol f dvlopmntin tratlstrrutinggllotyp irrto phnotyp provids
a mor fruitful pproah to undrstanding th intrations btwn volu. ntury (Hall 1992; Ri
titlnary and dvloprntal prosss (Rff d Kaufirran 198; Wilkins prossswr onsid
2002). Th volution of th gn()llland tl-rvoluti<rrr of ogirrlismalplrn. arly desriptions o{
typ, whih hav oftn n trtd s ifthy Wr sparatprsss,ar in. ast in trms of th s
xtrily tlund togthr y th prosss of dvlopnrnt (Figur 1). Th emryos of mor omp
\\,lly l1 mryo funtitlns has s ruh ttl do with ollgiallIrttnd smld simplr ons
ph,vlogntihistory as it dos with gnraring prtiular plrnotvpi out. tnth-ntury prfor
m. lnvrsly,tlr volutionary fat of a goup lf rlatd spisan t h s m p o s s .D u r i
influndlrs mulr y dvloprrrlltlrl hratristisas it is 1.ldult anatlmr.' tors notd that hikn
p h s i o | g vn. n l h . r v i o . that ontain faturs of
for instan, tht hum
w i t h l i m s p p a r i n gi
Tmporal Parallls still.
Karl rnst von Bar'
Rapitultion has by ttll1srrlightnd,ntrtailrd,lrnd spratdgrrr. vations of divrs nir
t i o n s l f i o l o g i s t s( d B r 1 9 4 0 ; G < l u l d| 9 7 7 ; u r h i l l 1 9 8 0 ) .T h r o g . tion (Hall 1'992i Rical
nition of parallls twn mrologil vt-ltslld rlationships among th mryos of vrtr
spis prdatd th onpt tlf lriologial vtllutitln, at [ast in its modrn rpsnt adults of oth
strsof trnsmtltation irtwnspis.Indd,th rvords uriutil and d. lowr spis do not a
ulopttwr usd intrhangalyto rfr tl th unfolding of oth indi. sn of fins in th m
vidual dvlopn-rnt arrd plrltillang until t Lrginningrlf th t.ntith ryos of distint group

208

)vlopmnt
ntal and voiutitlnary iollgy hav
lf iology as a distint sintifi dis-
h rasonsr not diffiult to find.
nporal prosss,and vry individ-
within, and rflts'a muh dpr
rlll forms th asis for th onpt
,pmntlitrally rplys volutionary
st nd most prsistntlyontrovr-
rtal iol<lgy,ut nlt its most illumi-
20021.
lg gllotyp into phnotyp provids
ing th intrationstwn volu-
Ratf and Kaufian 1983; Wilkins
th volutior-r of orgnismal phno-
t h yw r s P a r a t r o s s s. r i n -
; s so f d v l l p m n(tF i g u r 1 ) . T h
to do with oiogial ontt and
r a t i n g p t i t r l l rp h n l t y p i o u -
of a group o{ rlatd spisan
Lratristisas it is y adult anatlmy,
ntrtaind,rrdxaspratdgrrr-
J 1 9 7 7 1 h u r h i l l1 9 8 0 ) .T h r o g -
:al vnts nd rltionships among
;al volution,at last in its modrn
Indd,th words uolutio ad d-
l rfr to th unfolding of oth indi-
until th glnningof th twntith
uolutio d DeueloDmt 209
ev0li"rtr0nary
mehisms
g
t
i
bihemial deve|mntal rgantsmaI
genOtype
interat|Ons rOess8s nenolyp
Figur 1. Gnotyp to phnotyp.Th prosssof dvlopmnttransmut
gentiinformationinto organismalphnotyp,whih inludsanatomy'
havior,and physiolgy. Ths traits, in turn' intrat with th nvironment,
whrthy influnsurvivrship and rprodution. Naturl sltionand othr
evolutionarymhnismshang th rlativ frquenisof gnotypsin ah
sussivgnration.A omplt undrstandingof volution rquirs knowing
how prosssopratat ah lf thsstps.Until quit rntly,th wakst
link in our knowldg was dvlopmnt.
ntury (|1a|| 1992 Rihards 1992)..shat w rogniz today as distint
prossswr onsidrd intrialy intrtwind.
arly dsriptions of parallls twn dvlopmnt and volution wr
ast in trms of the scl ntur (hain of ing). Aristotl rognizd that
mryosof mor ompl raturspassd through stagsin whih thy r-
smld simplr ns (Gould 1977|. For harls Bonnt, an arly-igh-
tnth-nturyprformationist, volution and dvlopmnt Wr litrally
th sam pross. During th lat 1700s and arly 1800s svral invstiga-
tors notd that hikn and human mryos pass through a sris of stags
..lowr'' or ..infrior'' animals.
tht ontain faturs of Johann Mkl, notd,
for instan'that human mryos ontain gill slits and a tail at arly stags'
with lims apparing aftrwards' and dfinitiv mammalian fturs latr
still.
Karl rnst von Bar' who mad tnsiv and mtiulous firsthand osr.
vations of divrs animal mryos, pliitly rjtd th ida of rapitula-
tion (Hall 19921Riards \992). H itd th prsn of uniqu faturs in
th mryos of vrtrats,suh as th yolk sa) as vidn that thy did not
rprsntdults of othr spis;onvrsly,h notd that som faturs of
lowr spisdo not appar in th mryos of higher ons, suh as th ab-
snof fins in th mryos of irds (Bar 1 82 8 ). Bar also notd that th m-
brysof distint groups of animals did not pass through stagsrprsnting
2 1 0 ulutil d D uelopm t

othr Ons:..T mryil tlf vrtratis at t girrrringalla vrt-

rat'' (translationfrom Rihards |992, 59).
Dspit Ba's potnt ritiisnrs,support for th nrtin of rapitulation
prsistd for anothr ntury. Rapitr.rlationthory rlrd its znith in th
..ontognyr-
rvitingsof rrstHakl (1866, l875), whos satmntthat
pitrrlatsphyllgny'' (..di ontogni witr nihts ist als in kurz Rapitu-
lation dr Phylogni'';1866,2,7) r,ntldayrrapsulats flr rany iologists
th rltionships twn v<llutinnd dvllpmnt.Hakl, str>lrg sup-
portr of Darwin, viwd dvlopnrnt as an hiit of phylgnti history'
with th adults of anstorsdisplayd in squnduring th dvlopmntof an
individual. His rvoduts lpiting th progrssiv divrgrrlf initially simi-
l;rr vertrat mryos into distintiv fish, turtl, ird, and humn nrryos
hav om ions of lat-nintnth-nturyvolutionary dvlopmntal iol
ogy (Figur 2). Lss widly appriatdis Hakl's viw of rapitulirtionas
th irtual mhilnism that prodtrs v<llutionrylrang rtlrr thn sirnpla
pirssivrord of suh hirng.H onsidrd tis mhanism to onsist of two
prosss:trminal addition of new faturs onto th nd of dvlopl-rrnt and
ondnstion of xisting fatursinto vr mor .lbrvit]arlir st;lgs.
Hakl ognizd tlr istn of 1rtionsto strit rpitultion
( 1 8 7 5 ) ' i n l u d i n g t h o j t i o n sr a i s d y B a r ( 1 8 2 8 ) . H l l d t h m
Figur 2. Rapitultitln ;
rlgetic fturs and rgardd thm in mr.rhth sar light that most
natomist rnst Hrrkl '
phylogntiistsviw h<lrr-rolasy (faturstht hav lnvrgdin fttrrrrut
th volutitlnar history
without ommon anstry)today, as an ann()yingofusation of v()lution- dvlopl-nnt (Hakl 1 8
ary history. }Iakl idntifid two kinds of nlgnti faturs: hte- on of whih is sho,n h
rochro' or a hang in th ordr of dvlopmntvntsJtld btrltop,or among sis as dvllp
a hang in th grm lirr giving ris to strutur.Bot trms ar now usd thos of fish, r-rrphiia
in rnuh mor gnrl snss,to rfr to any l.rangin th timing or loation ll th mryos hav gill s
mryo hs til (middl
of a dvloprntal pross, rsptiviy (Raff 1996; Klingnrg 1998).
ws th prsl1f tlrs
Ironially, and not a litl onfusingl. th urrnt, mor gnral usgof th
t h i r s n i n l u l t s ,t h
tr1trochron encotpasssstitly rapitulatory tmporal hngs,as
wll as Hakl's original, nonrpitulatoryons.
Br was not th onl,v prson to attak rapitulatin thoryJ nor Was
Hakl its only apologist. Rrrrarkly,dirtsaorrt rapitulrtionwr
still going str()ngduring th 1930s as th modrn synth.siswas undr waY; among taxa, and of th
they may hav ontriutd to th ordial mutr.rlpathy that aros twn appliations wr all fir
voltltionay arrd dvllpnrntalbiology atlut this tim nd lastd trrrtilth somwhar diffrnt rirs
l t 1 9 8 0 s ( G o u l d 1 9 7 7 ; W i l k i n s 2 0 0 2 ) . A l t h o u g h t h i n f l u n o f r a p i t u - Not rany radsof l
latitln vntully fadd, omparativ mryology ontinud to pl n im- of volutiln taks pla
pOrtant rol in volutionarv ilogy. marshald all th vidn
tation 1ot1]spiswh
Bar, wlro rjtd th nl
mryos as vidn of volution, Phylogny' dvloprnntaltraits' D
and Homology plling vidn of on
strongst singl lilss of
If mryos do not infallily rapitrrlatvolutionary history, thy ntlnth- tr to As Gray (Sptm
lss provid vidnof th prssof volution.f phylogntirlationships giz (Burklr:rrdtt al. t99
 uoltio ltd DuloDlnt 211

is at th girrningalra1ya vrt-
se).
rt fr th nltitln of rapitulation
tiorr thory rahd its zrrith in tlr
,hosstatmntthat ..ontognyra-
tr nihts ist als in kurz Rpitu-
'dayrrapsulats fo rany iologists
Jvlopmnt.kl, strong su-
s an hiit of phylogntihistory,
qunduring tlr dvlopnrntof an
ogrssir. dir,grr of initilly simi_
lh, turtl,ird. and humn mryos
ury volutionardvlopmntal iol.
; H:rkl'svi'uvof rapitrtlation as
ltionaryhrrgrathr thn simply
rd this mharrismto onsist of two
sonto th n1of dvlopnrnt and
.mor rvitdarlir stags.
ptions to strit rapitulation
y Bar (1828). H alld thm
Figur2. Rapitrrltitln as nvisind1.Hakl. Th gratomparativ
in muh th sm light tat most
anatlt-t-tistrnst kl was tln of t strongstProPonntsof t notion that
s tht hav onvrgd in fom Ltt
thvoIutionr,v historyof 'r spisis rplavddrringits -rrylogial
nr-roying ofr.rstion of volution- -l
dvlomnt(Hakl tl66).His famos illustratinsof vrtratmryls,
nds of nognti features.. ht- onof whih is shown hr,lpitprgssivdivrgnin ovralisimilaritis
lopr-nntvnts' and htrltop' or amngspisas lvlopnllrtprods.Th mryos ar (from lft tl right)
t strutur.Both trms ilr now usd thosf a 6slr,arrrphiian. tr-rrtl, ird, pig, ow, rzrllit,and hurt-t. Not that
any hangin th timing o loation all tlr mryos lrvgill slits t an rl Stag(top ru) and tht th umn
Ly (Raff 1996; Klingnrg 199s). m r v o h s t i |( m i d d l r o t u , r i g h tt)l - r a t i s l t r \ o s t ( b o t t o m r l , r i g h t ) . I t
Wasth prslltlf thsplt>gntiaIlv nintftursin rrrlrryos, dspit
Llrnt'mor gnrl us:rgof th
tlrirsnirl dults,tha strgl}.i.rrssd Hkl.(Frlmkl 1866.)
rpitultr. tmporal hangs, as
ory ons'
ak rapitulation thoryt nor Ws
, dtsaor-rtapitulirtiorrlvr
modrn synthsiswas undr way; amollg ta' al1d of th lristory of spifi harirtrs.hs thr prtial
l mr'ltualapathv that aros twn appliationswr all first proposd during th nintnth ntury, ut for
aout this tim irnd lastd trntil th somrvhatdiffrrrtrasons thy rmairrimportant tody.
Although tlr influnof rapitr.r. N()t mn radrs of tlris olk lvilI nd to prsuadd that th pross
rrvologyontinud to play an im- of volution taks pla. Antiipatirrg a rathr diffrnt audin, Drwin
mrshaldall th vidn h ould fid to support th onpt of trnsmu-
tationi]mong,spiswhn lr wrot oll the rigi of Spis(1859). \Whr
.,spifi''
Bar,who rjtdth noti<lnof volr"rtion, spok of ..gnrl''and
ln, Phylogny, dvlopmntaltrits, Drwin intprtdsimilaritisamong mryos as om-
plling vidn of ommon dsnt' ..mrology is to m y far th
strongstsingl Iss of fats irr fvor of lrang of forms,'' lr l,vrot in a lt-
vlutionarylristory, thy nonth- tr to Asil Gra (Sptmr10, 1860) shortl afterth publirion of tl-rori-
llution,of phylgntiltionships gl (Burkhardtt al. 199, 350). mryologial similaritisrmin prsuasiv:
 rt
21 2 uolutiolt nd Dculrlmttt I
i

traits not manifst in adult anatomy' suh s th prsnof gill slits and a and sns or$ans prsr
tail in hurrratlmr1'<ls, ar tangil vidni dsntw,ithmodifitin. anstor or aros indr
A solld pratial appliation of omparativ mryology is irl phylogr- simila mryologial
ti analysis.Darwin pointd out in th rigi tht ..ommunity in mryoni sin orrspondn
strutur rr,lstltnmtrnitvof dsrrt''(18.i9,449). Sysrmtisrshav n 1994). nrrologial
applying this onpt nthtrsiastiallyvr sin. Sval signifiant insights mologis' suh as th
into phylognti rltionslrips am aout through us of this approah' origin of vrtrtja'
suh as th dis<lvr},thatrnals r arthopods tlrr thrr rnllr"rsks as vind y th rr
(Darwin 1854) and th ralization tat asidians ar losly rltd to vrt- (Burk and Fduia
brats (Grstang 1928). In oth ass'adults r highly lrivdanatomiallY, an our (d Br 1:
ut larva provid lr phlognti links. Sirrrilaitisirr mrylsor lrr. rains on of th nros,
providd th first vidnfor phylogntirlationshipsamong mny Othr. In rnt yas th
lvis anatomilly distint taa and oftn rm:rindth only dirt vidn to th prssion of
r-rrrtilth :ldvnt of rr-rolulr data sts.Th nrrlsf mrry lads r sd sd <ln th osrv
on ths mryologial ritria, sul] as Spirlia (for th spirl gomtry of during th dvlopm
mryoni ll divisions found in som anirals)and rnonoots (for th singl highly divrgnt anat
.s7ith
otyldon tlrat norishs th mryo of som flowring plants). tlr olland 1999). For
widsprad ppliation of DNA squnsir-rphylogntis,lrrryologial xprss not onl th
traits play a lss important rol in idntifying lds today, ut thy an po- funtion (myosin, at
vid an iortant rlity hk whrr rrw rlatilnshipsar proptlsd. trol th transriptiln
h squnof mrytllt>gial vnts has lso n invokd s asis for srvation that dvl
polarizing haratr-stattransftlrmtion in fornral phylognti analyss pattrns forms th z
(Nlson \978). his ltrlg|ttiL. rit('ri(,tllis zrsdon ir litrl :rdingof I994)' lgs (Pnganib
Br's laws: that rlir vnts duing dvloprrrnti1r mor gnral ;rnd ar homologous in ar
thrforaros arlir during phylogntihistory than latr ons. Th utility viningly rquirs o
o i t h i s i t i o n h s n q u s t i o n do n f i r s t p r i n i p l s( d Q u i r t l z 1 9 8 4 ) , rs of th sam gn
n d i t l a r l y f a i l s i n r n a n y r a l - w o r l d a s s( M a 1 9 8 9 ; R a f f l 9 9 6 ) . o u t - suh as spurious simi
group omparistln is far mor widly usd today, nd th ontogntirit- son 2001). Rigorous
riln is intrstingpinrarily as :rn ampl of th psistnf raglirr-rlationist homollgous strlltur
thinking a nturY aftr Hkl. among dutrosom
Th third pratial appliation of omparativ mrytllogyis s ritrion and of ody sgmnt
lf homollgy alon ] spifihartrs.of th tlrrpplitions rrrntiond tl 1'997: Azhnov
lrr,this rmains y th far th most impotant tod (Hll 1994; Wilkins
2002). Rihard own originally proposd th trms homolog and log
in 184 with rfnto his lloti()not th arl-rtyp' gnrlizlldy or- FItohrony
ganizatill-lshard among spis rvithin a larg gottp (Palrhn 1994).
Own, who was ostnsily strongly opposd to th idir of voltltion, intro- N{any vnts during
dud th trms t() ontrst twtl two kirrds tlf rrtomil sinrilitv partiular pa a
among spis:homology for orrsponlingprts of atl arhtypand nlrl- vnts; vn dvlop
ogy for nonhomologous traits with sirilrfuntions (Own 1848). Btlth trms in a onsistnt squ
wr soon propri:rtdy r,olutitlnarl'illogists(PnhrrI994) ltndar still vn largly indpn
trsdto distinguistwtwo funlrmntallv diffrntlristorialhypothss: most ilny hang in
ommon vrsusindpndntvtllutionaryrigins of a trait (Hall 1994). of othr aspts (Ra
Distingishing twn homllogr.rsand rralogous struturs is <lftn hang in th tinring
straightforward,ut in mll1,ssit is dinglydifliLrlt.A psistnthal- ld to th viw that
lng' for instan,is distinguishing,hthrth appndags,odv sgmnts' htrohroni (Goul

l s th prsnof gill slits and
t tlf dsntwith rodifiation.
rItivmryologyis in pylogn-
igi tat..ommunity in mryoni
1859' 449). Systmatistsl.rav
r sitr.Svral signifiant insights
lut through us of tl-risapproah,
arthropods rathr than mollusks
idiansr losly rltd to vrt_
Lltsr highl drivd anatomially,
s. Similaritisin mryos o lrva
rltionslripsmong many othr_
rmaindth onlv dirt vidn
h namsof many lads ar asd
ipiaiia (for th spirl g<lmtylf
imals)and monoOts(for th singl
som flowring plants). With th
s in phylogntis,mryologial
,ingldsttldy, ut th alr pro-
l rlationshipsar proposd.
las also rrinvrlklas a asis ftlr
t in flrnll phr,logrrtianalyss
lr is asd tlrr a lital rading of
ivloplrrntar mor gnral and
history thrrlatr tlns.lr utility
first prinipls (1Quiroz \984),
; s( M a l 9 8 9 ; R a f f 1 9 9 6 ) . o u t -
. d t d . r r . r l d t I t o n t ( ) g ( ' n t ir i
lf tlrpsistn of rapitulationist
' r t i v m l l r v o l . l g iys s . t r i r r i r l n
)f tlr thr rppliationsn-rntiond
lportanttoday (Hll l994; \/ilkins
d th trms homolog and Iog
aht1,p, a grrralizdod1,lr-
n larg group (Panen 1994).
l s dt o t h i J . r< l f v t l l t l t i o n ,i n t r o -
,o kinds tlf antomil sirnilarity
ing prts of i]n arhtvp and anal-
funtions(Own 1848). Btlth trrs
ologists(Panlrn1994) and ar still
rtallydiffrnthistorial hypotlrss.
originsof a trait (Hall 1994).
lnd arrllglusstruturs is lttn
:dinglydiffiult. A prsistnt hal-
-rrth ilppndags,ody
sgmnts,
uoltitl td D uellpmet 2 1
and snsorgans prsnt in diffrnt animal phyla ar drivd 1rom ommon
anstoror aros indpndntly (lor and lr)ilmr1997; Nilso22\. t
similar mryologial origin has long n onsidrd ritrion of lromology'
sin orrspondns in antomial rganization ar oftn larr (all
1994).mryologial omprisons hav idrrtifidsom initially surprising ho-
mologis,suh as th oigin of nrmrrralianar ossils from jawbons nd th
origin of vrtratjaws from gill arhs.This approah ontinus to usful'
as vind by th rnt dat aout digit homologis among arhosaurs
(Burkand Fdui 1997;Wagner arrd Gauthier 1999). Although xptions
an our (d Br 1971; Roth 1984), similarity of mbyologial origin r.
mains on of th rnost usful ritria of homology in anatomy'
In rntyars th mryologial rition of lrorologyhas n tndd
to th prssion of dvlopmntl rgulatory gns. This appliaion is
basdon th osrvtion that th sam rgulatory protin is oftn prsnt
duringth dvlopmntof horrrologousstrutursvn whn thy give ris to
highly divrgnt anatomis and vn in distntly rl:rtdtaa (Holland and
Hollnd \999). For instan, musl lls throughout th animl kingdom
xprss rrot only th strutural protins that ndow thm with tlrir spifi
funtion (myosin,tin' nd r-tinin)ut also rgultory protins that on-
trol th transription of ths strutural gns (ylD and MF.2). Th o-
srvation that dvlopmntal rgulat<lrygns ar prssd in similar
pattnsforms th asis of ontrovrsial prposals that ys (Quiring t al.
1994),lgs (Panganianet a|. 1997\, and sgmntation(Holla nd et a|. 1997)
ar homologous in arthropods and hordats.Framing suh argumnts on-
viningly rquirs nrpring orthologus gns (i.., orrsponding mm-
brs of th sm gn family) and ritially assssingaltrnativ hypothss,
suh as spurious similarity in gn prssion (Aouhif t,a|' 1997; David-
- son 2001). Rigorous appliations of gn-prssionritri:r to idntifyirrg
homologous struturs inlud ompaisons of th ntal nrvolls systm
among dutrostomgroups (Williams and Holland 1998; Low t al. 200.3)
and of ody sgmntsamong divrgntgroups of artlrropods (Avrof nd Pa-
te|1997i Azhanov et a|. 1999\'
Htrohrony
any vnts during dvllpnrnt happn in partiular squn and at a
partiular p aus th a triggrd by, or dpndnt upon, arlir
vnts;vn dvlopmntalvnts that ar not ausally rlatd oftn unfold
in a onsistnt squn aus of gloal oordination or simply aus
vnlargly indprrdntprosss r h tighrly orrtrolid. As a rsult, al_
most any hang in on asptof dvlopmentwill altr th rat or squn
of othr aspts (Raff 1996). Th rsult is a bterlhro'an volutionay
hangin tlr tinring of dvlopmntalprosss.This simpl onntion hs
ld to th viw tht all volutionary hang in dvlopmntis fundamntally
htrohroni(Gould 1977; Kinny and Namara 1991|'
21 1 d Dulopmt
F'ullutil

Mldrn idas aout volutionary hangs in th timing of dvlopmnt prsptiv that was soli
grW Out lf rapitr.rlationthory. Th aumulating wight of ptions and N4Namara 1991')'
stadily rodd th gnrality of akl's trminal addition and ondnsa- H t o h r o n yi n r l y
tion as volutionary mhanisms.Howvr, thr wr two additional fators fiation of dvlopmnt
that brought aout th dowrrfall of rapitulationthory: a lar undrstand- 1980s and 1990s. Dv
ing of th mhanismsthat undrli hrdity and th rognition that natural ryos and in omplx l
sltionn at troughout th lif yl (Gold 1977;kaff 1996; \Wilkins rohrony in th fossil rr
2002). By th 190s it was widly aptdthat th iogntilaw is nithr 1991)' Dvlopmntal
a mhanism of volutionary hang nor an aurat dsription of it. vn to suh pfosss as mor
th most voifrous ritis of rapitulatitlnthory rognizd,howvr,that p r i s o n s o f g n x p r s s
par:.rlllsfrquntly do ist twnvolutionry history and mryologial p o r l h a n g s .s o m i n
vnts. out of ths dats am nw and ultimatly mlr rwarding pr- Dikinson 1988; avn
s p tVl s. formations (.g., Wray l
On important insight Was that hangs in th timing of dvlopmntal lar that htrohrony
.!7l-
vnts lftn rprsnt adaptati()nsrathr than y-prldutsof hang. ould aris through a va
tr Garstng, a pugnaious and witty riti of rapitulation, assrtdthat A third important rns
..ontogny d<lsnlt rapitr-rlat phyitlgny, it rats it'' (Garstng 1922, rathr than pross (Raf
82). Garstang natly invrtd th aklian viw lf ontogny as a rord tims onsidrd to al
of phylti history ar-rdargud instad that ntural sltion ats on dvl- hang (Gould 1'9773
opmntal timing, thry produing hangsin anatomy. intrprtd original onption. To t
m a n y s g l i f i v o l u t i o n a r y h a n g si n a r l y d v l o p m n ta s a d a p t a t i o n s ,i n - traits, howvr, htro
l u d i n g i n t r p o l a t i o n s a n d l o s s s t l f l a r v a l s t a g s , h n g s i n m t h o d s o f annot aus hangs l
mtmorphosis, and altrations in th timing of rprodutin (Garstang nompasss a rih arr:
1 9 2 2 , 1 9 2 9 ) . S t p h nJ a y G o u l d t o o k u p t h i s t h m i n o t r l g n d h . mbros d Astors
loge 0977), rvhih tid htrohrony to th lassil ologial frm. sifiation of mods of h
work of r-sltion and K-sltion (r-sltionprdus adaptatilns'suh 1977; |eret a|.197
:rs rpid growth and arly rprodution' for suss in unprdital nvi- th varity of tmporal
ronmnts; K-sltionprodus th onvrsadaptationsin mor stal vidd a pris voaula
nivtlnmnts). any authlrs hav mphasizd th links twn ht- Htrohrony has fad
rohrony nd suh lif.history traits as fundity, timing of rprodution, opmntal volution, in
n d a d u l t o d y s i z ( . g . , K i r r n y a n d N m a r 1 9 9 1 ; R y a n a n d S m - pervasiv that doumnt
l i t s h 1 9 9 8 ) . T h i n p o r t a n tl s s o nt h a t m r g df r o m t h i s W O r k i s t h a t s i m - I/ilkins 2002). Th mp
pl hangs in dvllpmntal timing an prldu hirngs in organismal for volutionary hang
p h n o t y p t h a t a r o l o g i a l l yr l v a n t . hangs aris, nd how t
A sond, rlatd insight Was that volutionary hangs in dvlopmntal of phnotyp.
timing an ()urthroughout th lif yl. As arly as 189.5Frank Lilli sug-
gstd that adptatitlttsn aris in th arlist vnts of mryognsis.
(iarstang (1922) argud that nw traits ould appar at ny point in dvl- Dvlopmntal or
opmnt (not just th trminal addition advlatdy Hakl), and that su-
squnt modifiations ould shift thir tin-ringithr arlir r latr (not just A ntral gal of volutr
Hkl's ondnsation, involving progrssivshifts to an arlir position in ryp produ hngs in
ontogny).Dvlopmntal iologistsw losing intrstin volutionary pr- tntion on hrrri
sptivst th tim' howvr, and th fw volutionary iologists whtl wr idntifying whih ompl
studying htrorlnyfousd on lat lif-historystagsrathr than mbryos stalishing th numr
and larva. Most studis of htrohrony thrflr mphasizd hangs ifiations in traits. Ths
in dvlopmntal tiring rlativ to adult nat()myor to sxual maturity, a rn to th dvlopm
 uolutio d Dulomet 215

ngSin th timing of dvloprrrnt prsptivthat was solidifid y influntial ooks (Gould 1977;cKinne
rumrrltingrvight tlf ptions nd Namara 1991).
's trminal addition and ondnsa- trohrony in arly dvlopmnt ttratd rnwd intst as th runi-
'r, thrWrtwo additional fators fiation of dvlopmntal and volutionary iology gathrd pa during th
trrltilnthory:a lirr undrstand- 1980s and 1990s' Dvlopmntal iologists amind htrohrony in m-
lty and th rognition that natural bryos and in ompl lif yls, whil palorrtologistsdoumntd ht-
(Gould1977;Raff 1996;Wilkins rohrony in th fossil rord (Raff and Wray 1989; Kinny and Namara
d tht th iogntilaw is nithr 1991).Dvlopmntal biologists garrto apply th onpt of htrohrony
an aLlrtdsription of it. vrr to srrhprossss mrrphognsis,ll linag,rrd gn prssion. om-
)n thoryrognizd,hwvr' that parisonsof gn prssionamong losly rlated spisrvaldmany tm-
utionar1'history nd mryologial poral hangs' som in th asn of ovious anatomil orrlats (.g.,
nd Llltinratlymrrrwardirglr_ Dikinson 1988; avner 1992) and otlrrsorrlatdwith phnotypi trans-
formtions (.g.,Wray and Bly 19941 tverot and Patl |997).It am
gsirr th timing of dvlopmntal iar that htrohrony, lik othr kinds of phnotypi transformations'
.Wal-
r thn v-;lrodutsof hang. ould aisthrough a varity of volutionary mhanisms.
i t i o t r a p i t u l a t i o n a, s s r t dt h a t A third important insight was shift to viwing hrrohrony s pattrn
n y , i t r a t si t , ' ( G a r s t a n g 1 9 2 2 , rathr than pross (kaff 1996). Until quit rntly htrohrony Was som-
lian virv of ontogny as a ord timsonsidrdto an volutionary mhanism that produs phnotypi
l1tnilturl sltioll ats on dvl- hang(Gould 1977; Namra 19B8), a urious prsistnof akl
angs irr anatom\.. intrprtd original onption. To th tnt that dvlopmntal prosss ar hrital
r l y d v I o p m nats a d a p r a i < l n isn' - traits, howvr, htrohrony is an aspt of phnotype lik any othr and
'val stags,hangs in mthot]s of annot aus hangs in phnotyp (Raff 1996). Htrohrony nonthlss
timing <lfrprodution (Garstang nompasssa rih arra of phnotypi transformations. Gavin d Br's
p this thm tn togen d Ph- mbros d ncestors (1940\ was th first attmpt at a lmprhnsiv las-
r to th lassial ologial fram- sifitionof mods of htrohrony, arrd svral othrs hav followd (Gould
.Wray
:ltiorrprtldusdaptations, suh 7977;|rch t al. 79791Raff and 1,989I.hs analyssmphsizd
, for sussin unprdital nvi- th varity of tmporal hngs that ar possil in dvlopmnt and pro-
nvs adaptatins in mor stabl vidda prisvoabulary for dsriingtheslrangs.
r p h s i z dt h l i n k s l l t w l r l r t - Htrohrony has fadd somwht as a ntral thm in studis of dvl_
fundity,timing of rprodution, opmntalvolution, in part austmporal lrngsin dvlpmnt ar so
l MNamar 1991;Ryan and Sm- prvasivthat doumnting anothr as is no longr surprising (Raff 1996
mrgdfrom this rvtlrk is that sim- Wilkins 2002i' Th mphasis has shiftd to undrstandingth gnti asis
an pro{uharrgsin organismal for volutionaryhangs in phnotyp, th ologial ontt in wlrih suh
hangsaris,and how th organization of dvlopmntiassth volution
lutionar1.hangsir-rdvllprnntal of phnotyp.
. A s l ya s l 8 9 . 5 F a n k L i l l i s u g -
arlist vnts ()f mrygnsis.
ould appar at ay point irr dvl. Dvlopmntalonstaint and Dissoiaility
dvoat] ,vHakl)' and tht sub-
ming ithr arlir or latr (not just A ntralgoal of volutionary iology is undrstndinghow hangs in gno-
s s i v s h i f t s o a r l r l i r p t l s i t i o n i n typprodu hangs in phnotyp. volutionary gntiists hav fousd at-
. l o s i n gi l l t r s itn r ' t l l u t i n a yp r - t n t i o n o n h a r t r i z i n gg n t i v a r i t i o n w i t h p n o t y p i i m p a t ,
:w volutionaryillogistswh<lwr idntifyingwhih omponnts of this vaiation sltionoprats upon, and
-historystagsrathr than mryos stalishingth numr and rlativ impat of muttions that undrli mod-
ony thrfor mphsizd hangs ifiationsin traits' Ths qustions hav n addrssdlargly without rf-
lt anrtlnr1. lr to sul maturity' a rnto th dvlopmntal rhnisrsthat link gnotyp to phnotyp
216 uolutio d Dulopmet
evolutionary redL
sin th shism twnvolutionary and dvlopmntaliology during th
1930s and 1940s (ayr |9821Wilkins 2002), volutionary and population
gntiistshav gnrallytratd dvlopnrntas a lak ox whos ontents
hav littl to ontriut toward undrstndingvolutionary prosss.
Th validity of this approalr has n hallngd rpatdly. harls
.Whitman, Frogs
an arly volutionary 8ntiist'wondrd whthr ..th laws of
dvlopmnt lrrdsom lins of [phnotypi]variation nd favor othrs''
( 1 9 | 9 , 1 1 ) . B y t h 1 9 8 0 s s v r a l v o l u t i o n a r y i o l o g i s t sW r p r p a r dt o n -
swr in th ffirmativ (ayr 1982; Holdr 1983; aynard Srith t al.
1985; Athur 1988; Alerh 1989I. Th notion that th orgnization of dvl-
opmnt imposs iasson th prrrdutionof phnotyps smsrasonal-
to liv othrwis is to assum that vry possll phnotypi outom is
qually likly, whih is manifstly not th s (Alrh 1989; \/ilkins2002). Salamanders
Th tru hallnghas n idntifying th undrlying soursand lng-trm
onsqunsof suh dvlopmntalonstaints.
vidn of dvlopmntalonstraint has om from two sours.on is
forbidde morphologies, phnotyps that on an imagin and tht sm
funtionally rasonal ut ar nvr osrvd.Th sond is th rstritd
rang of phnotypsthat mrgfollowing natural or primntalprtura-
tion of devlopmnt. In oth ass som of th st ampls om from
t t r a p o dd i g i t p a t t r n s( F i g u r 3 ) ( H o l d r 1 9 8 ; A l r h a n d G a l 1 9 8 5 ) . B experimental redL
Although thes approahs onviningly dmonstratthat th prodution
of phnotypsan iasd,th aussof thsiasshv rmaind lusiv.
A phnotyp ould ..missing'' baus it is diffiult to produ dvlop- mitotiinhib
mntally, or ausof a physial or gomtri limitation, a lak of gnti
variation, or an unrognizdfitnssost. Distinguishing among thspossi-
ilitis has provn quit diffiult. vn whn it is possil to infr th xis-
tn of dvlopmntal onstrints y poss of limination' idntifying
what aspts of dvlopmnt ias phnotypi vition reiss furthr hal'
lngs,and fw lar ampls ist.
A rltd qustion is why suh iassist. on planation is that dvl-
opmnt is ativly ..uffrd''to produ onsistntphnotypi outomsin
spit of gntior nvironmntaIprturations.This ida was arfully arti- developinglim
ulatd y onrad \/addington(7957), who nmd it liztitl.Dvlop-
mntal biologists hav long rognizd a similr phnomnon thy ll
regultio:th aility of an mry()to onvrg on a normal phnotypfol-
Figur 3. Bisd produtio
lowing primntal manipulation. \7addingtonblivd nliztion to b
may rstitth rang lf pl
th produt of gnti assimilation' a pross whry a phnotyp that was fro-rprimntalprtur
initially invokd y th nvironmnt is susquntlygntiallyfid through r a n g o i p h n o t y p sr h i s
sltilnto maintain that phnotyp. Ivan Shmalhausn (1949) proposd i n d p n d r lt a s st l f d i g i t r
that stailizing sltion produd muh th sam rsult. othr possil amphiiirns.AIthough four
sours of dvlopmntalonstraint ar a progrssivloss of fliility s th thssult from th loss ol
prosssof dvlopmnt om urdnd y historial lgais (.Wimsatt tratingth dvlopingliml
A l ha n d G i r l ( 1 9 8 . 5w )
1986) and th liklihood that modifitions in arly dvlopmntwill hva
frogs and salmandrs.ln
roadr phnotypi impat thn latr modifiations (Arthur 1988; Gould
random ut instirdnfor
1989\. spis.(Bsdon Alrh l
 evolutionaryredutionin digit number
l dvlpmntaliologv dr-rringtlr
)02). volutionary and populatitln IV
nnts lak o whos ()ntnts
ding volutionaryprossS.
^fl/
\l
n lrllngdrpardly. harls
..th laws of Frogs
;t' wondrdwhthr
typil variirtionnd favor ohs''
raryiologistsWr prpard to an-
lldr 198; aynard Srnith t al.
tion tl"rtth organizlrtionof dvl-
Lof phnotypssmsasona1- ill

,.''/lil'
. . r yp o s i h l p h r l o p i o u o r i s
Salamanders
a s ( A l r h1 9 8 9 ;! i l k i n s 2 0 0 2 ) .
Lundrlyingsoursnd long-trm

\l
;ti]ints.
has lmfrom two sours. on is
lt on n irlagin nd tht s-r
srvd.Th sond is th rstritd
g natr-rral or primrrtalprturl"la-
of th st ampls om from
. 1 9 8 3 ;A l r h a n d G l 1 9 8 . 5 ) . B eperimentalreductionin digit number
;l1,dmonstrtthat th prodution
f thsiasshav rmaind lusiv.
' it is diffiult to prolu dvlop- mitotiinhibitor
)mtri limitation' a lak of gnti
:. Distinguishingamorrgths possi.
hn it is possilto infr th xis-
pross of limir"rtion,idntifying
otypi variation raiss furthr hal-

ist.on planatioll is tlrilt dvl.

onsistntphnotypi outOms in
developing
limb
ations.This ida was rfully arti-
ylronamd it lizti,,n.Dvlop-
l a similar phnomnon thy all
nornrl hnotyp fol-
onvrg <lt-t Figur.3.Biasd produtiln o1 phnotyps. Tlr orgzrniztin of dvlopmnt
ldington livdanalization t m;ivrstrit t reng of phnltyps it is apal of ;lroduing. vidn oms
sswhrya phnotyp that WaS from primntal prturation of dvloping mplriiarl lims, whih show a
rsquntly gntillyfixd through rngof phnotyps tht is sirilar to that flund in rlatd spis. (A) Svral
an Shmalhaus(1949) proposd indpndntass of digit rdution hav ourrd during th volution of
lr th sam rsult' othr possil mphiins.AlthoLrsh four-tod spis hav volvl in frogs nd salamandrs,
thsrsult from th ltlss of diffrnt digits (dnotd y oman numrals). (B) By
a progrssivloss of flxiility as th
tratingth dvloping lirns tlf amphiins with n inhiitor of mitlsis'
:nd y historiirllg.ris(W.imstt
Albrlrnd Gal (198.5)wr al to xrimntally rdu digit nr-rmbrin
ons in arly dvlopmntwill hav a frogsand slamandrs. lrl h as th rirng of rsillting phntltyps is not
rnodifiations(Arthur 1988; Gould random ut instad onforms to th rang f phnltyps found among rlatd
spis.(Baslon Alrh and Gal l 98.5')
218 uolutio d Dulopmt

If dvlopmnt somtlms ats as a onstraint on volutionry hang, within a population y n

how do modifiations in dvlopmntaris?Josph Ndham (193) pointd givn individual; th inr
out that fundamntal dvlopmental prosss' suh s ll division' mor- the mill of sltion undt
phognsis' and diffrntiation, an oftn ontinu indpndntly aftr - nt gnti akgrounds {
primntal prturations. H alld this phnomnon dissoibilit and nti variants that migh
assmlda long list of ampls.Th undrlying basis for dissoiailitymay thry bom visil t
diffr in dtail from s to as' ut a ommon gnral fatur is proaly drift as nutrl llls.\X
modularity, th poprty of partial or omplt funtional indpndn- gr to whih sltions
twn dvlopmntalposss(Grhart and Kirshner 19971Hansn 2003). tion and th dgrtl wl
Th volutionary signifian of dissoiaility is that it allows modifiations fat of lds (.Wst-
in partiular aspts of dvlopmnt without fata||ompromising th ntr-
pis (Grhart nd Kirshner 1997).
In gnti trms dissoiaility is roughly quivalnt to irumsrid volutinary vfsus
pliotropy (Wilkins 2002).In an influntial ook Ronald Fishr (1930) ar-
gud that mutations of small effet are far mor likly t form th gnti a- Th fild of v<llutilnar
sis of phnotypi hang; his argumnt turnd on th intuitiv assumption tion of how gnotypi d
that roadr pliotropy is' in gnral, mor likly to rsult in nt ngativ Batson, who oind th
fitnssonsqun.Fishr's viw has n rfind (rr 2002) ut rmains a sidr th asis for phn
asi xptation for most volutionary gntiists. terils flr th Stud o1
Dvlopmntal posssr somtims strongly dissoiatd from volu- tiit i th origi of !
tionary hangs in dult anatomy. Striking instanesof divrgnin dvl- are homelti (anothr tt
opmntal mhanisms with no ovious impat on adult morphology hav dupliations in anatmy
n doumntd in svral diffrnt lads, inluding nmatods (Goldstin n t i s p o V i d s s u f t i i
et a|. 1998; F6li and Strnbrg 1998I, arthropods (Patl t a|. 1994; Grtc win's hypothsis of rrat
et a|.1998), and sa urhins (Hnry and Raff 1990; Wry and Raff 7990|. nrodrn synthsis omplt
Som of ths ss involv hangs in lif history' suggsting that th way no-Dawinians pstulat
dvlopmnt prods, rathr than its adult produt, has n th targt of typi fft (Fishr 190)
sltion,as disussdlatr in this ssay. Rihard Goldshmidt (
If som ways of organizing dvlopmnt limit hang, othr ways of orga- dominant rol for marol
niztng it may failitat hang. Th proptis of dissoiaility and analriza- that h proposd ld to ,.
tion may ontriut to th as with whih modifiations aris in dvlopmnt implausil. It was not ut
(Grhart and Kirshner 1997; Strn 2000). Som authors hav argud that mutations gaind intll
hangs in th organizatton of dvlopmnt' or in its gnti asis' an ky (Wilkins 2002). Th wy
volutionary innovations with far-rhing ffts.h asi ida of a ky ogists' who argud that
volutionary innovation is that a novl fatur lvatsth ratio of spiation s u l t < t fm t l t i o n si n g (
to xtintion, prmits n xpansion of anatomial disparity, or oth' ovr nd Monod 1961;Britt
marovolutionary timsals (Lvinton 2001). Possil dvlopmntal - son 1975). Th ris of d
ampls inlud th pansion of Ho gn num arly in vrtratvolu- highlightd th potntia1
tion (Holland 1'992)and th apparanof holomtabolous dvlopmntin notypi transformations (
insts (Yang 2001). Ths hypothssar diffiult to tst' sin ah rpr- Dvlopmntal rgulat
snts a uniqu historial vnt' ut thy rais th possiility that th organi- iologists for svralras
zation of dvlopmnt might hav long-trm positiv impat on th bitbor and te1le
evolutionary fat of lad. 1 9 7 8 ; S t t t a l . 1 9 8 3 ) .l
.s7
ariv at a prolm that has not n adquatly addrssd:is anal- Batson's (1'894) homot
izatjon an impdimnt to or a failitator of volutionary hang? on th on into lgs (Figur 4). A rr
hand, anlization allows th aumulation lf additiv gnti variation s i d p l i r l t t l y t: h i 1

nstraint on volutionay hang'
) J o s p hN d h a m ( 1 9 3 3 )p o i n t d
s s s ,s u h a s l l d i v i s i o n , m o r -
ontinu indpndntly aftr -
phnomnon dissoibilit and
rlyingbasisfor dissoiility may
:nmongnralftur is proaly
pltfuntionalindpndn-
rd Kirshnr 19971Hansen 2003).
lity is that it allows modifiations
ut ftirllyompromising th ntr-
4hly quivalnt t iumsrid
al ook Ronald Fishr (1'930\ ar-
mor likly to form th gnti a-
r n do n t h i n t u i i v a s s u m p t i o n
l i k l y t o r s u I ti n n t n g a t i v
l r t i n d( o r 2 0 0 2 ) u t r m a i r r sa
:ntiists.
s s o n g l yd i s s o i a t df r o m v o l u -
g instansof divrgn in dvl-
mpat on adult mophology hav
s,irrludingnmatods(Goldstin
.thropods(Patlt l. 1994; Grbi
.sray
Raff 1990; and Raff 1990).
[ history, suggstingthat th way
u l t p r o d u t .h s n r h t r g t o f
't limit hang,othr ways of orga-
rtis of dissoiailityand analiza.
' modifiations aris in dvlopmnt
)). Som authors hav agud that
-tt,o in its gntiasis,an ky
ng ffts.Th asi ida of a ky
tturlvatsth ratio of spiation
'ntomialdisparity, or oth, ovr
2001). Possil dvlopmntal -
numr arly in vtrat volu-
of Irolomtaolousdvlopmnt in
" diffiultto tst' sin ah rpr-
:aisth possiilitvthat th organi-
long-trm positiv impat on th
n adquatlyaddrssd:is anal-
of volutionryhang? On th on
r r i o n o f a d d i t i v q , n t i v a i a t i o n
uiutild Dulo1lmt 21L)
within a population y minimizing its immdiat phnoypi impat On ny
givnindividr-ra[;rlr inrasdgnti variatiotl rnight latr ot-t-t
gist for
th mill of sltiorrunder altrd nvironmntal iumstansor irr diffr-
ntgntiakgrounds (Gison and Dworkin2004). On th othr hnd, g-
nti variants that might othrwis produ phnotypi onsqunsand
thrbybonrvisil to sltionmight b maskd and thrforfftivly
drift s nutrl llls.W still hav only ir limitd undrstanding of th d-
grto whih sltionstalislrdrnhanismsrhat uffr phnotvpi veria-
tion and th dgro whih suh mhanisms limit lr ias th vrlutionay
fatof lads (\ist-rhard 2003).
volutionaryvrsusDvlopmntalGntis
Th fild of voIutionarygntis,as notd arlir, largly ignord th qus-
tin of how gnotypi diffrnspodu phnotpi diffrns.William
BatSOn'who oind th trm gtis,Was On of t first iologists t() otl.
sidrth bsis flr plrrrotypivrilrtion and diffrrlsamOng spis.His
triIsfor t Stud of Vrit jr rtd it F|spilRgrtl t Diso-
tiuititl th rigi of Spis (l894) mphasizlth rol of hrrgsthat
arehomeoti (anothr trm h oind, to dsri s;latial transpositilnsnd
d u p l i a t i o n si n n i l t o m y ) .B a t s o n ( \ 9 0 2 ) l a t r a r g u d t h a t n d l i a n g -
ntisprovids stlffiient planatilr.r for th hrditr.drrrandd1.Dar.
win's hypothsis of ntural sltiln. Durilrg th l930s arrl l940s th
nrodrnsynthsisorpltd this histlri intllttrlmrrgr' and th arly
no-Darwinianspostulatd a dominant rol flr mutations of srrrallphnl-
t v p i f f t( F i s h r1 9 3 0 ) .
Rihard GoIdshmidt (1940) ukd th trnd of tl.rtims to dvtlat a
donrinlrtrol flr n1r()l11Llttions. vn t th tim, th gllti mlrrris-ts
thath propos1ld to ..hopful rrronstrs''(Goldshnlidt 1960, .3)6) slnd
implausil.It wirs not until th 1980s that Goldshmidt's onpt lf mro-
mutationsgaind intlltual tration, alit in onsidraly altrd guis
(\filkins 2002). Tlr way Was prpi1ldy svral ir.rfluntial r-nolulriol-
gists'who agud tlrt phnotpi iliffrnsmol1g spis er lglv a
rsultlf muttiorrs in gn gr.rlzrtion rathr than protin flttrtilt-t (Jao
n d o n o d 1 9 6 l ; B r i t t n n d D : r v i d s o n\ 9 6 9 1 K i r r g a n d \ / i l s o nl 9 7 ' 5 ;W i l -
son 1975).Th ris of dvlopmrrtalgntisduring th sr-rsqunt dad
highlightdth potntial rll of rgulatlrygrrsin mdiating dramllti ph-
notpitransformirtions(Raff and Klrr'rfmen1983)'
Dvlpmnralgulttlr1'gns aptud th imirgir-ration of Voltlti()llry
iologists for svrlasons.Tlr most famous rgultoygnslong t<lth
bithor ard tpedi comp|xs in Drosophil ml<lgstcr(Lwis
1978;Sott t al. 1983). Sptulrrlutantphnltypsin ths grrsrall
Batson's(1894) onroti diffrrrs,transforming' for instan, lll]tnna
intolgs(Figur4). A rmarkal fatrrrof tlls t-tltttirions is tlrir lrum-
sribdpliotrop\':thir plrrrotv1.li impt is oitrl quit lirir1lrrtsidth
220 uolution d Dt,eloDmt

hart nd Kirshirr 19j

protil-lsthat rgulat tl
ommullitiln anlong
trnsdus privds
as mirosisand ll dat
r and t|ldi t
found t.l ntlrin lr sq
prorins' In histoi
h otobr, :rnd rnm
pondn of loi that
fid in mlltant srn
Figur 4. Hmoti mutations. Th fruit {| DrosophiI mellgs,erhs n pri slr' rvhat dvIl1.l
an important rsarlr sujt for oth volutionar,v and dvlopn-rntl
opmnt:ommunitio
iologists. Hmoti muttions in this spis ar along th mlst famus and
i n t u r n a t i v a t sd i f f
sptaular muttions tht hav n rovrd from any organism. (A) Had of
a fly that is homozygous for wild-typ i-rlllsatte tenpdi lous. Not gOWr|lm ' t,phogIls
th smll, fathy ntnnl] (hit rrrt,). (B) Had oi a fly that is homozygous Th rvlth of informi
for Atepdi |oss-of-funtion allls. This mr.rttiontransforms th m o s t o f i t d u r i r r gt h p
antnna in into normolls' narly pftlgs. A rmarkal fatur of this and pliations. Fist, tl-rprr
many othr homoti mutations is thi highly loalizd phnotypi ffts. not unusual for phnot
(Photos Itudi Turnr.)
strutur or ll t1.1..l (\
urrl,vmbryos s1tim
tal vnts. This irplist
dlimitd asptsof n
homotiall,v transformd organ. This fft is ir gnti manifstation of 2000).
Ndhnr's onpt of dissoiation and' w now know' a rsult of tlr way Stlnd, many nluti
gn prssion is rgr.rlatd(Wray t al.200). Anothr intrstingfttrr histlry, or mating, syst
of th fly homoti loi is th rsmlan of mutant phnotyps to th and Kishnr 1997).W
anatomis of distantly rlatd inst spis.For instan,som mutations in sis for suh diffn
(ib produ four-rvingd fiis, ririlig rr srl anatoIl1h has bn ttions lvould tvpi:rll1
asnt in diptrans for rnor thn 200 million yzrrs(Lwis I97s). d n l < l l r s t r ntthe r v n i
Dvlopmntal gntiists hav idntifid hundrds f loi whos mutant pts of phnotyp th
phnotyps indit ky rols in dvlopmnt. During th 1990s loi wr monstrs sm lss fa
disovrd in plnts that smlth homoti gns of aninrls in svrl thr>ught.
rgards:thy produ dramati transformationsof floral rgn idntity' tlrir Tlrid, and prl.rpsr
phnotypi impat is gnrally irumsrid' and som muttions prldu opn th lk <l'xthat
flowrs that smlth floral organiztion of sal angiospms(Jak t al. tuy. Until quit rntl
1992; Ror:nslyt al. 7995)' Not all dvlopmntalrgultory gnsprodu vn' to som tl]t'v
homoti phnotyps whn thy r muttd. Svral loi in th nmatod s strt nritis; fw
enorbbditis elgs produ htrohoni phnotyps (Amros and and nrutations might o
Horvitz 1984). any mutations ar known that fft dvlopmntal pro. opmntl gntiiststho
sss; thy dlt speifi organs lr ll typs, ltr ody propotions' tionar,v ontxt. Th l
hang th timing of rpodution lr feunditv, modify olo pattrns or p o i l u n d e n d p < l s i t i v i l
.oday,
othr intgumntary struturs' and rat a host of othr spifi phno- ogy. thr is w
typs (Wilkins 199; Gilrt 2006|' glls r sujt to pol
As soon rrsit m possil to haatrizths gns at tlr molular 2007).
lvl, it ws lar tht most nod protins rvitlr rgulatory funtions (Gr-

)rosophil mltlgsterhs n
tionaryand dvlopmntal
)sar mongth most famus and
(A) Hd of
id frol ilnv orgi.tnism.
, at thtttpdilous. Not
B) Had f a fly tht is hmzygous
rismutationtrnsformsth
qs.A rmarkalfirturof this and
.lyloalizdphIrtypiffts.
ftis a gnti manifsttion of
W now know, a sult of th way
2003). Anothr intrstingfeatur
n of mutant phnotyps to th
s.For instan,som mutations in
ln anstralanatomy tlrt has bn
lion yars (Lwis 1978)'
d hundrds of loi whos mutant
mnt. Dllring th 1990s loi wr
|motignsof animals irl svral
ations of floral organ idntity, thir
id, and som mrrtationsprodu
on lf salangiosprrns(Jak t al.
opmntalrgul:1tory gns produ
:atd.Svralloi in th nmatod
hroni phnotvps (Amos and
)wn that afft dvlopmntal pro-
l l r y p s , l r r d y p r o p o r t i o n s '
:undity,modif olor pattns or
l a host of oth spifi phno-
trizthsgnsat th rolular
lins witlr rgulatry funtions (Gr.
uoltion d,Dulopnlt 221
hart irrrd Kirslrrrr 1997; Wilkins 2002). Som ar trnsriptiln tlltors'
protinsthat rgulat th prssion of othr gns;othrs r involvd in
ommuniation alnong lls (signals' reptors, nd th rhinry tat
tnsdusa privdsignal);e.tdsom grrlatky llular posss,su
.Whn
as mitosisanl ll dth. th holrrotigns of th Drr;sr4lhil bitho'
r ald tpdllzomplxs Wr isolatd, for instan, thy wr all
found to tlntein il squn motif htristi of rtain DNA-inding
protins.In lristorial nll to Bats<tn,this rrrltift-nknown as th
hmobl,nd rmrsof this gn firrnily ar :rlldox gns.Th pr-
pondranof loi that nod rgulatory protins among th gns idnti-
fid in lutnt sns ws irtlltrral|,v stisfving aus it rfltd
prislywhat dvlopmntl iologists onsidr tlr or prosssof dvl.
opmnt:tlmmunitin among lls ndows thm with distirrtfats,whih
in turn tivats diffrnt irttis of gns that mdiat th prosss
growth' n-rorplrtlgnsis, ani di f frrrtiirtiln.
h rvalthof information that has rnrgdfrorn dvloprnntalgntis'
most of it during th past 25 yrs, hs svral imp()rtant volutionay im-
pliations.First' th PrOssL]s lf dvllpmnttlftt limit pliotropy. It is
not unusual fo plrnotypsto irtrmsridtl prtiLllr natonrial
stluturor ll tvp (!ilkins l 993). vn nrutations that altr vnrs in
arlyrryrssotims do so without rating havo in latr dvlopmn-
tal r,nts.This irrrplistlrat n1utationstht ffr tlr dvloprrrrrt of stritly
dlimitd aspts of anatorrry an aris witlrin ntural populations (Strn
2000
).
Sond,many mutiltions rnilrridiffrrrsam()llgspisi nnatomy' lif
histor,or matin.systms (Raff and Kaufmarr 1983; Fitlr |997., Crhart
and Kishnr 1997|' Whthr or not ths mutations also rnimi th gnti
asisfo suh diffns,anl whthr ()r not individuals that er thsrnu-
tationswould 'pially sLrvivirr th 'ild (poinrs disussd lter), thy do
dmonstratthat vn simpl mutations n prodLlprofound hangs in s-
pts of phnotyp that ar ologiirlly rlvant; Goldshmidt's hopful
monstssm lss farfthd, at last :]t a gnti lvl, tan W on
thought.
Third, and phaps rost gnrally, dvlopmntal gntis has thrlwn
opnth lak rl that volutilnaryiollgistslagly ignord for half n-
tur.Utltil quit rrrtl1,
population gl1tiists't]tlal]tittivgntiists,and
vn't()som tnt' vllurionarygrrtiiststriltd gns for tlr most part
as abstratntitis;fw thought dply ablut how diffrnt kinds of gns
and mrrtlrtinsmight ontritlt to phnotvpi vllrrtion.lr-rvrsly, dvl-
oprrrntalgntiiststhought a lot alLttths isslts,ut zrrlyin an vllu.
tionar1.ontt. Th intrstion trvn ths prsptivs has lrd a
profourrdand plsitivimpt ln oth dvirpmntalnd volutiorrriol-
ogv' Trlday, thr is widsprad intrst irr undstanding r,vtkinds of
gnsr sujt to positiv sltior-r(Nilsn t al. 200.i; Haygood t al.
2007\.
222 uolutil,d Dulopmt

bom an ion of volu

Body Plans and th Ho Parado th knwn gn fmilis
systms in nimls dat
h runifiation rfdvlopmntirland volutionary iology was wll undr many to tlr unillulz
way by th aly |99s. A fotuitous omintiorrif vnts-th arli pu- Kirshn 1 9 L)7; ar r t>I
liation of insightftrlooks (Gould 1977; Rff and auf-rn19s)' thnial tal rols of ths rgular
advans (most notably th infusion lf molular thniqus into virtually and r"lbidlpsz zlr ll
vry faet of iologv), improvmnts in nlytial tllls(spiallyphylog- zoans.
nti mthods), and a sris of iting disovris (nror on ths latr)_ Prhaps \.n1orr
prplld th mrger forwrd. uh of th initial itrnt rvllvd tins ar pssd in ro
around th volution of animal t>dyplans. frnt phyla (Grhart l
As th first dvloprnntalrgulator,vgnswr hrratrizl at rnol- gns providd th firs
ular lvl, a majo supris mrgd:homologous dvlopmntalrgulator.v othr orthologous rgt
gns ar shrd among organis-rswith vry diffrnt ody plarrs.Th first tantly rlt]animal ph
lar instan was th liztion tat li-12, Noth, nd Gl , impotnt to ovrlook th fat that
rgulatory gns indpndntly idntifid in nmatods, arthropods, and s i m i l a r d o m l i n so f x p r
h o d a t s ,a r h o r n o l o g o u s( G r n w l d 1 9 8 5 ; Y o h m t l ' 1 9 8 8 ) .B u t t h i s that thir dvlopr-r-rn
finding was quikly ovrshadowd y th disovry tht vrtrtsontain f i a t i t l no f a n i m l l s 1 D a v
lox gns homlogous to thos in l1is,and tht thy ar lustrd in th sirrrilaritisirr grrprt
g n o m i n t h s a m r l t i v o r d r ( A k a r 1 9 s 9 ) . I } yt h m i d - 1 9 9 0 st h l i s t o f trats hav n usr
rgulatory gns shard among th mlljtlr modl systn1sof dvloprnntal tor (DRortis and S
iologv ran to svaldozn (Raff 1996; Ghart nd Kirslrnr 1997). Sur. l o n g - t i n t n s t o r ss
vys of othr phyla rvald a rmarkirly road phylognti distriution, volvs many ssumption
irrluding asal nrtz)ngroups (suh as nidrians :rnd tnlphlrs)arrd of th ilitrian ns
anatomially oti ons (suh as inodrms ilnd rnollr:sks).It am Wilkins 20021.
ommonpla to spak of a shrd ,.tollkit''of rrimal dvlopmnt (arroll
tal.2001).
All of this was ompltly unxptd.Animals in diffrnt phyla ar so
anatomiaily distint that idntifying hlologoLlssrrutufsand dtrmin.
ing phylognti rlationshipsrmain ontntious (NiIsn 2oo2). This ph-
"l /
Okl
notypi distintnss hs fomd on lf th rrtrl tlrms of omp.rrativ ln.|
natomy for as long s th fild hs istd (HalI l992: Raff 1996\, Th no- Ap
tion f a ody plan is that ah phylum of ar-rimalshas uniqu lmination D
Dlt
of ntomial trits, most of whilr r rhitural(sulr s oiy s1,rm. hlt
try), mryologil (suh as tlr gomtry of ll division), or oth (suh as A
th nunrr <lfprirnary grnr layrs).h oiy-plnonpr has 1plvinflu.
nd idas out th divrsifitilntlf animls up tO th prsnt (GouId
1 9 8 9 ; H a l l | 9 9 2 ; k a i i 1 9 9 6 ; A r t h u r | 9 9 7 ) . D v l o p m r . r t li < l l g i s t sf ,o r Figur 5. h Ho parad<
thir pat, hv llng rognizd rnv prtlftlr.rnddiffrns in th wlry in flis nd mi dspitth ;
whih asi prosssof pattrnirrgand mlphlgrrsis ths nimls. h mosr f
tlLlrin mrvos of
stalish positiln f od
diffrent phvl (Gilrt anl Rtrni 1997). No otr pditd ht rrryoi
hlmtllog<lusgns pttrn
th sam asi st of gnswould turn ollt to rgulatdvlt;prnlrt thrugh- hordts. Th-s htlmolog
oLlt th animal kingdom. during dvlopmnt ur pl
Yt it is so. ontmporar txtloks<lfdvl<lpmntirl or vlutionarv i- Kirshn 1997; Wilkins 2
ology usually lntairran illustration that shows th sirnilar gnlrniorgani. of dvlopmntal gn nt
zation and prssirlnof Ho grlsin Drsrlphil nl tttous:this img hs Wry 2003.)
 ultil ttd D,l<lplet 22

boman ion of voiutilnarydvlopmntlilogy (Figur 5). Narly all

th knlwn gn farnilis that nod transription fators arrd ll signlirrg
systlnsin animals dat irk at last to th anstr o{ all ilatrians, and
manv to tlr unilIulr nstor of plarrts and :lnirnals (Grlrart and
iology Was Wll undr
llutior-rary
Kirshnr1997.,arroll t al.2001; wilkins202). Th urrnt dvlopmn.
intionof vnts-th rlir pu-
tl rols of tlrsrgulatory protins in rlodl organiss suh s Drlsopbilr,t
'aff ad Kaufrrranl98), thnial
and rbidlpsls ar larly diffrnt from thir original funtions in proto-
olulr thniqusinto virtually
zoans.
ralytiltools (spiliyphylog-
rhapsvn mor rrnrkaly, sonr of ths onsrvd rgultory pro-
isovris(mor on ths latr)-
tinsar prssdin roughly sinrilar rgions of mrvos that long to dif-
i th iitial itmt-ttrvolvd
f r r rpt h y l i l ( G r h r t a n l K i r s l r n e r 1 9 9 7 . , r r o l l t a l . 2 0 0 1 ) . h H o x
gnsprovidd th first irnd still th most dramati ampl, ut many
n., *.,. haratriz] at a nlol-
oth orthllogor-rsrgultory glls shorv similar prssion amlng dis-
ologousdvlopmrrtalrgtrlatory
tantly rlatd animal phyla (Figur 5). In all th xitmnt it has n asy
ry diffrntody plans. Th first
to ovrlook th ft tht .llstdvlopmntal rgultory gns do zol hv
I 2 ' | i , i h . . r n d ( i / r 1 ,i m p o r r n t
similar drains lf prssion irr diffrnt phyla; this osrvation suggsts
l in nmatods' arthropods, and
thatthir dvlpmntal ols hav volvd onsidraly during th divrsi.
) 8 5 ;Y o h m t a l . 1 9 8 8 ) . B u t t h i s
fiationof irnimals (Daviison200 l; '0ilkins2002). Th sorrrtims.striking
d i s o v r t h a t v r t l r l t s n t a i n
similaritisin gn prssion tht ar shard twn arthropods and vr-
and tirt thy ar lrrstrdin th
trtshav n usd to ronstrut th ntorny of th iltrian ns-
l 9 8 9 ) . B y t h m i d - 1 9 9 0 st h l i s t o f
tor (DRortis and Sasai 1996; arroll t al. 2001). Ronstruting
: modl systmsof dvloprnntal
long-tintl]nstorssolly on tlr sis of g-prssionsimilaritis in-
] r h r t a n d K i s l r n r1 9 9 7 ) . S u r -
volvsmany ssulnptions nd muh gusswork, howvr, and th anatoy
y rod phyllgntidistriution,
o f t h i l i t r i a na n s t o r m a i n s u n r t a i n ( r w i n a n d D a v i d s o n 2 0 2
; n i d r i a n s n d t n o p h o r s ) n d
\)flilkins2002).
:ldrmsnd rollusks). It am
it'' of iml 1vlopnrnt (arroll

LH2 Ho-t
Anirals in diffrnt phyla ar so Dl
nologous strLltursnd dtrmin.
tntiotls(Nilsn 2002)' This ph. td
'lr ntai thnls of omptiv |nS
BMP4
.d (Hall 1992 kaff |996). Th no. Ap t:holin
D
a n i m a l sh a s a t r n i q u o m i n t i o n D
rhittural(suh as ody symm. hh Nkr2.2
' of ll divisitln)o Ap Ms
, r oth (suh s lG-H
lody-pIanol1ptlras dply inflr.r-
animals up ttl th prsnt (Gould
Figur5. Ho prado. Homllogous gns pi]ttrn t mrvos of fruit
97). Dvloprrrntaliolgists, fo
flisnd mi dspit th plrylogrrtidistan nd phrrotypi disprity lf
rofourddiffnsin th way in
thesanimls. Th mlst fmous of ths r gns lf th Ho lmplx, whih
rrorphgnsislur ir-rmryos of stalishposition of l.od rgions |ong th ntoPostrior is. Mnv othr
7).No on prditlthat rnany of honr<llogous gns pttrn th rins, hrts, ys, and lims of arthrolds and
t to rgulatlvlopmntthrough- hordts.Tlrs htlnrlogol.lsglls ar prssd in roughly similar pattrns
duringdvlrlpmnt ut odu r,ry diffnt phnor1'ps(Grhart ar-rd
l dvlopmntlor rllutilnaryi- Kirshnr 1997; Wilkins22). Tlris asi rsr.rltsuggststnsiv ..rwiring''
of dv,lopmrltalgn ntworks drrring th ours of volution (s txt). (From
s h o w ' t h r i r r l i I l rg t l < l t n i() r g n i -
lfray 2003.)
osl1lhil and mous: this imag has
224 uolutio d Dulopmt

What is lar at this point is that th sam asi omplmnt of dvlop- and rgulatoy squn
mntal rgulatory gns' som of whih ar prssd in roughly th sam r- a o l i s h i n gs u h i n t r r
gions of th mryo' an produ a wid divrsity of nimal ody plans. of muttions, singl as
Figuring out how this lurs is on of th grand hallngs of volutionary t l. 200J). Mny prot
dvlopmntal iology (Ghart and Kirshner 1997; arroll t al. 2001; nuli aids, and small
Davidson 2001). A dtaild rsolution of this Ho parado lis many yars d r a m t i f f t s. hngi
in th futur, ut hangs in th organization of dvlopmntl gn nt- tion or altr affinity dr:
works ar likly to fom an importnt part of th solution (arroll t al. an rat or modify a i
.Wry
2001; Wilkins 2002; t l. 200). rgion.
Two asi kinds of rwiring within gn ntworks hav n doumentd, hangs in dvlopm
roughly orrsponding to quantitativ and qualitativ hangs. Th first, or pat on th history of lif
quantittiv' atgory involvs a hang in th loation or timing of prs- and why gn ntwlrks
sion of a rgulatory gn. Suh hangs an aris whn a ntwork is altrd
upstram of th gn of intst.Th loation or timing of th dvlopmntal
prosssth gn ontrols will hang as a rsult, potntially altring organ- Frm ly{arovoluti
ismal phnotyp. For xampl, th spatial xtnt of Ho gn xprssion dif-
frs in ways that orrspond to sgmnt idntity in rustaans (Avrof and Phnotypi hangs,inlt
Pate| 1997) and to vrtrl anatomy in ttrapods (Burk t al. 1995). Th variation within populati
sond, or qualitativ, kind of ntwork rwiring ntails a hang in whih d- mntal volution at thrs l
vlopmntal prosss ar rgulatd. Ths hangs an our whn a nt. aout th gnti asis ft
work is altrd dwnstrm of th gn of intrst.For instan,signalingvia wild populations (as opp
th rptor Noth gults spifiation of nurnal ll fat in insts and th la) or aout what k
spifiation of whit lood ll fat in vrtrats (Hitzlr and Simpson tion in ntwork funtlon
1991; Guidos 2002), involving two vry diffrnt sts of fftor gns. Sim. happn ovr llng tims
ilarly, th tansription fator rahyury rgulatsnotohord dvlopmntin This pauity of inform
urhordats and hordats ut plays no suh rol in othr phyla, whih lak D v | l p m n t al i o l o g i s
this strutur; instad, it rgulats dvlopmnt of th gut and othr stru- influns of gnti an
turs (Thnau and Sholtz 2003). hv workd largly witl
Dozns of ass r now known whr an volutionary hang in th x- gnt that omprisons 1
prssion prfil or downstrm targts f a rgulatory protin has altrd nisms. For thir part' po
anatomy. Som ass nompass hangs at th sal of ntir ody plans, a n d h v f t l u s dp r i m r
suh as th transition from rdial to ilatral symmtry during arly mt- protins, protin strutL
zoan volution (Martindl t al. 2002) and th latr transition ak to radial rathr than gn intrt
.Wray
symmtry in hinodrms (Low and 7997). Othrs involv modifia- hr ar, of ours'
tions in organs' suh as th tansition from fin to lim during th origin of variation in th oding :
ttrapods (Shuin et a|.7997), or volutinary innovations, suh as arthro. u l a t o r y g n s ( . g . ,A y a
pod wings (Avrof and ohn 1997). vn subtl diffrns in anatomy ar T h s o p u l a t i o n - l v ll
oftn th rsult of modifiations in gn ntworks: ampls inlud diffr- o f s g r g t i n gv a r i i o n
ns in ristl pattrns in flis (Strn 1998; Skar and Simpson 2000) and opmntl mhanisrns
olor pattrns on uttrflywings (Brakfildet a|. 1996i Bruntti t al. 2001; toris (.g.' Wray and
Gompl t al. 2005). 1998), or among popu
How do hangs in dvlopmntal gn ntwoks aris? Thr ar many t a l . 2 0 0 4 ) , a n r v a ll
ways in whih this an happn-any spifi intration etwn maromol. hangs in sltion. (
uls is a link in a gn ntwork' and any diffrn in intrations altrs th out population gn
ntwork. Th two most ommon and important kinds of ntwork hangs opmnt rmlins a gleri
ar proaly intrations twn protins and intrations twn protins vlopmntal iology.

am bsi omplmnt of dvlop-
' prssdin roughly th sam r-
l divsit1,of niml ody plans.
grand hallngsof volutionary
'shnr19971 arroll t al. 273
this Hox pradox lis many yars
zation of dvlopmntal gn nt-
lart ot th solution (Carroll t al.
e ntworkshav n doumntd,
d qualitativhangs.Th first, or
n th loation or timing of xprs-
:an aris whn a ntwork is altrd
tion or timing of th dvlopmntal
a s u l tp
' o r n t i e l l ya l t e r i n go r g a n -
xtnt of Hox gn xprssion dif-
idntityin Llstaans(Avof and
ttrpods(Burk t al. 1995). Th
wiring ntailsa hang in whih d-
]s hangsan our whn a nt-
: intrst.For instan,signaling via
Lof nuronlll fat in insts nd
vrtrats(Hitzlr and Simpson
jiffrnt sts of fftor gns. Sim-
:gulatsntohord dvlopmnt in
,uhol in othr phyla, whih lak
)pmnt of tlr gut and oth stru-
n volutionryhang in th -
f a gulatoryprotin has altrd
s at th sl of ntir body plans,
ttrl symmtry during aly mta-
nd th latr transition ak to radial
a 7997). othrs ir.rvoivmodifia-
om fin to lim during th origin of
ionar innovations,suh as rthro-
:n sutl diflrnsin anatomy ar
ntworks: ampls inlud diff-
)98; Skar and Simpson 2000) and
ld t al. 1'9961'Brunettit al. 2001;
r ntwoks aris? Thr ar many
:ifi itrationtwnmaromol-
l diffrnin intrirtions altrs th
:lportntkinds of ntwork hangs
r s n d i n t r . t r i o nbs t w
uolutil dd D eU|opmt 22.t
and gulilt()rysqttnsin DNA (Davidson 200 l). Crating, rnodifvirrg'or
aolishingsuh intrations an <lurwith th sirnplstand most ommon
of mutations' singl as sustitr-rti<lns
(Grhrt nd Kirsirn 1997 wra
t al. 200). Many protins ssoiatspifiallywith othr protins or witlr
nuliails,and sr.n:rll
hangsin inding affinity or spifiit,van produ
dranratiffts.hngirrg singl nrirroilid n rt a spifi intra-
tion or altr affinity dramatially; similarly' hanging singl nulotid
enrtlr nrodify inding sit fr tr.lnsripti()nfator ilr a rgulato1'
rglon.
hrngs i dvlopmntalgn trtwokslaly hv had a profound im_
pat On th histlry of lif. As disussd nxt, hwvr, undrstandirrghow
nd ,hy gn ntwrks r ..rwird'' rrnainsa ntrl hallng.
Frm Marovolution to irovolution
hnotypiangs,inluding vt>lutionay hangsin lvloprrrnt, gin s
varii.ttion within populatilns.Rltivly fw str.rdis hv addrssddvlop-
mntalvolution at this mirovolutionary sal. As sult, w know littl
aboLrtth gnti sis flr nrodi6atir-rs in dvloprntalgn ntrvorks irr
wild ppulations (as opposd to What indud mutations an aOmplish in
thla) or aotlt wt kinls of volutilrraryml-r:rnisls s()rtgnti \,ri-
tion in ntwork funtion (s opposd tl simply osrving tlrt hn{:saIr
happnovr long tirnsals)(Purugganar2000; Wry t al. 200).
This pauity of informtion drivs from tw<lhistoril trrrdsin rsarlr.
Dvlopmntalilogists, in th first instan, lrav stivn to liminat th
influnsof grrtird rrvironmrrtlvri:rti<lfrorrr thir rral'ssan.l
havworkd largly with modl systmsthat ar so phylogrrtiallydivr-
gnt that omparisons ild littl inflrrrrtionbotrt volrrtilnaryr-nh.
nisms.Fo thir part' popr-rlation gntiistshav largl1,ignord dvl<lpmnt
andhavfor"rsd pimarily on nzymsralthrthirn dvlopmntalrgtllatory
protins,rotin strutur rtlrr tharr gn prssilln, alrd singl gngs
rathrthan gn intratilns.
Thr ar' of <lurs,prions. Svral stlldis hitv xirnlind gni
vritionin th tldirrg nd ls-rgulatlrysqunsoI dvlopmntal rg-
u l a t o r yg n s ( . g . ,A y a l a a n d H a r t l 1 9 9 ; I - u 1 w i ga n d K r i t m n 1 9 9 . 5 ) .
Thspopulation-lvl arralyssprovid a glirr-rpslf th n.1turand lvl
of sgrgatingvariirtion ftlr fw rgul:ltory gns. lmparisons of dvl_
o r n n t anl t h a n i s m sa o n ] l < l s l yr l a t d s p i sw i t h l i s t i n tl i [ h i s -
t o r i s( . g . ,l / r a y a n d R a f f 1 9 9 0 ; S w a l l a n d J f f r v 1 9 9 6 ; G r i t a l .
1 9 9 8 ) ,o r a ] o n g p o p r - r l a t i l nrsv i t h d i s t i n t r o r p h o l l g l s( . g . ' S l r a p i r o
t a l . 2 0 0 4 ) ' n r v a l h o w s p i f i d v l l p m n t arlr r h n i s m rs s p o n d t o
h a n g si n s l t i r n .O v r a 1 l , t l r o u g h , t h g n r a l l a k t l f i n f o r l a t i n
a o u tp o p u l t i o n g n t i sa n d s r a l l . s a l v l u t i o r r r y h : l n g si n d v l -
o p m e n tr m a i n s a g l a i n g g p i n t h r r t n i f i a t i o no f v o l u t i o n a r y r r n d d _
np o t i n s vloptr-rti.rl iology.
226 uolutild Duelopmet

On unrsolvd ut rathr asi qustion is what kind of gntivarition ls-rgultory rgions ((

ontriuts to volutionary hangs in dvlopmnt. Th larg-fft muta- gns suggstthat pr
tions lovd y dvlopmntal gntiists ltr phnotyp in dramati ways, (ihalak and Nolr 20(
somtimsmimikins diffrnsamong spis.Ths mutations ar proa-
l not gnrally good mdls for th gnti asis of volutionary hang, in
part aus th phnotyps ar rarly Wll intgratd in trms of organismal ologial Dvlop
funtion (Budd 1999; arroll t al. 2001). This dos not man that mutations
of iarg fft play no rol in th volution of dvlopmnt, ut dtting L i g h v a n V a l r r( 1 9 7 3 , 4
thir influn is a signifiant hallng. At th ppulation sal, variants of mnt by ology.'' Th s
larg fft ar ptd to xdingly rar, and mutations of small fft trations twn whl
ar xptdto prdominat in th wild; vn at th lvl of los intrspis disipIins within iolo
omprisons, multipl mutations ar typially fid at ah lous' and dis- m U s t - r J a f u n i o n
sting out thir individul ontriutions is dingly diffiult (\/ilkins in a mann onsistntW
2002). a mans to dstination;r
Anothr asi qustion is how muh gnti variation that influns d- oth aspts of dvlopn
vlopmnt is prsnt in natural populations. Additiv gnti variation that far lss ttntion (Gilrt
undriis a spifi aspof anatomy provids an indirt masur' sin it tion in th dvllpnrrr
must at som lvl rprsnt modifiations in dvlopmnt. But y itslf' rOutsto th sm phno
quantitativ gntis provids no information aout what kinds of gns ar o U t s r . r t u . l l Iqy u i t r
rsponsil fo trait diffns,nor how muttions in thos gns altr d. tion on dvlopmnt p r
vlopmntal prosss.Rntly, th quantittivgntisof gn prssion ral phnomna.
has attratd ttntion s a Way to bgin ridging this gap. It has beom Dvlopmnt in many
lr that natural populations haror abundant gntivariation that affts gnrations nd lntras
gn prssion (.g.,olksiak et a|.22i Rokmn and \Way2021\Yttt- dvlopnrntal digrssr
kopp t a\. 204). Howvr, stlishinga onntion twn gntially rprodutiv adult. In pri
s dd i f f r n si n 8 n p r s s i o na n d r g a n i s m a lp h n o t y p p o s si t s o w n o n a u r e I s l t i o nf o r
hallngs.Molular and quantitativ gntisprlvid ways of tsting this gag (fturs that hav
link, and onvining assar ginningto appar (Brm t a\.2002; Shapiro othr aspts of organls
et a|. 204; Gompl t al. 2005). ar proaly tru tO soln
A third asi qustion is what volutionary mhanismsoprat on gnti to dptiv (Garstang
variation that influns dvlopmnt. Th long-trm onsrvation f m- aid in disrsal, provid ir
ryos that forms th asis of Bar's laws is famous, ut it is unlar whthr loal xtintion (dw
its aus is dvlopmntal onstraint or prsistnt ngativ or stailizing latd to spis that lak
sltion.A numr of studishav impliatd partiular volutionary mh- mnt an aompany thi
anisms that oprat on dvlopmntal prosss' inluding rlad nga- Grbi t al. 1998; Bkh
t i v s l t i o n ( S t r i k l r t a l . 2 0 0 1 ) , h a r a t r d i s p l a m n t( W r a 1 9 9 6 ; ing a partiular adult od
G r i t a L .| 9 9 8 ) , s u a l s l t i o n( K o p p t a l . 2 0 0 0 ) ' a n d d i f t ( T r u a n d r o u t ( S t r a t h m a n n1 9 8 . 5
Hag 2001). shifts in ways that nhan
During th past fw yars dvlopmntalgn ntworks hav lso nim- ltion has opratd on th
pliatd in spiation.Th Dozhansky.Mullr modl of spitionproposs Altrnativ dvlopm
that gnti isoltion an aris aus of inomptiilitis in intrations within spis.Rhaditi
among gns at many loi (Dozhansk 796; ullr 1942|. Baus thy mpl: adults drivd fro
involv so many intrations' dvlopmntalgn ntworks ar good andi- idntial, ut undrlyr
dats for suh inompatiilitis (Johnson and Portr 2000; !ilkins 2002). (6lix2004). (Daurstag
Svral loi involvd in produtivisoltion hav bn haratrizdat th a n l t r n t i v s e g t a k ni
molular lvl. and most ithr nod transriotion fators r rsidwithin t o s u v i v u n t i l o n d i r i t l

l i s w h t k i n d o f 8 n t i v r i a t i o n
llopmnt.Th larg-fft muta-
l t p h n o y P i n d r l l m a t i w a y s .
lis.Ths mutations ar proa-
'i asisof volutionary hallg' in
l intgrtdin trms of orgnismal
T h i sd sn o t m n t h t m u a t i o n s
on of dvlopmnt' ut dtting
t th plpulationsal, varints of
rar,and mutations of small fft
' n t t h l v lo f l o s i n t r s p i s
ally fixd at ah lous, and dis_
; is dinglydiffiult (Wilkins
:ntir,ritiorrthat irrflunsd_
ns. Additiv gnti variation that
lvids an indirt msur' sin it
lns in dvlopmnt.But y itslf,
rion aout what kinds of gns ar
mutationsirr thos gns altr d-
rtitativg n t i so f g n p r s s i o n
r ridging this gap. It hs om
ndant gnti variation that ffts
}; Rokman and.Wray 2002; \/itt-
l a onntion twn gntially
, r g a n i s m pl h n o t y p p o s si t s o w n
ntisprovid ways of tsting this
O appar(Brnrt a|.202.,Shapir
r a r ym h a n i s m o s p r t o n g n t i
.h
l o n g . t r m o n s r v t i o no f m -
is famous' ut it is unl whth
. p r s i s t nnt g a r i v o r s t i l i z i n g
;atd partiularvolutionary mh-
prosss'inluding rlad ng-
art displzrmnt{Wra 1996;
rp t al. 2000), and drift (Tru and
l gnntworks hav also n im.
'{ullr rnodl f spiirtion poposes
o f i n o m p r i i l i r i isn i n r r a t i o n s
1936; Mull 1942\' Baus thy
ntal gn ntworks a good andi-
n and Portr 2000; \7ilkins 2002).
ltion hv n haratrizd at th
ransriptionfators or rsid within
L't,llutillt
l Dt,ellplttt 227
rgions (orr and Prsgravs 2000), w,hil survys of mny
ls-rsultlrv
gnssuggstthar prssion diffrnsplay rllin pr<llutiv
isllation
(ihalakand Nolr 2003).
lgialDevlopmntalvolutionary Biology
Lighvan Vln (1973, 488) nltdthat ..volutionis the ontol of dvlop-
mnty llogy..'h sirnpliit tlf this statmntlis tlr divrsity of in-
trationstwn what hav n studid trditionally s thr sparar
disiplinswithirr iology (V/st-rlrrd2003). f)vlopmrrt not orr[y
mustr-rata funtioning adult organism vry gnration ut nrust do so
in a rrlnnronsistntwitlr srrrvivl.In othr rvords, lvlopmntis lrot just
a mansto dstination;it is a partiular rout to that dstinti<ln.Although
oth sptsof dvlopmnt ar larl1.inrportant' tlr sond has rivd
far lssattntion(Gilrt 2001; wst-rhr1200). Thr is littl rogrri-
tion in the dvlopmntal iology litri1turtlrat altrnativ dvllpmntal
roLltsto th sanrphnorypi OLltomvn is.Altrnativ dvlopmrrtal
routsar atually quit ommn, howvr, and th imprint of natural sl-
tion orrdr,lopmrrt pr s' raththn tlrotrghits produt, is vidritin sr,-
ralphnomna.
Dvlpn-rnt in m:rny ttkryotsis indirt: ltr-rating hploil-diploid
gnerationsand ()ntrastinglarval-adLrltmorphologis ar two widsprd
dvlopmntaldigssi<lns from dirt rout that links zgot'to sually
rprodutivdult. Irr pinipl, ompl lif yls luid ithr th rsult
of ntural sltion for a partiular dvlopmntal rlut or historial g-
gag(fatLrrs that hv lost thir ftlntiln ut annot dltd aus
othrasptsof organismal funtion ar uilt around thm)' Although oth
arpoalvtru t() som xtnt' rnany fatursof tmpllif yls PPr
t b adaptiv (Grstang 1929; Strathnrann 1985). F<linstn,larva ry
aid irl dispsl,provid ssto flod,vad prdatos,or minimiz risks of
loalxtirrtion(N{dlv:td1995). lvletnyspiswith iarv r losly r_
latdto spisthat lak thm, and tnsiv diffrnsin arly dvlop-
.W.re,v
mnt n 2lompnythis lif-historv ()lltrast (.g., rrd Raff 1990;
Grbi t a|. 199\ tskhamt al. 2003). Suh ss dmonstrattlrat uild-
ing ir prtitrlrdr"rltod,vpln ]lsnot rqui partiula dvlopmntl
rout(Strathmann198.5).Man lf ths ass ar orlatd with ologil
shiftsin ways that nhan orgar-rismlfuntion' suggstingtht ntural s-
ltionhas tlpratdon tlr undrlying dvllnrntlrnhnisms.
Altrnativdvlopmrrtalrouts to th sam phr"rtyp mmonly ist
,ithina spis.Rhaditid nriltodspovid partiulrl wll-srudidx-
ampl:adults drivd from nortnal lr daur l:rrval stgs ar antonrially
idntial,ut undrlying dvltlpmIrtlpr()sssn 1uitdiffl-lt
(Fli2004). (Daur stags'fror th Crrnan word raning ..nduing,''ir
analtrntiv stagtakn in unfavralnvirrlmnts,nalingth oganisn]s
to survivuntil onditi<ltrsimprov.) Asu[ rproduion nd rgnraion
228 uolutio d Deulopmnt

provid additionl ampls. Both phnomn r ommon and ar prsnt

in phylogntially divrs multillular orgnisrns;oth dn-ronstrthow
Prospts
vstly diffrnt dvlopntirlr<lutsn onv l on olnmon nd points.
Aftr half a ntury of l
In spiswhr oth llnirrgan]sxual prodtrtitlI1 our] adults ar litr.
lutionry illogists
ally indistinguishl dspit having stirrtd lvlopmntin vry ]iffrnt
nmnt. ontemporar1
ways. Lik th lif-liistory hangs disussdin th prding paragraph. r-
imprint of volutinary
generationand sul produtionar dvlopmntaladpttionsgnlly ..toolkit''
of potins th:
assoiatd with partiular nvirnmntal irumstns.Asxul rprodu-
karyots, and omparat
tion may volv from rgnration(Bly and lVr:ty 2001), and gnration
dvlopmntl gn nt
may volv as an laoration of wound-haling mhnisms.
part' hs n influn
Dvlopmnt is rrly lrrdwird. Most multillLrlr organisms shlw
opmntz.rl iologists ha.
sorrrdgrof phntypi pl:rstiityin rsponsto th nvironmnt thy .
.Wst-lrard twen gnotyp and pht
prin during dvlopr-rnt (Shliting irnd Pigliui 1998;
of singl loi to modl
2003)' Ths rspol.lssin som ass r simph,sid ffts of prturation
qui1ntitativ gntis ts
that ar not uffrd ,vanalization, ut in tnany tlthr ass rsponssto
funtional anlysis ratht
th nvironmnt ar adrptiv.mpls inlud' th growth hait and lf
vlopmntal and volut
shap of many plants, irrduil dfnssir-rmany plants and animls, and
old p:rrtnrs ar k wl
partitioning rsourstwngrowth and rprodLrtionin many plants nd
poriv rlationship.
anirnals (Havll 1990; Slrliting and Pigliui 1998; \/st-brhard 200).
Dranrti ass of phnltr,piplastiity involv s dtrmintion, whih is
tt-l-\erture
dpndnt in sonl turtls and ollilirrs (rws t al' 1994) AKNoWLEDG'1l
and is st y soial olltxt in som fishs (Grldwil.rr al. 2003).
Plyphnism is a sttstof phnotypi plastiitvwhr disrtmorphs d. Thnks to Dan Sha
suportdy th National!
vlop in sponsto nvirrnmntalus. Pllyphnismhas n partiularly
SpAdministration.
wll studid in insts:rnany uttrflisand phids hav sasonal morphs,
som watr stridrs produ wingd and nonwingd rnorphs dpnding on w-
tr lvl, and, prhaps most sptaularly, th slil irrstsdvllp into dis- E}ILIoGRAPHY
.s7st_rherr1
tint sts(Niihout nd 'rVlrlr1982; 2003). In polyphnisnl
A o u h i f . . H . , N 4 .A k m ,
tlr srn gnotyp an prodll two or mtlr distil-rtphrror1,ps that r t
N . H . P t |R' . A . R t
last as diffrnt s spis within a gnus nd il-l som ass a gld dirl dvlopmntalgns.T
mor so. A o u h i f , . H . , n d G . A . '
h volutionary oigins and dvlopnrntale.rss of phnotypi plastiity volutionof wing poly
ar not wll undrstood (Shliting and Pigliui 1998; wst-brhrd A z h l t o vA ' . ' A . P o p a d i :'
200). onsidral dt has ntrdaround th qustilnswhthr thr hornlgy of arthropo
..plastiity gt-ts''and whthr plastiity is a produt or y. A k m , N , I .1 9 8 9 .H o a n dF
ar distint
vtrts. ell 57: 34
produt f sltion (Via t l. 1995; Shiiting nl Pigliui 1998). Th - A l b h .P . l 9 8 c . T h l l g i
prssin of mny gns is srrsirivto nvironmnt:rlotllitions, and this dvloprnnt nd volt-
snsitivityprovids il gnal mans of trans]r-ritlg trnl us into phno. A I r h ,P . , a n d . A . G a l .
typi rsponssnd su.]sts that thr is no nd to posit distirrtplastiitv trnd:DigitIduti
Alrh, P., S.J. Gould, G. J
gns (.!Vrayet al.200)' Diffrns in gn prssilnaompany th d.
lltognyand phylog
vlopmnt of diffrnt sts in s and ants (vns nd Whlr 2001; Alrt, P. S., and D. L. Ridd
Aouhif and $ry 2002)' whil th gnti asis for polyphnism hs n dr.rrlik|ara' Dul
workd out in dtail for tlr daur larv f th nrnatodeorhbiditis l- Amos, V., and H. R. orr
gns (t|ert and Riddl 1988). Ths approahs suggstthat hangs in lrhbditis lgs
gl-lxprssionmay b t last s important as harrgsin gn funtilndu- Athur, W. l988. AThal1,

ing th volution of plastiity. John Wily and Sos.


lna ar ommon nd ar pst1t
rganisms;oth dmonstrat how
OnvrgOn ommon nd points.
our' adults a litr-
:prldutitln
d dvlopmnt in vry diffrnt
d in th prdingparagraph, r-
lvlopmntaladapttionsgnrally
irumstans.Asual rprodu.
.ndWrav 200 l ). lld rgntin
[ingmhanisms.
lst multillular organisms show
iponsto th nvillnmnt thy -
LrrdPigliui 1 998; !st-rhard
simply sid fftsof prturation
' in mt-tythr ss rsponss to
inlud th growth hait nd lf
in many ;llants ard animls, and
d rpllr'rtitln in rny plants arrd
liui1998; l W s t -
nvolvs dtrmintion,rvhih is
r d r o o d i l i a n s( ( i r w s t a l . 1 9 9 4 )
( G d w i n t a l . 2 0 0 ) .
llilstiity .lrrdist mophs d-
Polyphr"rism has n prtiularl
and aphids hav saslnalmorphs,
onr,vingd rtlphsdpndingon wa-
, th soial instsdvllpinto dis-
/st-rhrd 2003), In polyphnism
ro distint phnotypes tht r t
rus and in som ass a good dal
lntal i1ssof plrrrotypiplastiity
nd Pigliui 1998; \Wst-rhard
ound th qustirlswhrlrr thr
th plastiity is prodr-rtor y-
r l i t i n g n d P i g l i u
nvironmntalonditions, nd this
ansduirlgtrnlus irrt<lphrro.
s no nd to posit distint plastiity
8Ilprssionaompany th d-
nd ants (vnsand Whlr 2001;
rtiasisfor polyphnism has n
of tlr nratod ()0orbbiditis eI-
pproalrssuggstthat lrangs in
tnt as hngsin gn funtion dur-
t,olutiod DueloDmt 229
Prospts
Aftr half a ntury of largly ignoring h othr, dvlopmntal and vo-
lutionaryiologists rntly ntrd a phas of prfound riproal nlight-
nmnt. ontmporry dvlopmntal iology as an unmistakal
imprint of volr-rtionary onpts:phylognti mthods rvld th shard
..toolkit'' protins
of that rgulat dvlopmntal prosssthroughout u-
karyots'and omparativ mthods ar now routinly usd to hartriz
dvlopmntalgn ntwrks. ontrp<lrayvolutionry biology, for its
part' has n inflund by th pwful funtional prsptivthat dvl-
opmntl iologists hav rought to undrstanding th rlationship -
twngnotyp nd phnotyp: attntion is shifting from astrat modls
of singl loi to modls of gn ntworks asd on ral organisrns, and
quantitativgntis is now viw1as th first stp in idntifying loi for
funtionalanalysis rather than s an nd in itslf. Th rintgration of d-
vlopmntaland volutionary iology is not yt omplt, ut ths two
o l d p a r t n r sa r a k w h t h y l o n g :i n o n s n s u a l n d m u t u a l l y s u p -
portivrlatinship.
r h a r2d0 0 ) .
AKNoWLEDGMNs
Thanksto Dn MShaand ihal Rusfor insightfulommnts'My rsirrh is
supportdby th NatioalSiIr
Furrdtion
nd th Ntioalonutisnd
SpaAdrninistration.
B|LIoGRAPHY
.W.
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20 uolutio d Duekrymnt

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N4ost animals liv solitar1

individuals forag, mov
highly soial animals our
long.trm rlationships a
spis liv in groups an
want to know why som I
'W
want to know why o
tiv and omptitiv tndt
many soitis. \/also w
uals and th sxs dvlo
not. Answrs to thsqu
to animals to larn aout .
answring ths nd othr
y problms. Bhavior, lik
th far that what animals
to shy away from asking
aout th adaptiv valu

Foundations in Natu

U n d r s a n d i n gw h y s p i
q u i r s d r w i n g i d sf r o m
half ntury ths idas, t
bn wovn into a ohr
iorists had to larify hw t
th mundn to th fasrn
to ask' ..Why ar thy
(196) proposd that thr
tions. Thr wr standard
] volutiOnf devlopmnt.
bg1'47:675-684.
r o i h i n o d r md v l o p m n ti s
t |20, Supplmnt:97_106.
l h o f f ,M . P i z r .M . V . R o k m n '

Soial Bhaviorand
transription1 rgultion rn
20 177-I4I9.
ralgulatorygns and hinodrm

Lsof linagfoundr lls in th

Soiobiology
th r ogrmm . D ullpment I

adaptivradiations:A ompartson Dnil l. Rubenstein

uolutitl Dullpmet : 59_72'
. h rlrh bditis !gs li- 12
h ovrll similarity to Drlsopbil

Most nimals liv solitary livs' Apart from oming togthr to mat' most
individuls forag, mov aout, anl void prdatos on thir own. Yt as
highly soial animals ourslvs, W a fasinatd by spis that dvlop
.W
long.trm riationships and liv soial livs. want to know why suh
.!e
spisliv in groups and why th groups tak th form that thy do.
want to know why som groups prsist, whras Othrs ar mor transitory.
W want to know why ooprtion oms in many typs and why oopra-
tiv and omptitiv tndrrisshp most rlationshipsnd thus oist in
mny soitis.\Walso want to know why partiular rols twn individ-
uals and th ss dvlop and why som appar univrsl, whil othrs do
not. Answrs to ths qustions ontinu to m]'ut sin w oftn look
to animalsto lrn aout ourslvs'th ility to fous th lens of sinon
answingthsand other qustionsabout human soiality has n loudd
y prolms.Bhavior, lik all oth fatursof th phnotyp,volvs, and
thfar that what nimals do, humans must also do has oftn ausd popl
to shy away from asking' or at tims vn bing allowd to ask' qustions
aboutth adaptiv valu of soial bhavior.

Foundationsin Natural History

Undrstandingwhy spis hibit th pattrns of soial havior thy do r-

quisdrawing idas from volution, ology, and thology. During th past
half ntury thes idas, ah spun fom diffrnt modS of thought, hav
bnwovn into ohrnt ut lngthningonptual loth. Fist, hav_
ioristshad to lrify how thy xplaind th appranof a havior. From
thmundan to th fasinating' th havior of animals indus th urious
...!hyar
to ask, thy having in this way?'' In sminal artil Tinrgn
(1963)proposd that thr wr four ways of answring ths ..why'' qrrs-
tions.Thr wr stndard far that fousdon issuslf immdizrtausation,

27
28 Soil Behuiord Soiobioklg

ontognti dvlopmnt' nd volutionary history. What h addd was a baus saf nst sits.
fourth that foused on unravling a havior's funtion y idntifying how it safst trs. \With opp
augmntd an ator's survival or rprodutiv valu. This mphasis on x- polygyny and right m
plaining havio in trms of sltivnfitsto individuals hlpd mov - tis of many othr gro
plnations of soial funtions away from what is good for groups to what is parativ lns (Lak 196
good for th individuals that ompos ths groups (\/illiams1966). Tinr- mthods of atgorizin
gn's insights idntifida onfusion that oftn prvadsth study of havior, among spis hav
namly, that ther ar roadly two typs of planations:proimat and ul. omparativ mthod t
timat. Idntifying triggrs,or rlasrs,of a havior or th timing of its ap. among havioral out
paran during ontogny yilds proximat planations austhy fous road-rush undrsta
on th mahinry within animls tht gnrats th havior. Idntifying yond what ritis (.g
how a bhavior altrs an individual's fitnss or th way in whih historial gain a mhanisti un
antdntsdtrmin what matrial onstituts th rprtoir of a spis why partiular soial r
yilds ultimat xplanations;thy fous on th volutionary roots and ons- irumstans was n
quns of havior. of ours' this dihotomy is lurry sin mhanisti x p l a i n i n d i v i d u [d i
pattrns hav funtional onsquns,and as a rsult thy too volv. But
for th most part' kping th lvl of planationslar whn aounting for
why animals havth way thy do was a major rkthrough.If this pra- mrging Thorti
ti had n aptdwidly, many of th argumntsthat hav plagud th
study of soial havir, spially as it rlats to humans, ould hav n Two diffrnt thoris
avoidd. ogists. Why animals sh
Sond, sarhingfor a havior'sfuntion, or adaptiv valu, nssitats thmslvs,is a qustio
assssingits rlationship to th nvironmnt.Long for Darwin, iologists was puzzld aout wh;
.W.hat
intrprtd morphologial traits in rlation to th nvironmnt. Dar. through ourrd wh
win did Was dmonstratthat suh adaptationsould aris without a rator' Until th mid-1960s th
thus omplling arly naturalists to sarh for nvironmntal prssurs tht dwrds (7962) am
ould serv as sltiv,or shaping' gnts.on of th arlistand most om- mans of rgulating po
plling studis that dmonstratdthat the adaptiv valu of ntir soial sys- opratd. Williams (19
tms ould xplaind y diffrns in ology Was rook's (1964a, ial intrations augm
1964) study of wavrirds.By xamining th foraging, nsting' grouping, Hn sltion atd
and mating havior of ovr 100 spis of Afrian Wavrs that oupid individuals, thn aou
haitats ranging from tropial forststo grasslandsand svannas,rook was straly hlp ripints
al to dmonstratthat ologial fators assoiatdwith th distriution of prolm. Hamilton's in
food and prdators Wr rsponsilfor th road soial diffrnsthat th only dirtly through
spisxhiitd. Thos spisthat livd in forststndd to solitary' in- rlativs. Th ominat
stivorous,trritorial, draly olord' and monogamous. Crook argud that clusiue fitess and provl
ths traits ovarid baus insts wr hard to find. Sin fding young vn stril asts of ins
rquird frqunt visits to th nst y fathrs and mothrs, monogamy and prodution of rlativs
pairwise dfnse of aras whr insts would rppar ld to monogamy fiint,'' and Hamilton
and trritoriality. rook also suggstdthat uilding rypti nstswithin tr- ration of te kiship
ritoris and th volution of dra oloration would rdu prdation. on- drivd y a rlativ m
vrsely, thos spis that livd in savannas and othr opn landsaps rlativ dvalued y th
tndd to soial, trsof sds'olonial, and polygynous' with mals ov- for undrstanding th
rd in rightly olord fathrs.Sin sds oftn ar lumpd, Crook - and has n supportd
livd that nfitsssoiatdwith groups ing al to sarh fo sdsovr That nfits and ot
wid aras ld to th volution of soiality and that llniality mrgd for th dvlopmnt of .
 Soil Beuior d Soio|iolog 29
.!7hat aussaf nst sits wr rar, induing mny pairs to nst togthr in th
r history. h addd was a
lr's funtion y idntifying how it safsttrs. !ith opportunities for ukoldry prvalnt, rook argued that
tiv valu. This mphasls on x- polygynyand right mal oloration wr also favord. Over th yars soi-
its to individulshlpd lnov - tisof many othr groups of animals hav bn analzd by us of this om.
hat is good for grouPs to what is p a r t i v l n s ( L a k | 9 6 8 i l a r n a n 1 9 7 4 i K r u u k 1 9 7 5 ; F r i k 1 9 7 5 ) .A l t h o u g h
r groups(Williams 1966). Tin- mthods of atgorizing havior and lassifying volutionary rlationships
n prv.rdsth study of havio' among spis hav n rfind y ovrlaying havio on phylognis, th
[ plrrtitlns: proimat and ul- omparativmthod y itslf, with its rlian on statistial orrlations
hvioror th timing of its zlp_ among havioral outoms nd eologial fors, ould only provid a
planationsausthy fous road-rushundrstandingof th volution of animal soitis.To mov -
nratsth havior. Idntifying yond what ritis (.g.,Lwontin 1979) |aimd wr ..just.sostoris'' and
ss or th way in whih historial gain a mhanisti undrstanding of ologial ausation, ttr gsp of
titutsth rprtoir of a spis why partiular soil rlationships among individuals dvlopd in spifi
t h r ' o l u t i o n e r ryo o t s a n d o n s _ irumstanswas ndd, and this rquird th dvlopmnt of thoris to
ltom is lurry sin mhanisti plainindividual dision mking.
d as a rsult th too volv. But
. n t i o n s l rw h n o u n t i n gf o r
Lmajor rakthror'rgh. If this pra- mrgingThortial Framworks
argumntsthat hav plagud th
:latsto humans, ould hav n Two diffrnt thoris mrgd, on from soial thorists and on from ol-
ogists.Why animls should hlp others, oftn t th xpns of rproduing
i o n ' o r a d a p t i v v l u ,n s s i t t s thmslvs' is a qustion that ppld biologists for nturis.vn Darwin
nt. Long forDrwin, iologists was puzzld aout why so mny soial insts wr stril. A major rak-
ln to th nvironmnt. What Dar- throughourrd whn Hamilton (1,964)proposd th ida of hi sltio.
rionsould aiswithout a rator' Until th mid-1960s thr was onfusion out th units of sltion.Wynn-
r for nvironmntlprssurs that dwards (1962) hampiond th viw that soial havior volvd as a
. on of th arlistand most om- mansof rgulating populations; hn goup havior ws whre sltion
adaptivvalu of lltir soial sys- optd.\(illiams (1966) ritiizd this viw y dmonstratingthat most so-
in ologwas Crook's (1964a, ial intrationsaugmntd th fitnss of individuls that ngagd in thm.
n g t h f o r g i n ,l 1 s t i n gg, r o u p i n g , Hnsltionatd on individuals. Br'rtif bnfitsof bing soial aru to
of AIrin wavrs that oupid individuals' thn aounting for th volution of altruisti ats that dmon-
asslandsand savannas,rook was straly hlp ripints ut aru no dirt nfits for altruists boms a
i sslitd with th distribution of prolm.amilton's insight was that individuals an pass on thir gnsno
h road soial diffrnsth.1tth only dirtly thr<lughoffsprirrg ut als ir"rdirtlyvia th offspring of los
in forststndd to b solitary, in- rltivs.Th ornination of ths two nefitsprodus an individual's ln-
d monogamous.rook argud tht Iusiufitess and provids a fitnss aounting for th volution of altruism.
: hard to find. Sin fding young vn stril asts of insts an volv if thy an suffiintly nhan th r-
:hrsilnd mothrs, monogamy nd produtionof rltivs.But th dvil is in th dtails of what onstituts,.suf-
would rappar ld to monogamy fiint,'' and Hamilton providd a alulus for dtrmining this with th
Latulldingrypti nsts within tr- rationof the kiship oeffiit. His rul that th nt rprodutiv nfit
tion woLlld rdu pdation. on- drivdby a rlativ must gratr than th ost inurd y th altruisti
anns and othr opn larrdsaps rlativdvalud by th riprol of rh kinship offiintprovidd the ky
al, and polygynous,with mals v. for undrstanding th volution of th rnost fundamntal soial haviors
ds oftn ar lumpd, rook b- andhs n supportd y many studis on insts and vrtrats.
s ingabl to sarlrfor sds ovr That nfitsaIrd osts ar fftd y kinship also ratsth onditions
ality and that <llor.rility mrgd forth dvloDmntof onflit within familis and twnth ss (Trivrs
240 Soil Bbuior ]Sociobiolog

1972, 19741Trivrs and Har I976). Sin parnts ar rlatd qually to all
thir offspring, ut offsping ar mor losly rlatd to tlrmslvs than to providd th lns for
thir silings, onflit ovr ptdinvstmnty parnts will oftn mrg. nting options. And v
In gnral, offspring will attmpt to prvnt parnts from qually sprading mining whih s th.
rsoursamong all thir offspring. Howvr, th rltionship bven parntal volutinary history a
invstmnt and sual omptition nray hav n th most profound tn. Whil th soial th
sion of kinship thory for shaping th study of soial volution. Armd with part of th polygyny-t
th rsults of Batman,s(1948) study of fruit flis that showd that rprodu- rang of ll1aring svstl
tion y mals and fmals was limitd by diffrnt fators, Trivrs proposd widr rang of o[
that whn on s invsts mor in th raring of ffspring than th othr, potntial for polygyn
mmbrs of th lattr will ompt for mmrs of th formr. In ssn,th tion of th limiting s
sx that invstsmor in raring omsa limiting rsour'and th s with find it diffiult to ontl
..ss'' in groups, or rsour
rsours should invst hvily in mating tivitis, inludirrgfind.
ing and guarding mmrs of th panting sx. grgat without inurr
Th sond thory mrgd 1rom ology. Pionring work y Orians, as in th N4aynard Sm
.Willson mals; if fmals wr
Vrnr' and showd that th mating systmof rd-wingdlakirds
dpendd on how ologial onditions afftd th ss in diffrnt ways for polygyny ould nol
(Orians 1969). Thy slrowd that within on population oth polgyny and
monogamy ould oist, and that th lravior of fmals indud this
mating-systm variation. Thy argud that on landsaps with rsour mrgnof a S
paths that varid in quality, mals on those of th highst quality ould d-
fnd nough rsoursto suPport th luths of m:rny fmals. on thos of Th lmntsof a synt
lowr quality, howvr, trritorial mals would luky if thy w :rlto h a m l 9 8 6 ) . F m a l sr v
.Wht aus thir soial rl
sunough rsoursto support th offspring rfon tmal. mad
thir modl uniqu was that it dpndd on assssmnts y frrrals-th dmands of th physi
polygyny thrshold-to dtrmin whthr mating polygynously or monoga- sminal rviw, ths
mously was in a fmal'sst volutionary intrst' and prdation prssu
h undrlying prmis that fitnss dpndd orr what othrs wr doing pndd mostly on solv
st th stag for aynad Smith's (|977) insight tlrat th volution of th saf sites for thmslv
fou major mating systms is a gm in whih what is st for on sx d- mts. oniy aftr ss
pnds on what is st for th othr. Maynard Smith proposd tlrat whthr lndsps wr dtr
On s should invst in aring urrnt offspring should dpnd on how fo mting opportunit
likly th othr s Was to ontinu raring thos offspring. Sin parnts' in- ations y rdingma
trsts ar similar ut not always idntial, Maynirrd Smith showd that hirvior ultimatly ld
thr was a pair of strtgis' on for th ml and on tr th frrral,tht, y suadults and dv
whn prformd togthr, would lad to an volutionarily stl stratgy g n a t i o n s l s o a r i
(SS). Suh a stratgywould our only if it did not pay ithr parnt to d- itis (Rubnstin 198
part from his or hr stratgy.Thus it was possil for iparntal ar, uni- Th synthti mod]
parntal ar proffrd y ith s' or no parntal ar at all to an SS wr bundnt, sp
undr spifi ologil onditions. IWhatth nvionmnt dtrminsis th would low nough t
tnt to whih pantalar is nddto rr offspring. If only on parnt is aging or antiprdator
ndd,thn the liklihood of itlrpartnr dsrtingdpndson th asof and th groups tht fo
finding an additinal mat. For fmals in partir'rlar' this liklihood is also dfndd y singl ma
afftd y th xtnt to whih prnting rdus hr frtility in futur rpro- sult. Flowvr, if rsou
dutiv pisods.Fo th first tim soial asptsof mating havior ould tion among fmales rr
haatrizd nd tid to srlifi sltiv fors. Suzrl asvmrtis would vary, nd fmal
than dfnding unst
 Socil Behuior d Sociobiolog 241
partsar rrlatd qually to all
providd th lns for idntifying how rsours and prdation afftd par-
sly rlatd t thmslvs tharr to
nting options. And whn only on prnt ws rquird for raring, dtr-
nnt parntswill oftn mrg.
mining whih s tha should dpndd on trad-offs shapd y oth
1t parntsfrom qually sprading
volutionary history and rrnt ologial irumstans.
:, th rlationshiptwnparntal
!7hil th soial thorists Wr panding th havioral dision-making
v n th most profound xtn-
part of th polygyny-thrshold modl to aount for th volution of th full
y of soial volution. Armd with
rangof mating systms' ologists wr panding th modl to aount for
rit flisthat showd that rprodu-
widr rang of ologial dtminnts.For mlen nd Orling (1977), the
Jiffrntfators,Trivrs proposd
potnti:rlfr polygyny dpndd on oth th spatial and tmporal distriu-
aring of offspring than th othr'
tion of th limiting sx. If fmals wr widly sattrd' thn mals would
rsof th fornrr.In ssn,th
find it diffiult to ontrol mor than on fmal. Altrnativly, if tmalslivd
limitingrsur,and th sx with
in groups, or rsours wr lumpd so that fmals ould tmporaril ag-
:l mating ativitis'inluding find.
ggatwithout inurring muh omptition, thn polygyny ould rsult. But
g s.
as in th aynd Smith modl' muh would dpnd on th havior of f-
ogy. Pioning work y orians'
mals; if fmals wr rptiv mostly at th sam tim, thn th potntial
lg systlof rd-wingdlakids
for polygyny ould not ralizd.
fftdth ss in diffrnt ways
ln population oth polygyny and
havior f fmls indud this
hat on landsaps with rsour
mrgnof a SynthtiThory
os of th highstquality ould d- .!Vrang_
Th lmntsof synthti thory wr rystallizing (Rubnstin nd
hsof many fmals.on thos of
ham 1986). Fmals wr viwd as th driving for of soial volution -
vould luky if thy w al to
:fspringof on fmal. What mad austhir soial rltionships rfltd thir bst rsPons to mting th
dmarrdsof th physial nvironmnt,and, as Aland r (|974) argud in his
l on assssmntsby fmals-th
sminal rviw, ths largly fll into two atgoris: rsour distriution
r matingpolygynously or monoga-
and prdation Prssur. Maimizing rprodutiv suess y fmals d-
r intrst.
pnddmostly on solving prolms posd y aquiring rsours or finding
lndd on wht othrs wr doing
saf sits for thmslvs or thir young rathr than on finding or suring
) insight that th volution of th
mats. only ftr assoiatiorrsamong fmls and thir distribuions on
whih what is bst for on sx d-
landsapswr detrnrinddid mal stratgisassoiatd with ompting
nard Smith proposd that whthr
for mating opportunitis om into pIay' Susquntrsponssy fmals to
offspring shold dpnd on how
tionsby brdingmls and, in turn, rsponssy rdingmals to fmal
g thos offsprirrg.Sin parnts' in.
bhavioultimatly ld to an volutionarily stal mating systm.But ations
ial, aynard Smith sl.lowd that
by suadultsand dvlpmnt of rlationships among mmrs of diffrnt
mal and on for th fmal. that.
gnratinsalso am into play as soial strutursdvlopd in omplx so-
an volutinarily stl stratgy
itis(Rubnstin1986, 1994).
f it did not pay ithr pant to d-
Th synthti modl prditd that in nvironmnts in whih rsolJrs
s possilfor biparntal ar, uni-
wr abundant, spially whn thy wr vnly distriutd, omptition
ro prntal areat all to an SS
would low nough to prmir fmlsto aggrgat.If suffiintlylrge for-
t th nvironmntdtrmins is th
agingr antiprdator nfits ould drivd y ths aggrgating fmals,
r a ro f f s p r i n g I. f o n l y o n p a r n t i s
and th ]roupsthat formd we not too larg, thn ths groups ould
rr dstingdpndson th s of
dfnddy singl nrals, and so aild harrn_dfns polygvny would r-
n partiular' this liklihood is also
sult.Howvr, if rsursWr mor pathily distriutd, so that ompti-
r d u sh r f r t i l i t yi n f u t u r p r o -
tion among fmals was priodially intnsifid, thn fmal group sizs
lsptsf mtirrghavior ould
.tiv fos. Sxual asymmtis would vry, arrd fmal assoiationswould om mor tratrsitry.Rathr
than dfnding unstal groups of fmls, mals instad would attmpt to
242 Socil Behuior nd Soiobiolog

dfnd rsour paths sought y fmals. In thse systms of rsour- m an aility to su

dfns polygyny th most al mals would dfnd th st paths and thus it was pathily distriu
gain ass to th largst numr of fmals for th longst priods of tim. If rduk (Redu sp
rsours wr distriutd not only pathily ut in paths that wr larg, th st paths that f
wr widly sparatd' and flutuatd sasonally in aundan, thn omp- distriutd, whih oft
tition among fmals would low. This would failitat th formation of as land (Turotrgs
larg groups providd that fmals ould rang widly and follow th shifting vn th lowst-qual
loations of paks in food aundan. als would thus ford to ithr spis viwd lrg <
follow ths larg groups and ompt for' and thn tnd' on rprodutiv sour omptition wa
.!7
fmal at a tim (tllndering polg) or position thmslvs at th intrs. with many fmals.
tion of fmal migratory routs and wait for fmals to visit tem (lekking of a small sugroup o
polgfl.In ithr as intns mal-mal omptition would gnrata Systms dvlopd.
mating systm asd on mal-dominan polygyny, and in th lattr as f. Th sam sorts of
mals would affordd th quisit opportunity of simultanously om- and havioral disr
.$hnvr othr taxa. A rif sur
paring many mals for hoosing with whom to mat. rsours
wr sparsly ut somwhat vnly distriutd, high lvls of omptition p t n s .F o t h q u i
would prvnt fmals from forming groups. As a rsult, individual fmals als hors (qus
would dfnd individual trritoris, thus nsuring a rgular supply of a r- diffrnt rprodutiv t
nwing rsour. Sin solitary individuals who wr sarhing for mmrs dfnd suh groups, al
of th opposit s would fa hightndprdation risk' pairs would ftn ar widly disprsd,a
shr trritoris,and monogamy would rsult. ass (qus hemioos
Until mpirial studison antlop (Jarman 1974) and ats (Bradbury and oli onstraints fror l
Vhrnamp 1977a,1977) showd how ology atually shapd fmal dis- toris along travlingrr
triutions' many of th undrlying assumptions f this modl wr supportd points (Runstin 19E
only y andotal orrlations.Jarman,s lassi study of Afrian antlopsil- In flids, fmals rl
lustrats how nvironmntal fors intratd with physiologial onstraints dnsly woodd au
to shap th mating systm of spis. By shwing how ody siz afftd and onsumd for
th ways in whih diffrnt spis privd and thn rspondd to th dis- whr oth pry and r
triution and aundan of forag and prdators on grasslands, Jarman tions of fmals form l
showd why partiular soial systms inrasd survival and rprodutiv vourd (Pakr \986)'
prospts for partiular spis.H argud that th smallst-odidspis, thr is prssur for mi
suh as did-dik (doqu hirhii)' duikrs (phlophs sPP., Sluicpr mals. Thus th only h
spp.), suni (Neotrgus moshtus), and klipspringers (reotrgL|s orotr- lo). |n this plntiful l
gs), quird limitd amounts of high-quality vgtation. But aus of tary aus it an saf
thir small siz, suh food itms oftn appard to widly sattrd. High For anids, monoga
lvls of omptition and intnsifid risks of prdation mad trritoiality foes (Vulpes spp.) sor
and monogamy appar th st stratgis.Pairs gnrally livd in wodd or dogs (Lo pits) ;
shru-rih aras whr moistur nald vgtation to grow and rnw itslf (Mohlman t986). T
wll into th dry sason. By signaling trritorial ownrship via snt rathr to hlp nourish|rt
than y mans of sound or visual disply, thssmall-odidspisrdud verak jakl (is l
th hans that any of a larg numr of arnivors would pry upon thm. trs that annot thI
As spis inrasd in ody siz' oth physiologially dtrmind ditary smallst spis, howt
nds and th way aptabl forag bam distributd on th landsap small for a fmal; thr
hangd. Sin rypsis am an untnal antipdator stratgy for largr yars whn pry rsou
and mor widly ranging spis, forming groups bam th st strtgy thir soon-to--fun
for suh spis to lowr prdation risk. Fortunatly, with largr siz also to disprs. Thy thrf

ls. In ths systms of rsour-
ld dfndth st paths and thus
:s for th longst priods of tim. If
.ily ut i paths that wr lrg,
sonallyin abundan' thn omp-
; would failitat th formation of
angwidly and follow th shifting
als would thus ford to ithr
'r, and thn tnd, on rprodutiv
position thmslvs at th intrs-
- for fmals to visit tem (lk.king
ral omptition would gnrat a
polygyny, nd in th lattr as f-
ppotunityof simultanously om-
.s7hnvr
whom to mat. rsous
riutd,high lvls of omptition
ups. As a rsult, individual fmals
nsuringa rgular supply of a r*
ls who wr srhing for mmrs
d prdationrisk, pairs would oftn
sult.
"nan 1974) and ts (Bradury and
ologyatully shapd fmal dis-
ptions of this modl wr supportd
lassistudy of Afian ntlops il-
atdwith physiologial onstraints
By showing how ody siz afftd
:ivd and thn rspondd to th dis-
J prdators or-rgrsslands, Jarman
nrasd survival and rprodutiv
rdtht th smallst-odidspis,
<rs(phllpbus spp., Slvipr
klipspringrs (reotrgus oreotr-
-quality vgtation. But baus of
'padto widly sattrd. High
;ks of prdation mad trritoriality
s. airs gnrallylivd in wodd or
vgtationto grow and rnw itslf
:rritorial ownrship via snt rthr
', thssmall-bodidspisdud vrak 1aka|(is mesomeld, th hlprs ar young from prvious lit.
f rnivorswould pry upon thm.
r physiologiallydtrmind ditary
amdistriutd on th landsap
lbl antiprdator stratgy for largr
ng groups bam th st stratgy
l. Fortunatly,with largr siz also
Soil Bahuior d Soioliolog 24
am an aility to susiston mor aundant, lowr-quality vgtation..i7hn
it was pathily distriutd' as Was th s for impala (Apcrlsmelmpus),
duk(Redu spp.), and sm gzlls (Gzll spp.), mals dfndd
th st paths tht fmls prfrrd. !hn th vgtation was mor vnly
distriutd,whih oftn rsultdsimply from th fat that largr spissuh
as land (Turotrgus orv) and ap uffal o (Scros cffer) ould us
Vn th lowst-quality items, largr groups formd. Baus th largst
spisviwd lrg ontinuous swards of a landsap as aptal, r-
souomptition was virtually liminatd,nd many mals ould assoiat
rvith many fnrals.With suh high lvls of rnale_nralomptition' dfns
of a smal] sugr'rpof frals anrimpossil, and dominan dfns
sstmsdvlopd.
Th sam sorts of onntions btwn hanging ologial irumstans
and havioral disitln making shad th typs of soiality xhiitd in
oth taa. A rif survy of mammals illustrats som of th mor gnral
pattrns.For th quids, th los assoiation twn food and watr
n-
als hors (quus bllus) and plains zera (quus burhelli) fmals of
diffrntrprodutivstatsto assoiatprmanntly.Thus mals ar al to
dfnd suh groups, nd so-alld harms form. .Whn ths two rsours
ar widly disprsd, as fo Gvy's zra (quus grui) and th Asiati wild
ass (quus hmioous), fmals of diffrnt stats ar prludd by mta.
oli onstraintsfrom foraging togthr.As a sult,mls ompt for trri_
torisalong trvling routs that tak fmalsfrom fding aras to watring
points (Runstin1986, 1994\'
In flids, fmls rmain sparatdwhn food is sar and haitats ar
dnslywoodd ausin suh irumstansindividul pry an b aught
and nsumd for omptitos an intrvn.In mor opn haitats and
whr oth pry and omptitors ar lrg arrd mulr mor numrous, oali-
tions of fmals frm to help hold on to kills until thy ar ompltly d-
vourd (Pakr 19s6). If ths fmal olitions ar thmslvs larg, thn
thris prssurfor mals to ggrgtto ontrol rprodutiv assto f-
mals.Thus th only higly soial flid is th svanna-living|ton (Pnthr
Ieo)' In this plntiful landsap th lopari (Pnthr prdus) rmains soli-
tary auseit an safly ah its lag pry in trs.
For anids, monogamy is th rul. But variations do our: small-odied
foes (Vulps spp.) somtims hiit polygyny, and larg_odidhunting
dogs (Lrl pitus) arrd tirnr wlvs (is lupus) dvlop polyanlry
(ohlman 19s6). Typially anids nd hlp from norrrding
foagrs
to hlp nurish latating mothrs. Fr th midsizd iakals. suh as th sil-
trs tht annot thmslvs find sussful rding loations. For th
smallst spis, howvr, prpartum invstmnt in young is rlativly
small for a fmal; thus hr mat provids all th hlp that is ndd. In
yarswhn pry rsours ar vry high, omptition twn mothrs
and
thirsoon-to--funddaughtrs is low, and thir dughtrs ar not ford
to disprs.Thy thrtrprovid nighorirrg mls with additionl mating
244 Soil Behuior d Soiobiolog

opportunitis, and polygyny rsults. For th largst-odid spis, how- or food rsours oftt
vr, fmal prpartum invstmnt is vry high, and thy nd all th hlp p r o d u d d i r i n sl i
thy an gt. To nlist th support of othr adult mals that must hunt o- failing to prdu an
oprativly to aptur larg pry' a dominant fmal not only kills th off- forming whn ondit
spring of othr fmals in th group ut also mats with thir mats so that But whn thy form'
ths mals hav as if thy ar th sirs of th dominant fmal's young. as many studis hav
Hn, dpnding on siz-dtrmindmtaoli investmntsand nds' so- If family omposit
ial systms of anids an vary from polygyny to polyandry, and thy som- hang. Baus los
tims us th srvis of juvnil or dult nonrdrs to hlp rar spring rarly attmpt
offspring. sx. Thus omptitior
As ths xampls illustrat, th omparativ mthod nfitd trmn- nitis rmains low. If,
dously y th dvlopmnt of soioologial thory that mphasizd th thn from the prsp
importan of fmals and thir rlationship to th nvironmnt in strutur- will likly produ .
ing soitis.But asptsof kinship and dmography also om into play. As mat and rmains in
amilton's rul illustrats, th strongr th dgr of rlatdnss among offspring of this pir,
rlativs, th mor likly altruists ar to nhan th rprodutiv opportu- dgr to whih th r
nitis of kin whil inurring osts assoiatd with diminishd prsonal r- ratd will dpnd on
prodution. Thus in th prvious xampls, it is not surprising to find that ship twn th tw
th oalitions that form among lionssswhn thy ar protting kills and prodution mong l
among male lions whn thy ar dfnding mating opportunitis with f- valu of 1 whn th r
mals ar formd most oftn among full silings (Pakr 1986); nor is it sur- whn rding is om
prising that th hlprs jakal adults kp to rear additional offspring are podution ours wil
thmslvsth full silings of th offspring ing raisd (Mohlman 1986). by dominants if suo
In gnral, strong kinship lowrs th thrshold for th apparan of altruis- nats if thy lav; (3
ti and ooprativ havior and may sustantially afft th osts of living and (4) th magnitud
in groups. rdrs. $hn oppo
Th strutur of jakal soitis not only highlights th importan of kin high, th dominan d
sltion ut also illustratesth fat that mny soitisonsist of adults of th nfits of having
diffrnt gnrations' som of whih ar for offspring that ar not nssar. breder will slt
ily thir own. any insts in th ordrs Hymnptr and Isoptra and (Rv and Ratniks 1
aout " of birds and mammals show srrh systms omposd of oopra- options lswhr and
tiv brdrs (mln 1997)' Up to this point th mrging synthti thory of damag on th sta
soiality has largly tratd all adult fmals and mals as indpndntdi- .!7rg
1992), Groov
sion makrs; rsponding t ologial onditions shapd th st rspons (Faaorg et a|. 1995),
for fmals, whih in turn onstraind th st rsponss of mals. But in 1994) have shown, do
many soitis mature offspring rmain and help parnts rar additional off. likly whn hlprs ar
spring. Ultimatly, undrstanding why individuals forgo prsonal rprodu- kin sltion.
tion to hlp los rlativs rprodu povids insights into how th alan But as might p
btwn dspoti and galitarin soial rlationships, as wll as th dynamis and laving disions t
of familis, is dtrmind y intrationsmong ology, kinship, nd domi. qustiond whthr d
nan (mln 1'995). groups (lutton.Brok
In most spis, offspring lav hom for rding. But in many soial ativ brding soitis
insts, primats, and othr mammals and irds, on sx rtins tis to par- tions asd on kinship :
nts, somtims prmanntly' rating multignrational familis. Gnrally, suh as mutualisms tha
offspring postpon disprsing whn rding opportunitis ar limitd ls- that mrg as mrka
whr. lgial onstraints assoiatdwith limitd rding sits, mats' om into play.

: th lrgst-odidspis' how-
:y high, and thy nd all th hlp
rr adult mals that nrust hunt o.
inant fmi not only kilis th off-
r l s o n r t sw i t h t l r i r m a t s s o t h t
:s of th dominant fmal's young.
:taoliinvstmntsand nds' so-
gyll to polyandry, and thy som-
adult nonbrdrs to hlp rar
partivmthod nfitd trmn-
ogial thry that mphasizd th
; h i pr o t h n v i r o n t n n ti n s t r u t u r -
lmographyalso om into play. As
: th dgr of rlatdnss among
n h a n t h r p r o d u t i v o p p t u -
iatd with dimirrishd prsonal r-
lls, it is not surprising to find that
i whn thy ar protting kills and
ding mating opportunitis with f-
s i l i n g s( P a k rl 9 8 6 ) ;n o r i s i t s u r -
)p to rar additional offspring ar
ing bing raisd (Mohlman 1986).
sholdfo th app;rranof altruis-
tstantiallyafft th osts of living
nly highlights th importan of kin
t many soitisonsist of adults of
for offspringthat ar not nssar-
rs L{ymnoptra and Isoptrzr and
suh systmsomposd of oopra-
int th mging synthti thory of
ralsand mals as indpndnt di-
:onditionsshapd th st rspons
th st rsponssof mals. But in
and hlp parnts rar dditiona| off-
ndividtllsforgo prsonal rprodu'
ovids insights irrto how th bln
.lationships,as wll s th dynamis
; mong ology' kinship, and domi-
forbrding.But in many soial
n d i r d s ,o n s r t a i n st i s t o p a r -
nultignrtional familis. Gnrally,
ldingopportunitis ar limitd ls-
l w i t h l i m i t d r d i n g s i r s ,t n a t s '
So il B ehtlior d So ci lbiollg z+\
or food rsoursoftn mak th option of stying hom and hlping parnts
produadditional silings ttr than disprsing arly and almost rtainly
failing to produ any offspring. Thus marry family groups will unstal,
formingwhn onditions ilre poor and disnding whn onditions good.
But whn thy form, intrgnrationaloopration should ommonpla,
as many studis hav shown.
If family onrposition hangs, howv, thn soial dynamis an also
hang.Bause los inrding oftn has dltriousffts' rtirrdoff.
springrarly attmpt to mat with a prnr or los rlativ of th pposit
s.Thus omptition with th sam-sxdparnt for rprodutiv opportu-
nitisrmains low. If, howvr, a parnt dis and is rplad y a nonrlativ,
thn fom th prsptivof nondisprsingadults, nw mirting opportunitis
will likly produ onflit. Similarly' if a nondispsing adult aquirs
mat and rmains in th group and limitd rsours ar onsuled y th
offspringof this pair, thn omptition wi1l hightnd.In ithr as th
dgrto whih th rprodutiv ativitisof th nondisprsingadult ar tol.
ratdwill dpnd on th stngthof th dominan-suordinrrtionrlation-
ship twn th two. Reproductiue she, desris th distriution of
rprodution among rding adults within familis and an rng from a
valuof l whn th rdrin harg garnrs all rprodution to a valu of 0
whn rding is ompltly egalitarirr.Th dgr to whih sharing ot r-
produtin ours will dpend upon four onditions: (1)th rrfitdrivd
y dominants if suordinats stay; (2) th rprodutiv suss of subordi-
natsif thy lv; (3) th gnti rlatdnssamong potntial ordrs;
and (4) th mgnitud of th domirran rlationshi btwn potntial o-
brdrs. \ff/hn opportunitis for th suordinat to rd lswh a
high, th dominan diffnamong rdrsis small, kinship is low' and
th nfitsof having hlps to th dominant r high, thn th dominant
rdrwill seleted to pay high sti?lg inntiu and p itttiz,
(Rvand Rtniks 1993) to nsur that a hlpr hlps sin l-rlprshav
options lswhr and, aus of narly qual fighting aility, an inflit
damag on th stalishd rdr. As studis on -atrs(mln and
Wrge 1992), Groov-illd anis (Vhrnamp 1978), Galipagos Hawk
(Faaorget a|. |995|, and mongoss (Crl nd \Wasr1991; Kan t al.
1994) ave shown, dominant rdrs making ..onssions'' to hlprs is
likly whn hlprs ar not los kin sine thy gain littl indirt nfitvi
kin sltion.
But as might b xptd, variants to this st of ruls that shp staying
and laving disions of hlprs our. lutton-Brok and ollagus hav
qustiondwhthr dominants hav full ontrol ovr rprodution within
gloups (lutton-Brok t al. 2000). Thir studishv shown that in oopr_
ativ rding soitisof mrkats, lvls of onssions do rrot fit prdi.
tionsbasd on kinship and th nds of dominants' As a rsult, othr ftors'
suhas mutualisms that nhan rprodution for vry group mmr and
that mgas mrkat groups gt lrg (lutton-Bok t l. 2002.), ma
ominto play.
246 Soil Behuior nd Soiobiolog

a sukr dos tt
Additional Soial Glus b-product mutulis
of what a partnr d
As mrkats illustrat, not all soitisnd hld togthr y th glu of kin- Studis on a varit
ship. Rlationships uilt around oopration rathr than altruism or asd Long-trm studis of
on mutualism and riproity an hav just as strong an fft and provid Pakr 1991) and St
th opportunitis for diffrnt typs of soitis to volv. Rlationships ar y adopting individu
uilt upon rpatdintrationsthat an involv omptitivand ooprativ ilarly, strong rlation
lmnts. Manipulating th natur of prsonal rltionships and thos of fantiidal mals or
ompanions should also dtrminth xistnand haratristisof many prdtors providd si
soitis. ing alon.
Trivrs (1972a) Was th first to suggstthat riproity ould play an im-
portant rol in shaping soitis.If individuals Wr to aid a strangr y in-
rasing its survival or futur rprodution at som ost tO slf in trms of Sial Variants al
diminishd rprodution' thn suh havior ould only maintaind in a
population if th ripint wr to vntually rpay th osts inurrd by As th xampls of fa
th altruist. This onpt has n formalizd as th prisoner's dilemm, dnsity and th intn
whr two individuals hav two altrnativs-ooprat or dft. utual trritoris with suffi
oopration, usually y staying tru to ah othr dspit attmpts y au. rsponsibl for favori
thoritis to ajol on Partnr to tufn on th othr, provids a highr r- of hlprs. Thus dm
ward (R) than if thy both dft and ar punishd /P/. If only on partnr sx ratio and ag stru
turns on th othr' hwvr, then th siz of th tmptation (T) nssaryto vitl ats, will shap
rak th partnrs' oopration nals th hat to do st of all. on- spis variation as
vrsly' th partnr who rmains ooprativ to th nd is th sukr and pl, wr ag spifi
rivsth lowst rward (S/.As long as payoffs ar ordrd > R > P > s, thn high rding s
dfting is favord sin it provids th gratst short.trm nfit. If, tionships would gn
howvr, suh situations our frquntly and th nmr of individuals in of motality Wr r
a soity is small, thn pairs of individuals will find thmslvs intrating ommon. Sin som
tim and again. In suh a stting, dfting is no longr th st stratgy. th variation in anid
Rathr, oopration dos bst' and th tati tht oftn provids th grat. uls an hav importa
st long-trm rWard is tit-for.tt (TFT; Alrod and amiltorr 1981)' in Th synthti modt
whih intratants ooprat initially and ontinu to do so until a partnf nvironrnntal fatur
dfts. If dftin ours' thn th othr partnr dfts and ontinus to mals. In many ass1
math th havior of th initial dftor. If th initial dftor again oop- systm of soial org
rats' thn th rspondr is instantly forgiving and ooprats as wll. TFT onditions. In quids
an rsist invasion y a stratgy of pur dftion. But sin it is qually as transitory groups wh
rwarding as a stratgy of always ooprating, TFT an suprsddby populations mov fr<
suh a stratgy via drift. Sin a purly ooprativ stratgy n invadd hang. As was foun
by th slfish stratgy of dftion, th stratgy of tit-for-tat annot truly Stats, whr food an
an SS, ut it oms los. diffring nds an
Th logi of this argumnt suggsts that struturing soitis on rl. aras ar ford to s
proity alon will diffiult. Kinship an always provid a strong foun. spond y dfndingtl
dation sin it rdus th nd of fully rpaying dlayd nfits. By and harmlike soiti
oopating mutually to solv ommon prolms, partnrs may nfitto Soial variation' ho
suh a lrg tnt that th tmptation to dft rmains low nough that Variation in sial str

hld togthry th glu of kin-
t rirthrthan altruism or asd
as stongan fft and provid
:isto volv.Rlationshis r
l v m p t i t i v n d o o p r a t i v
lnal rlationshipsand ths of
n and hartistisof mny
l r t r i p r o i t y u l d p l a a n i m -
lls wr to aid a Strangr y in-
at som ost to slf in trms of
r ould only maintaind in a
lly rpay th osts inurrd y
zd s the prislr's dilmm,
s_ooprat or dft. utual
h othr dspit attmpts y au-
,h othr, pror,ids highr r-
unishd /P). If only on partnr
th tmptation (T) nssary to
hat to do st of all. on-
iv to th nd is th sukr and
ryoffsar odrd T> R > P > s,
gratstshort-trm nfit. If,
nd th numr of individuals in
w i l l f i n d t h m s l v si n t r a t i n g
g is no longr th st stratgy.
'i that oftn prvids th grat-
lrod and Hamiltorr 1981), in
:ol-ltinu to do sl until a prtnr
p a r t l . l d f t s n d o n t i n u s t o
[ h i n i t i ld f o g a i n o o p -
,ing and oopratsas wll. TFT
ftion.But sin it is qually as
ting' TFT an suprsdd y
prativstratgyan invdd
:gyof tit.for-tat annot truly
a t s t r u t u r i n gs o i t i s o n r
r lwys provid ir strong foun.
' rpaying dlayd nfits. By
'olms'partnrs may n{it to
dftrmains low nough that
Socil Bhuior d Soioliolog 247
a s u k r d o s t t r t h a n a d f t o r . . W s t - r h a r d( 1 9 8 9 ) t r m s t h i s
b-produt mutulist us oopration is univrslly st rgardlss
of what a partnr dos.
Studison a varity of spissuggstthat mutualism an shap soiality.
Long-trm sttrdisof lions y Pakr (Pakr and Ruttan 1988; Shl and
Pkr 1'991')and Standr (1992) showd that only y hunting togthr and
by adopting individul rols wr lions liklv to ing down lrg pry. Sim_
ilarly,strong rltionshipsamong fmalsto prott us from attaks y in-
fantiidl mals or to hold on to pry in th fa of othr marauding
prdtorsprovidd simultanous nfitsthat old not obtaind y at-
ing alon.
Soial Variants and Altrnativ Stratgis
As th ampls of fmilis with ooprtiv brdrsshow, high population
dnsity and th intnsifid omptition it ngndrs in th sarh for sr.rital
trritories with suffiint food or haital urrows is th fator ultimatly
rsponsilfor favoring th stalishmntof oalitions nd th rruitmnt
of hlprs.Thus dmographi ftors, suh as popultion dnsity, as wll as
sx ratio and ag stutur that rsult frorn diffrns in phenotyp-spifi
vital rats, will shap pattrns of soial organization and lad to within-
spisvariation as ologial irumstanshang. If mortality, for xm-
pl, wr ag spifiand high for prrprodutivfmals than for mals'
thn high rding sx ratios (mals/fmals)wuld rsult, and mating rla-
tiorrshipswould gnrally om mor polyandrous. If s.spifi pattrns
of mortality wr rvrsd, howvr, thn polygyny would om mor
ommon. Sin so of ths nrortality onrns wr also rsponsil for
th vriation irr arrid soil strutu dsrid arlir' ths mortality shd-
ulsan hv irpotant onsquns.
Th synthti modl of soiality that is mrging shows that a varity of
nvironmntalfturs influn th pattrns of soility xhiitd y ani-
mals' ]n many ass ths pattrs ar flxibl, and spis an vary in th
systmof soial organization thy hiit dpnding upon nvironmntal
onditins. In quids' for ampl, although fmal asss typially liv in
transitorygroups whos mmrshiphangs (Runstein1986,1994) wen
populations mov from arid to msi aras, soial rlationships an also
hang.As was found on an island off th ost of th southastrnUnitd
Stats'whr food and Watr ar aundant and los togthr,fmals with
diffringnds an oals into prmannt groups. als, whih in arid
i - arsar ford to stalish trritoris along routs to and from watr, r-
spond y dfnding ths groups muh lik mals of horss, thir los kin,
and harmlik soitismrg.
Soil variation, howvr' is not limitd to diffrnsamong popultions.
Vrition in soial stratgisan volv within populations sin not all indi-
248 Soil Behuior d Soiobiolog

viduals ar qual; diffrensin ownrship and past prinan afftth

outom of intrations.If individuals irrtratrpatdly,thn th natur of Undrlying N
thir intrations will hang ovr tim so th?1tontsts nd mor quikly and Control to S
paaly, with suordinats sumitting to dominants. In small or modratly
sizd groups, hirarhisform and soial rols om odifid. Although all Dision making
individuals' rgardlssof rnk, nfit whn srious fighting is rdud - parisons of osts
aus oth winnrs and loss bnefit y rduing wastd tinr on outoms tiulr soil v
that r virtually assurd, dominants typially driv th gratstrprodu- prossd nl us
tiv nfits. In tndd fmilis of ooprativ rdrs, dominan plays nt lvl of arrly
an important rol in dtrmining whthr rprodution is mor or lss ithr why indivi
qually apportiond among adult group mmrs. was triggrd.
!0hn dominan is xtrrland kinship is low, rprodutiv options for ining th two t<
adults ar limitd as thy id thir tim prodingalong norml ontogn- soial hvior.
ti trajtorisn rout to rproduing.In suh situations shortutsto rd- on forr of ir
ing ar oftn favored, spially if mortality rats of adults ar siz or ag ganizs havior
spifi and ar gratr for largr and oldr mals. Trading off longvity with physial or soia
low rats of rprodution for opious rprodution ovr a slrort priod an at th right tim,
lad to disrt siz polyrnorphisms-so-alld altrnativ rnal mating rquirs olrdin
stratgis-within ss within popultions. oftn th typial stratgy animals ar rird
adoptd y oldr and larg mals of dfnding harms or rsoursyilds mate' matur ]a
th most rprodutiv gains during a rding sason' ut it oftn inurs a for th transition
high ost as wll. Baus displaying' fighting, attrating th attntion of faturs must
Prdators, and dlaying rprodution whil gowing ar all ostly ativitis, whthr thy at
mals that adopt lss it.tstntanously sussfulbt also lss ostly tatis thn oordirrirtin
an flourish. formation florv. L
Th maintnan of suh altrnativ mating pattrns is ommon among possibl.
many spis of insts, fish' amphibians, irds, and mamnrals. For am. Anothr typ o
pl, in lugill sunfish (Lepomis mrobirus) mals typially dfnd nst ior mrgs fro-
sits whr thy display' attrat fmals' i:rrrdthn ftiliz nd guard th havior viwd
ggs thy lay (Gross and harnov 1980). Sin only th largst mals hav sal thy appar
th aility to dfnd nst sits, thy must dlay rding for ovr svn may assmbl in
yars. As a rsult' smallr and youngr mals hav volvd various ukold- ditions, ut whtt
ing stratgisthat may in ft qually sussfulvolutionarily. In on, and sizs dpnds
th so-alld snak gins rding at two yars of g' Although vry pulsion, attrtio
small, suh snaks ar virtually all tsts and baus of thir inonspiu. ont for th siz
ousnss an div from th surfa just as a mating pair rlasssprm and Using a st of hit
ggs. By xuding larg volums of sprm' som frtiliz a fw ggs. In th omptition and
othr, mals dlay rding for as long as do fmals. By ing th sam lagus (|996\ s
siz and olo as fmals, ths mals join th mating pair and apparntly tndnis to frag
fool th displaying rl ir-rtothinking that h is ourting two fmlesrthr indu ths triln
than just on. As th original pair rlss its gamts' th fmal mimi los nighors. I\
dos too, thus frtilizing som of th fmal's ggs. For ths two stratgis ffts of niglro
to qually sussful altrnativS' th osts and nfits of ah must dimnsions' ut
vary invrsly with th frquny of individuals that adopt ths tatis.Al. align with th ori
though thy somtims do, as in th as of th lugill sunfish, in othr in individual ha
spis thy do not. and floks of irdr

) and pst prin an afft th
rtrpatdly,thn th natur of
that ontstsnd mor quikly and
l dominants.In small or modratly
olsomodifid. Although all
hn srious fighting is rdud -
rduing wrstdtim on outoms
ially driv th gratstrprodu-
prativrdrs,dominan plays
,hr rprodution is mor or lss
nmrs.
rip is low' rprdutiv optiorrs for
prodingalng normal ontogn-
r suh situationsshortuts to rd-
r l i t y r a t so f d u l t s a r s i z o r a g
r mals. Trading off longvity with
prodution ovr a short priod an
so-alld altnativ mal mating
rtions. ofln th typial stratgy
fndingharms or rsours yilds
ding sason' ut it oftn inurs a
ighting, ttating th attntion of
ril growing ar all ostly ativitiS,
rssfr'rlut lso lss ostl tatis
mating pattrns is ommon among
l s , i r d s ,a n d m a m m a l s . F o r x a m -
,hirus)mals typially dfnd nst
:s, nd thn frtiliz and grrard th
l). Sin only th largst ma]s hav
rust dlay brding for ovr svn
nals hav volvd various ukold-
ly sussfulvolutionarily. In on,
: two yrs of ag. Although vry
s and ausof thir inonspiu-
ls a mating pair rlases sprm and
mt som fertl|ize a fw ggs' In th
3 as do fmals. By bing th sam
oin th rating pair nd apparntly
rt h is ourting two females rathr
lssits gamts' th fmal mimi
m a l . s g g s .F o t h s t w o s t r t g i s
h osts and nfits of ah must
ividuals that adopt ths tatis. Al-
as of th lugill sunfish, in othr
SlilBhuior d S<lciobiolog 249
Undrlyinghanisms:From Hormonal
ontrol t Slf-organization
Disionrnking undrlis th mgnof arrimal soitis.Although om-
parisonstrf osts and bnfitsprovid th ultimat planation of whn par.
tiul soial variations fvord y sltion, th wy irrformation is
prossdand usd in making soia1disins rquirs fousing on a diffr-
nt lvlof anerlysis.In th pst, studisof siality fousd on only on sa|:
ithrwhy individuls intratdth wy thy did or on how suh havior
was triggrd.Mor rntly' studishav gun moving aross sals,om-
ining th two to provid llor intgrtdundrstnding of th auss of
soial havior.
on form of intgration amins how informtiorr withir-rorganisms or-
ganizshavilr.Sin nluh of havior is onditioned on fatrrrsof th
physial or soial trvironmnt'nsuring that appropriat haviors appar
at th right tim, in th right Ontt'and in th prsnof th right
artnrs
rquirsoordination. Hormons ar ordinators par xlln.\/hntwo
animis a rady t() ll1t' thy mr-rstoth in th right mod; whn thy
mat,matur gamtsmr"rst rdy and waiting; and whn th ag o siz
for th trnsition frrmsuadult to rnatur adult is rahd,many phenotypi
fatursmrrst rorganizd.\ihih sustanesontrol what prosssand
whthr thy at rr th rain or oth organs an vary' ut initiating and
thnoordinating and intgratirrgmany asptsof soil bhavior rqui in-
formation flow. Ultimatlv th prosssinvolvd onstrain what havior is
possil.
Arrothtyp of irrtgrationinvolvs undrstanding how olltiv hav-
ior mrgsfrom th ations of individuals (Lvin |999). Oftn pattrns ot
bhaviorviwd at on sl apper imposd fom aov' whras at nothr
saithy appar to mrg from low as if slf.organized.Thus animals
may assrnblin partiuizrrhitats ausof prvaiiirrgnvirontnntalon-
ditions,ut whthr or not thy form tightly knit groups of partiular shaps
and sizs dpds on how thy intrat with ah othr. Simpl ruls of r-
pulsion,atttion, and alignmnt ssoiatdwith intrtion zons an a-
ount for th siz, shap, omposition, and dirtion of group movmnts.
Using a st of hirirrhil dision rules tht aland repulsion to avoid
omptition and attration to rdu risks of prdation, Guron and ol-
lagus(I996) showd that groups displayd a wid varity of shaps and
tndnisto fragmnt and slf-sort bv phnotyp. All that ws rquird to
indu ths transitions was information aout th spd and position of
losenighors.Mrrh dpnddon th istnof rrutralzon whr th
efftsof nighors did not altr intrinsi tndnis.Similar models in thr
dimnsions,ut whr th psnof a nutrl zon allowd individuals to
alignwith th orintirtionof a nrst nighor, rvald how small hangs
in individul havior ould dramatially hang th shap of shools of fish
and floks of irds (ouzin nd Kraus 2003). Dpnding on tlr strngth of
250 Soil Bhuior d Sociobillog

orintation, groups ould mov from disorganizd swarms to highly polar. is no on animal
izd onformations. In fat' strngth of alignmnt playd a ritial rol in un- grs y oopra
drstanding ladrship, information transfr' and onsnsus dision making thos of most t}
within groups vn whn informd individuals did not know th quality of and mor variti
thir information or whthr thy wr in th majority or minority (ouzin diffus. But this .
t al. 2005). not apply to thos
nssarily invali<
on of th mos
From Animals to Popl an plain a n
rothr's phnorr
Humans ar animals, so it sms slf-vidntthat volutionary prosss posd to thir ov
should shap human soiality. First prinipls, spiallythos that undrli strat8y appars
th synthti modl of soiality in nonhuman animals, hav stimulatd and qual, a parnt sh
organizd th study of human soial havior. Not surprisingly, onntions as with a sibling's
among partiular faturs of soial volution and partiular aspts of hu- spially tru in
man natur hav n idntifid. Nowhr is this mor apparnt thn in th (Flinn 1981; Iron
ara of mat hoi. In rodnts gns assoiatd with th majr histoompat- rih' and as a rst
ibility ompl (MH) afft ll rognition and immun funtion. During rais their hildr
strus fmals assoiat with mals that diffr gntially from thmslvs' philandring is hi
whras whn thy ar prgnnt, thy assoiat with fmals that ar gn- is ommon' and l
tially similar. Assssmntis via odor and suggststhat MH-speifi assoi- vn if offspring i
ations ar volutionarily adaptiv. Divrsifying th gen pool of offspring sister's hildrn t
should improv thir immun funtion, and assoiating with los kin during ship thory, mn z
prgnany should rdu osts and inras nfits assoiatd with soiality th st of a ad j
.sfdkind
(Pnn and Potts 1999). Whn and ollagus(1995) prformd a Applying st.t
study to s if human fmals showd similar sual prfrns, h found man soial hav
that womn in th frtil phas of thir yl idntifid th odor of T-shirts biology sought to
from mals with whom thy shard H gnsas lss ..pleasant''than thos havior of isti
of mals with whom thy shard fwr gns. Apparntly th sam undrly- hop was sltiv
ing fors and mhanisms mployd y animals may oprating in hu- that had opratd
mans whn it oms to hoosing mats. But y no mns do ths rsults likly to tru' i
dmonstrat that prosss that shap th soiosxual havior of animals for idntifying h
will nssarily shap th havior of humans in similar ways. Unfortunatly, among and tw
th links twn animals and popl ar oftn prsntd as univrsally tru most animal soi
whn in fat thy show variation dpnding on th natur of sltiv fors. losly to thir n
By ignoring th ontingny of sltion, laims of univrsality of human so- als an mas
ial institutions and thir immutal natur ar likly to fals. As many of Prhaps spur
th ampls of animal soiality plord rlir hav dmonstratd, partiu- psyhologists lso
lar typs of soial rlationships and soial systms thriv only undr a limitd prsptiv, but t
st of nvironmntal onditions. Why, thn, should W xpt that only on sought to idntify
st of human rlationships or only on typ of soity should volv? lutionary tim. Fo
It is somwhat ironi that politial thory itslf is a human onstrut with anisms tht our
ah prediatd on a diffrnt st of assumptionsaout human natur. on- ostraizing heats
srvativ philosophis oftn dpit humans as fundamntally slfish, ompt- solv suh prol
itiv, and somwhat invariant, whil mor liral philsophis oftn assum usd on rprodu
that humans ar flxil and adaptabl.As is th s with all animals, thr mhanisms that

rganizd swarms to highly polar-
]nmntplayd a ritial rol in un-
r, and onsnsus disilrrmaking
Juals did not know th quality of
th mjority or minority (ouzin
.idnt tht evolutionary
Prosss
pls,spiallythos that undrli
mn animals' hav stimulatd nd
v i o r .N o t s u r p r i s i g l y' o l ] n
tion and partiular aspts of hu-
is this mor apparnt than in th
i t dw i t h t h m j o r h i s t o o m p a t -
tion and immun funtion. During
diffr gntiall from thmslvs,
soitwith fmals that ar gn-
suggststhat -spifi assoi-
'sifying th gn pool of offspring
rd assoiatingwith los kin during
s nfitsassoiatdwith soiality
and ollagus(1995) prformd a
rnilar sul prfrns' l.rfound
:yl idntifid tlr odor lf T-shirts
..plasant''than thos
) gnsas lss
ns.Apparrntly th sam undrly-
l animals may oprating in hu-
But b no mans do ths rsults
:l soiosxualhavior of arrimals
rans in similar ways. Unfortunatly,
: oftn Prsntdas univrsally tru
ing on th natur of sltivfors.
laims of univrsalityof human so-
rr ar likly to fals. As many of
Larlirhav drnonstrtd'partiu-
l systmsthriv only undr a limitd
rn,should We pt that only on
.pof soityshould vo[v?
ory itslf is a human onstrut with
mptions out human natur. on-
ns s fundamntallyslfish' ompt-
r liral philosophis oftn assum
As is th as with all anirals, thr
So iI B huior d Soio bioll91, z . \ I
is no on animal natur. Aggrssion and dominan ar offst to varying d-
grslry ooperation and altruism. What sts human soitis pat from
thosof most thr animals is thir ornplity. Mo lvls of orgenizatiorr
and mor varitisof rlationshipsan form, thus making intrations mor
diffus'But this dos not man th ruls that strutur animal soitis do
not apply to thos of humans. Prhaps thy ar insuffiint,ut th ar not
nssarilyinvalid.
on of th most omplling ampls whr volutionry uls of soiality
an plain novl st of human rlrionships is th so_alld mothr's
brothr'sphnomnon, in whih mn invst in thir sistrs' hildrn as op-
posd to thir own hildrn (Alxandr 1979a, 1979)' At first glalr this
stratgyappears to violat th most si ruls lf kinship. All ls ing
qual,a parnt sharson vragtwi as many gnswith its own offspring
t i o n S as with a siling'soffspring. But in som soitisntlt all ls is qual. This is
.Wst
espially tru in marry Afrian soitis, suh as that of tlr Tongir
(Flinn 1981; Irons 1981). Th land upon whih ths hortiulturalists liv is
rih, and as a rsult, fmalsn driv nough rsoursfrom thir laor to
raisthir hildn witlrout th aid of thir husrrds.In suh ulturs fmal
philandringis high' and fmals an initiat divor. As a rsult, ukoldry
is ommon' and mn ar not likly to faths of thir putativ lrildrn.
vn if offsprir.rgar only half sis, mn will mr losly rlatd to thir
sistr'shildren than to thir own ptativ offspring. Thus, aording to kin-
ship tory,mn r havingvolutinrily rationlly sin thy ar making
th bst of a d job and invsting in th losstgnti rltiv vilal.
Applying ost-nfitand gn-aountingprinipls to th study of hu_
mn soil hvior is th ornrstonof soioiology. uh of rly soio-
bioiogy sought to plain humar"rity's soial roots nd thus studid th soial
haviorof isting ultures minimally impatd y lrVstrnsoitis.Th
hop was sltivprssursthat urrntl1,opratd Wr th sm as thos
that had opratd in tlr past. Although th assumption of ontinuitv is un-
likly t b tru, it dos not invlidat th undrlying soioiologil sarh
for idntifying how nvironmntal prssurs shap htrnran rlatitllrships
among and twnss and within and twnfamilis' Morovr, s irr
most animal soitis,individuals in ths non-!strnizdsoitisar tid
ioslyto thir nvironrnt.hus logil, soial, nd podr-rtiv vri-
alsan rasurdto idntify pattrns of ustion.
haps spurrd on y th Darwinian invstigation of human soiality,
psyholgistsaiso ga nrirringhunrn soial havior from an daptiv
prsptiv'ut thir approah was fundamntally diffrnt aus th
soughtto idntify how natural sltionshapd th human psyh Ovr vo-
lutionarytir. For psyhologists,th r.rltimatqustion ws, ,.Ar th mh-
nisnrsthat our rains us to mak disions aout hoosing lnats or
ostraizinghats th ons that w woul] pt if thy wr .dsignd' to
s o l v s u h p r o l m s ? ' ' ( B a r r t t t a l . 2 0 0 2 ) . T h u s w h i l s o i o i o l o g i s t sf l -
usdon rpodutivoutomSto masuradaptation and oftn igrrordth
mhnismsthat undrli assssmnt and dision making' volutionary
2.'2 Socil Bhuior d Soiobiolog

psyhologists fousd on idntifying tlr mhanisms of assssllnt nd in- Barrtt,L., R. Dunar,ni

formation prossing with littl gard for rprodutiv ouoms. Printon,NJ: rint
Just as an
intgratd approh tl th study of soial havior has mrged withirr th Batn,A. J. 1948.Irrtra
.W.,
biologial tradition' n approah that intgratsmhanisti and funtional Bradury'J. and S. L.
i n m l l o n u r i dts
qusts for undrstandingth zrdptivvalu of th hurnan mind nd th roi
2: 1-17.
it plays in shaping soial havior would th st of oth worlds. It would 1977.Soilorg
ut aross sals and povid nrr onrplt insights ilrto wh humns e- invstmnt pattrns.B
hav th way thy do. lutton-Brok,. H., P. N
Although humans r animals, thy ar ptionalanimals in many ways. I,. Sharp,R. Kansk
The langug thy us is mor ompl thn tlr ommuniation systmsof ontriutions to av
Pro,!igsof th R
othr animals; th ways in whih thy manipulat and div :rnd giv sup-
lutton-Brok,T. H., A. F
port to nd liit it from othrs ar mor sutl than in any othr spis;ernd G' M. MIlrath.200
oniy humans hav dvlopd ultural noms and trditions tht an nrod- oopr:tt ir'hlvii
ifid and passd on t<l hildrn. volutionary ruls will hav muh to say o u z i n 'I . D . , a n d . | .K r u
aout th adaptiv vlu of ts haviors. And whil w sarh for links vrtrats. due
twnpartiular hutnan prtis and partiulr irltmstans'W slrluld ouzirr,I. D.' J. Krus,N
n dd i s i rnn k i ig
not abandon th srh for univrsalsthat might undrli th organizationof
rl, S. R., and P. . W
humansoialit. lon]ooss (logI
Human soitisoltn ornsudividd into olltivsorposdof in. 2:7-15.
dividuals who ar similar gntially or phnotypially. Prhaps this is - rook, J. H. 1964a'Th
arrs simiirity tightns onntions and rpatd intatilns strngthn in th wvrirds(P
1964.Fild
soial rlationships. Th sam tndnisfor individuals to assoit with oth-
r t i nw a v rh i r d s
rs lik thmslvsar oftn sn in animal soitisas thy gt lagr (Run- 2 1 7 - 2 55 .
stin nd Hak 2004). Diffrrrt nvironmnts will ontinu to favor rtin D:rrvkins'k. 1'976.Th S
typs of intration ovr othrs, ut farniliarity will rdu unrtainty and in- mln, S. T. l99.5.An vo
rasth liklihood that oopration,whthr sdn altruism,riproity, Ntilldmof
1 9 9 7 .P r d i i nf
or mutualism, will strong. Languag is a tool tht humans us to rinfor
N. B. Dvis,ds.,B
onntions' to mt out punishmnt, and to roadast th ploits of oop-
o o r d :t s l k w 5l l
rators. In gnal, idas-th mms of Rihard Dawkins (1976)-an e mlrr,S. T.' and L. W. o
transmittd among gnrations just lik gns, with good ons sprading and nratingsstrs. Slcr
ad ons disapparing'Th volution of soial norms that tol virtu may mln, S. T., and P. tl. w
th tra pi of th qution rhat diffrntiatshumarrs frm Inost othr so- of hlprsmongb
Faorg,J., P. G. Parkr,
ial spis. Tsting this proposition is ssntial lf w r vr to know what
Naranjo,nd T. A. $
onnts and sts us apart from othr spis,ut it will b mad virtually im- GalapagosHawk 1B
possil if it is intrtwindwith plitiaI agndas. .l/.,l' d S,'..i.ll,i,
F l i n n , . 1 9 8 1 . U t r i n v
marriag prfrn
lLloGRAPY d ). W. Tinkl, d
d Nu Theor', 4
Aland'R. D. I974. Th volutionof soialhvior.ulRuitl., -l
of oklg Frik, H. W' 975. r,ol
d Sstmtls5: 32.5-383. Zitshrift fr Tirps
1979a.Drinism d m fiirs. Sattl:Univrsityof \X/shinston Gross, M. R., and . I-.
Pss. sunfish. Prodigs
1979.r.olutionnd ultur'In N' . hgnonand .W'Irons'ds.. G u r t l n ' S ' . s . . l . v i n .;
uolutiorBiolog d Hum Bh,luilr: thrrl|ltlklgilPrsptit,, h r d s :F o m i n d i v i d r
59-78.North Situt,: Duuryrss. 85-98.
A l r o d ,R . , a n d W . D . H a m i l t o n .1 9 8 1 .T h v o l u t i o n
o f o o p r a t i o nS . i e 2 1 1 : milton, \. D. 1964. T
1390-1-196. of l lrtiI tlhli

mhanismsof assssmntand in-
. rprdutivOutoms.
Just as an
l hvior has mrgd withirr th
gratsmehanisti nd funtional
u of th human mind and th rol
) th bstlf oth worlds. It would
lltinsightsinto why humans -
ptionalanimals in many ways.
han th ommuniation systms of
rnipulatand div and giv sup-
utl thn in any othr spis; and
rnsand traditiotlstht an mod-
rrary ruls will hav rnuh to say
ors. And whil w sarh 1r |inks
)artiularirumstans,w should
t might undrlith organization of
ld into olltivsomposd of in-
phnotypially. Prhaps this is -
d rpatd intrations strngthn
or individualsto assoiatwith oth-
.l soietisas thy gt larg (Run-
nnts will ontinu to favor rtain
LritywiIl rduunrtaintyand in-
r l l r a s do n l t r u i s m .r
; a tool that humans us to rinfor
d to rodastth xploits of oop-
I Rihard Dawkins (7976)-an
,ns,with good orrs sprading and
oial norms that tol virtr.rmay
:ntiatshumans from most oth so-
sntialif w ar vr to know what
:is,ut it will b mad virtually im-
rgndas.
ri bhavior. ttl Ruitu lf olog
. Sttl;Univrsity of Washingtorr
t. hagnon nd 1X/.Irons, ds.'
;,r:.
! |t tpo lgil rsptit''
l,oiutionof oprarion. Sin 211:
Slil Bhuior d Soiobiolog 25.]
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z'- l-t/-
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)86. Soiology:Origins and trnds. In

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-709.
r lions:h rol of tlr indivi]ul.
45454.
lf tology. Zitshrift fr Tirpsbologi
 th human lin, a
forms from aplikt
uriousl, thir
lngd during th
lngs inludd th.
Human volution animal world rs
1926,1929). Othr
ommon anstor
r Iv4.NIIfnr kind of
..pr-ap''
i n g a p s ( . g . ,N a p
so divrgnt in sh
gn of th humz
Th nintnth-
(.g.,Grgor 1'9|
lr rihnss of th human fossil rord llows rasonal onfidnin r- until th last thr
onstruting <.lur volutilnaryhistory' This rord dmonstrtsthat humn. lonly vois (Fla
like forms appad y aout 6 million yars ago and aumulatd mor Among th man
atrd mor human hratrististhrough tim. As Darwin and his ontm- tionary studis du
poris prditd, th human linag shows lr vidn of dsntwith appliation of moi
modifiation. Prhps th st dmonstrtion of tlris is th xpansion of th omparing th str
rain through tim (Tal 1)' Fossil spis that wr larly humanlik in protins of noth
ody and drrtition 6 to 3 million yars ago had p-sizd br:rinsthat Wr ti rlatdnssam(
orr-thirdth siz of thos of nrodrn lrumans.Just aftr 2 rlillion yarsago to most omparati
som fossils show an inras to aout half th siz of Homo spis, and tim, humans Wt
ovr th pst half million yars, modr:rrrin siz apprs in th fossil than Afrian aps
rord (onro 200.5; Shwi1rtz and Tttrsall 2002, 200, 2005; Dlson wr far mor dist
.\X/ils
t al. 2000). (Sarih and
Although Darwin nd othr arly volutionists wr knly interstdin firmd ths arly
h u n l . r n v o l u r i o n n d n l a d g , rs t r i d s i r l i n t r p r t i n 8 o m p r t i v m o r - (.g., alunrin) wl
phology fom an vllutionarypoint of viw, vry fw prmodrn humn fos- 1,967).onstany
sils wer known in thir day (Huxly 1863). By th turn f th twntith lutionary linags
ntury' its f Nandrtals nd ..Java 1n'' ntrd th onsiousnssof divrgn and thr
volutionary iologists (Kith 1915). B1,th turn of th twtlt-firstntury' Sarih and.Wilsorl
thousands of prmodrn fossils had found thir wy into th world's mus- of svral mamm
ums, and knowldg of humn volutiorl had grown sptaularly (Dlson From this thy a
t l . 2 0 0 0 : o n r o y 2 0 0 5 ; K l i n 1 9 9 9 ; S h w a r t z i r n d T a t t r s a l l2 0 0 2 ' 2 0 0 ' mans and Afrian
2005; Holloway t aL.2004). Th rihnss of th rord is du to th intlli- finmnts pla th
gn' skill, zrnddiligrrof th fildwlrkrswho find th fossils and dtr. urrnt stimats(
min tlril iologil and glogial oIrtxts. 1989; Chn and L
trson t a|' 206
th advnt of DNl
our Pla in Natur latd to th Afri
ary linags is rl
It was lar to Drwin and his ontmporaristhat popl sh:.rrda ommon A sond impor
anstor with living aps mor losly thzrnwith any oth anirnal' Thomas th mthods of
Huly (1863) Was a partiularly strong advoat ftlr an aplik anstryof annot appli
256

n animal world rsmldth tarsir mor losly than any ap (Jons 1918,
lows rasonalonfidn in r-
rord dmonstratsthat humn-
/ars ago and aumulatd mor
r i m .A s D a r w i n a n d h i s o n t m -
ws lar vidnof dsnt with
ion of this is th pansion of th
s that wr larly humanlik in
3o had ap-sizdbrains that wr
ans.Just aftr2 million yars ago
rlf th siz of Homo spis, and
L rain siz appars in th fossil
r s a l l2 0 0 2 , 2 0 0 3 , 2 0 0 5 ; D l s o n
rtionistswr knly intrstd in
i n i n t r p r t i n g o m p r t i v m o -
v' vry fw prmodrn human fos-
63). By th turn of th twntith
tan'' ntrdth onsiousnss of
l turn of th twnty-first ntury,
l thir way into th world's mus-
had grown sptaularly (Dlson
7wartzand Tattrsa||2002. 2oo.
; of th rord is du to th intlli.
rs who find th fossils and dtr-
\LS.
rristhat popl shard a ommon
rn with any othr animal. Thomas
Ldvoatfor an aplik anstry of
Hm uolutio 257
th human lin, and Hakl (1s6s) imagind and namd a spifi sris of
forms from aplik to humanlik.
uriously, thir ap-man thory (as it was latr alld) was sriously hal-
lngd during th first svn dads of th twntith ntury. Ths hal-
lngs inludd th viw tht th last ommon anStor w shard with th
|926, |929). other arly twntith-nqurysintistsronstrutdth last
ommon anstor as mor monkylik than aplik (Straus 1949) or as som
..pr-ap''without long fingrsand uppr.ody spializationsof liv-
kind of
ing aps (.g.,Napir and Davis 7959I. Bausmodrn humans appar to b
so divrgnt in shap and bhavior from othr nimals, th tim of divr-
gn of th human volutionary linag was oftn givn as quit anint.
..ap.man'' thry ontinud to hav spPortrs
Th nintnth-ntury
(.g.,Grgor \9 |6, 1926, 7927a, 1927, 1'928,19 41 sashurn1 95 1 ), ut
until th last thr dads of th twntith entuy ths wr rlativly
lonly vois (Flagl and Jungrs 1982).
Among th many important hangs in viw and mthod in human volu-
tionary studis during th last fw dads of th twntith ntury was th
appliatin of molular iology. Immunology providd th first insight. By
omparing th strngth of th immun systm,s rspons y on spis to
proteinsof anothr spis,Goodman (1963\ quantifid an stimatof gn-
ti rlatdnssamong primat spis. Th rsults wr shoking at th tim
to most omparativ anatomists.Contrary to prvailing undrstandingat th
tim, humans wr mor similar to Afrian aps (himpanz and gorilla)
than Afrian aps Wr to th Asian grat p (orangutan). Monky spis
wr far mor distantly rlatd. Mor sophistiatdimmunologial mthods
(Sarihand.!ilson 1966) and DNA hyridization (Hoyr et a|'.1964) con-
firmd ths arly rsults and rvald that hangs in som maromoluls
.Wilson
(.g.,alumin) wr rlativly onstant through tim (Sarih and
1967).onstany of hang prmittd stimatsof tim of divrgnof vo-
lutionary linagsas long s th rat ould aliratd y a known dat of
..molularlok.''
divrgnand thus provid a asis for dvlopmntof a
'Wilson
Sarihand (\967) usd th dat of 60 million yars for th divrgn
of svral mammalian ordrs to alulat th rat of hang in alumin.
From this thy alulatd that th divrgn of th linags that ld to hu-
mans and Afrian grat ps ourrd 4.2 million yrs ago. Susquntr-
finmntspla th dat somwhr twn 10 and 5 million yars y most
urrntstimats(Kumar and Hdgs 1998; Kumar t al. 2005; Holms t al.
1989;hn and Li 2001; Arnason t al. 2000; Pilam and Young 2004;Pat-
trsont a|.2006; Glazko and satoshi 2003; Stauffr t al.200]). !7ith
th advnt of DNA squning it is now lar that humans ar vry losly r-
latdto th Afrin grat aps)and th tim of divrgnof thir volution.
ary linagsis rlativly short from th prsptivof mammlian volution.
A sond important hang in th latr prt of th twntith ntury was in
th mthods of omparing morphology of spis. Molular approahs
annot applid to fossiis muh oldr than 70'000 yars (Kraus t al.
l 1. Speis,dats,ody size, rain siz,nd postriotoot siz in arly hominids

ass Staura Brain Postanine

Taonl Dats2 Ma1 Fmal Ml Fmal V5 wight6 tooth ara7 Q' MQ,
troglodtes
Shlthropus thdests
xtant
7-6
49 41,

;
65
95 294
:Y
rrori tugesis 6 +l lJl 329
Ardip itb eus kd|| 5.2-5.8 .)/
u strlop i tll us msis 4.z_.9 ; 428
A ust r l op it h t's [r nsi s 3.9-3.0 45 29 151 4; 4; 460 2..4 1.7
us trlop it ll cus fri nus .0_2.4 4l 30 18 115 452 448 516 2.7 2.
Prnthropus ethiop ius 1-7 11
41 47 688
Prthropus boisi 2'_I.4 49 t+ 17 r24 521 514 756 2.7 2.7
Prnthr ops rt:lbustus 1.9-1.4 40 .)z 132 110 530 523 588 3.0 2.2
Astrlopitbes grbi 2.5- 4s0 446
Hrllllo bilis 7.9-r.6 -)/ ; 131 100 612 601 478 .6 1.9
Hlmo rudolfesis 2.4-1.6 60 51 160 150 752 76 572 -1.1 1.5
Homo ergstr 1.9-1.7 66 56 180 160 871. 849 - ) // 3. 0.9
Homo eretus 1.1-0.01 66 56 I,050 1,01.9 402 4.0 1.0
Horo bidelrgesis o.6-0.2 77 56 1,\94 \,156 389 4.2 0.9
Late H omo n.ldrtb.llsis o-2-o.o3 77 66 ,t o. -/ .l
7.62
58 7,414 33-5 4.7 0.7

Homo floresiesis 0.09-0.01 29 t06 380 78 2.6 1.8

Homo spies tant 58 49 775 t61 1'350 4 5.8 0.9
1. Tatlnomr,sdon Klin (1999)rvit dditionsfrom Brunt t al. (2002),Snutt al. (2001)'Brown t al. (2004),and HiI-S|assi t l. (2004).
2. Dts (in millions of y:rrsg) r from lin \1999)' wih dditionsfrorn Bruntt al. (2002),Snutt l. (2001),Br<lr-l,n t a|, (2004)anl Hail-Slssi
t al' (2004).
. Body rassstimats(in kg) ar from nr 0992\ ptfor th follor-l,ing: o. tugsisis frorn Nkatsuksat al. (2007)'whr 4] is th mid-
point of thir r llg nd thy did not spify s; . msis mal is from L ky t al. ( 199.5); . lsisfmal is alul:rtdfrlm th rtio of lrll and
 s
A str l op it h us me.si 4.2-3.9 51 JJ +L6 l.a

ustrlop itheus frns 3.9-3.0 45 29 ; r* 48 44 460 2.4 1.7

3.0-2.4 41 30 138 115 452 448 516 2.7 2.0
str lop itheus fr i us
rthrlpsthiop ius 2.7-2.2 410 407 688
2.3-1.4 49 17 124 52L 5\4 756 2.7 2.7
Prthrops boisei -)a

1.9-1.4 40 110 530 52 s88 3.0 2.2

Prthropus robustus L t -'tL

ustr lop ithus grhi 2.5-? 450 446

;. 7 L ; 100 612 61 478 .6 1.9
Homo hbilis 1.9-r.6 -)/
Homo rudolfsis 2.4-1.6 6 51 160 150 752 76 572 3.1 1.5
Homo rgster r.9-r.7 66 56 180 160 871. 849 77 3.3 0.9
1.1-0.01 66 56 1,050 1,01,9 402 4.0 1.0
Homo rectus
0.6-0.2 77 56 1,194 1,156 89 4.2 0.9
Homo hidelbrgensis
Late Homo nedertblnsis 0.2-0.03 77 66 1,414 1.,362 335 4.7 0.7
",

.l
omo floresisis 0.09-0.01 29 106 3B0 78 2.6 .8
Homo sIpies xtant 58 49 t75 16,|' 1,350 4 s.8 0.9
1. Taxtlnlmysdon Klin (1999)with dditionsfronr Brunt t al. (2002),Snutt l. (2001),Brown t al. (2004)'and Hail-Slssi t l. (2004).
2. l)ats (in millilns<lfyrs g) r from KIin (1999),,ith additilnsfrom Brunt t al. (2002),Snutt al. (2001),Brrlwn t al. (2004) and Hil-Slssi
t l. (2004)'
. Bod mss stimts(in kg) ar flmHnr (|992-)eptfor th fol|wing:. tugsisis fr<lmNktsuksat al. (2007)'whr 41 is th rrrid-
point <lfthir rang and thy did not spifys; . msisml is from Laky t al. (1995);. lsisfmalis lulatdfrom th rtio of ml and
fml in . frtlsis;. rgstr'H. rts,H. heidlbrgzs,and |ate H. drthIsisr from Ruff et al. (1998); and H. florsizsisis from Brwn t
l. (2004)
4. Stturstimts(in n.r)ar from Hnr (991) pt for . tugeesis'whih is from Nktsuksat al, |207)'whr 13l is th midpoint of thir
.Walkr
rng and thy did nlt spify s . rgstr,whih is from Ruff nd and lat . drthlsis'whih r
(1993); H, rtus,H. hidIbrgesls,
fronl Ruff t l. (1998);and H. |lorsiesis, Brown t al. (2004).
whih is frln.r
5. V is ranil paityin fror sourslistd in MHnry (1994a;1994)u,ith th addition of S. thdsls(Zollikofr t 1.2005);,L.444-2 (540
) to . frsis(imbl,rsonlommuntatin);Stw 505 (51.5) to . fl.ius(lnrlyt t. 1998);P. thi<.lpius (Walkt al. 1986);KGA 10-525
(545 ) to P. bisi (Suw t a|. 1997|; and BOU-VP-12/1 0 (4.50) to . grhi (sfaw ct aI. |999). ' rtus,H. hidlbrgsls,and |ate . dthl-
sis ar from Ruff t 1.(1998), and H. |loresizslsis from Brlwnt l. (2004).
6. Brin wight is alulatdfrom formula (6) in Ruff t al. (1998).
7. Postanintoot ar (in mm2)is th sum of prdutsof bual-lingualnd rsio-distllrrgthsof l,., ', r"rdM. irnd is takn from MHnr |1994)
with th ddition of S. tchdsisstimtdfrom TM266-02-154-1 reportdin Brunt t l. (2002);(). tugsisfrlm Snutt al. (200.lI;' Ar. kdbb from
il-SIssi t al. (2004); r. rmidus frtlmlihitt al. (1994); . msisfrom I,kyt l. (1995); H. H.
rtus, hidlbrgsls,nd lat ' n-
drthIsisfrorn Wlpoff (1971,1982I; and H. |lorsisis from Brown t al. (2004).
8. Q is th nphlizationquotintalulatds ritmss dividd y (11.22xbody mass().'6) from Mrtin (1981).
9. MQ is th mgadontialquotintdrivdas ostnit-t tooth ar dividd by (12.1.5bodymass()86) from Hnry (1988).
260 Hum uolutil

2071r|ando t l. 2006), so morphology is th only guid tl undrstand- lins of dsnt. F

ing rlationships.los volutionary rlationshipsan infrrd from simi- 1996). Baus Hl
larity in morphology (s Figur 1)' ut not withut arful analysisof th ary divrgen, it is
auss of rsmlans.volutionary iologists hav always n awarof is uniqu voluti
this, of ours, ut a formal approah to mak omparisons of form only traits that ar not
took hold in th latr part of th twntith ntury. shard uniqu tra
Willi Hnnig (I966) formalizd th approah to omparing morphologyof ..shard
drivd.''
spis y noting that rsmlan of form did not nssarily imply los h powr of th
phylognti affinity. Similaity may du to sharing primitiv hratri hlps ronil th
tis that ar not hlpful in dtrmining iologial rlationships.Aspetsof th fossil rord. As
th dntition of th 28-million-yar.ol d Propliopitbus, for ampl, rsm- now rflt our tru
l thos of modrn humans mor than thos of modrn aps (Kurtn 1972), (orangutans, gorillz
but ths ar du to th rtntilnof anstraltraits and not to th volution mans into Homini
of uniqu traits shard only y dsendantsand not presnt in othr losly that rognizs th
rlatd spis.Somtims similaritis r du to paralll or onvrgntvO. lad as th family
lution rathr than dsntfrom a ommon anstor.Th wings of irds'a, familis Gorillina
and uttrflis ar similar, ut not us thy volvd from th sam mans). It is a usfu
wingd anstor. Th trm homopls is usd to dsri th phnomnon organi forms and t
whr organisms shar uniqu traits that volvd indpndntlyin sparat

Bfor Hominir
ll0mo
reus
our los gnti r
{n'.,,./
origin of our volul
r"l
natly, th fossil r
panzs' and humar
bttr known. Disr
rvaling. In sdim
forms appar that ]
(Pilbam 1982). Siu
uniquly orang-lik
profil of th fa t
faturs that among
Th fa of th As
th Afrian lad (g
(flakedston of ths living sp
tifds may not' y th logi,
unknown)
ost palontologis
qualify uropan f.
ago), ouropitb,
(1'2.5_| million y
--\
as part of th Afri
Sahelanthropus
thadensis Afria for ]\4ion
spars' but dilignt
Figur 1. A working hypothsisof volutionryrlationshipswithin th human pitheus (16 to 14 r
fossil rord,rdrawn from R. G. Klin. 15 million yas ago
 umn uolution 261

4y is th only guid to undrstarrd-
ionshipsan infrrd from simi-
.rotwitlrotrt arful arralysis of tlr
logistshv alwas n awar of
l mak omparisorrsof form only
1 ntry.
roah to omparing morphology of
rr-rdid not nssarily imply los
u to sharing priritiv haratris-
liologial lationships.Aspts of
"olioithcus, for ampl' rsm-
.osof nrodrn aps (Kurtn 1972),
stral trits and not to th volrrtion
lts and not prsnt in othr losly
: du to paralll or onvrgnt vo-
l nsto.Th wings of irds' ts,
ausr thy volvd from th s:rm
usd to dsribth phnomnon
v o l v di n d p n d n t l yi n s p a r a t
lins of dsnt. Homoplasy is prvasiv in human volution (Hnry
1996|.Becaus Hnnig's mthod fouss on th haratristis of volution-
ary divrgn, it is oftn rfrrd to as ldistis' ld in this voaulary
is a uniqu volutionary ranh f th tr of lif d6nd y shring uniqu
traits that ar not shrd in losly rlatd ranhs. nnig alld suh
shard uniqu traits ..synapomorphis,'' whih is usually translatd as
..shard
drivd.''
Th powr of this logi pvds volutionary iology (Hull 1988) and
hlps ronil th findings of molular iology with th intrprttion of
th fossil rord. As a rsult, th trms usd to lassify our biologial rlativs
now rflt our tru gnti rlationships. Th old tradition plad grat aps
(orangutans'gorillas, and himpanzs) into th family Pongida and hu-
mans into on-rirrida'but th tu gnti affinitis for a rlassifiation
that rognizs th Asian grat ap as th family Pongida and th Afrian
lad as th family Hominid. \/ithin th family Honrinida ar th su.
familis Grillina (gorilla), Panina (himpanz), and Hominina (hu-
mans).It is a usful rmindr that w ar tightly nstd in th Lrshof lif's
oganiforms and that th origin of our uniqu linag is vry rnt.

BforHminins
ll0mo Artifat raditions
f/oresiens 1"
,t". st"'* g"

la .
J Upper PaIeo|ithi
our los gnti rlationship to th Afrian grat aps implis an Afrian
n(........-
[ )i - iddle stone Age & origin of our volutionary linag. Fossil disovrisonfirm this. Unfortu-

ff::'|'
\j!g]g|g!gg]i. natly' th fossil ord of th origin of th Afrian lad (gorills, him-
A [-) panzs'and hunrans)is poorly doumntd.Th Asian lad of our family is
ffi ."uteanffi ttrknown. Disovris in th foothills of th Himalyas ar partiularly
tt 13
rvaling.In sdimntsthat dat twn aout 12 and 8 million yars ago'
v

,.'"'r'i,
lParanthropus 1 . 6 5-
forms appar tht rsml tlr ofangutan in mny spifi faial faturs
|
J,.'",|^
d7\ oldo*n fl) (Pilam 1982|. Siupithcus is th gnus nam that inluds fossils with th
\_)/ \,
uniquly orang-lik faturs of narrow spir twn th ys, ..prowlik',
2.5
profilof th fa low th nos' onstrution of th palt, and matly othr
fatusthat among th living spis our in th rnguta n (Pongt)'
Th fa of th Asian Iade (Siupithelts and Pogl|ontrasts with that of
,,.^".:*,*, th Afrian lad (grillas,himpanzs,and humans), ut th rsmlan
, of ths living spis may du to sharing primitiv haratristis and
( f | a k ds t o n e
theUs atifts may not' y th logi of ladistis, usful in dtrminingtru rlationships.
unknown)
ost palontologists apt ths rsmlns as suffiintly r"rniquto
qualify uropan fossil gna suh as Dropithus (8_13 million yas

I-'
.
ago),Ournopitheus (9_10 million yars ago), and prhaps Pirolpithes

l.
(12.5-1million yars ago) (Moyi-Soli et a|' 2004; Bgurl and rwrd 2005)
as part of th Afrian lad (Bgun 1994;F|eag|e 1999). Th fossil rord in
Afria for ion mmrs of Hominida (Afrian aps and humans) is
spars' ut dilignt attmpts ontinu to promis nw disovries' en.
within th humn
nary rltionships pitbus(16 to 74 million yas ago) and rlatd gnra (qutoris, 16 to
i5 million yars ago) appar to part of th Afrian lad with living nrm-
262 Humn ulllttio

brs, ut ah has its own spializationsthat mk thm lss attrtivas for sltion that fav
andidts for anstry of livirrg spisof ominid (rossin and Bn- g r s .A n r h h o m i n i
t 1997\. of th formn lltag
T.h palonvironmntwithin whih our nstorsvolvd was inflund howvr, Sbeltbr
profoundl y drmati flutuationsirr world limats ovr th past 7 millio t h h u m n l i n a g .I
yars (Vra et a|. 19951Bromag and Shnk 1999; Potts 2007; Pikford mans. It has normou
2006; Bhrnsmyr2006).Intnspriods of old tmpratursin highlati. ps or arlir mm'
tuds rsultdin pisodsof drought in th tropis' Thes apprd:rt16_13' having n Issdv
6.5-5, nd 2.8 million yars ago' and ginning at 2.5 million yars ago' th h disovr of S
arth prind21 yls of glaiation in highr lvations and ltituds n s so f i h l B r u n t
aompanid y rdution in rainfall in quatorial lands, spiallyAfria. Chadian studnt who
Haitats fvord y hominids (anstorsof th Afrian aps and humans) Afrian Rift, whr
ontratd and xpandd. Th fossil rord of hominids in th iddl nd minds us to mind{l
Lat ion is frustratingly inomplt, and almost nothing is known of th Almost all of th .
linags tht ld to himpanzs and gorillas. Suprisingly, muh more is omes from th Afri'
known of th fossil rord of our own rnh, th Hominina. vnts'many lt i<
mnts layrd y vol.
nrurally ouringp<
Th arlist Hominins of ash in thiopia, K
th arly part of th s
Rogr-ritionof th rlir mrrsof Homirrina is guidd by undstanding ourrd.
gaind from moltrlr iology and ladistis.Th molular lok rakets Th fist haptr in
th im of divrgn somwh twn 10 and .5 million yars y most to just lss than 6 mill
urnt stimats.By th logi of ladistis, th arlist astor of humans 2001; Pikford et a|,2
an rognizd y th prsnof traits uniqu to th linag.This is ob- this spis omind '
vious ut is asily onfusd y naiv viws of pr-Hnnigin omparativ appar to adaprd
morphology that ontinu to plagu human volutionary studis.If it looks J u n g r s2 0 0 8 ) .
.W.
lik an ap' it is not nssrily ludd from th human ralm. now Slightly latr in tim
know f n traordinary rord of an aplik-oming-lrumanlik s- thiopian spimns rt
qun of fossils.Th arlistmmr of this squnlolks vry aplik.By Gabril t a|.200l; H
ldisti logi, that is atly what on would pt. with rrori tugs
Th arlist andidat for mmrship in th sufarrrily Holinina most rspts,ut it d
Shlnthropus thdsis (Brunt et a\. 2002). This prious fossil skull ths is anin tooth
frorrr had was found in sdirnntsthat ontilind funa linkd with databl sil or modrn aps an
sdimnts in thiopia nd North Afria 7 to 6 million yars ago. Its ovrall the ability to hyprxtt
apparan is quit aplik' inluding an stimatdrain siz slightly low It livd in a losd wo
th avrgfor living himpanzs (Tal 1). But y th logi of ldistis,it
sharswith latr hominins uniqu taits that dfin our linag.Th mostim-
potant of ths is th anir-rthat, unlik that of ps and primats in gn- Austrlopithecus i
rl, dos not projt or war against its lowr prmolar. This is a signifiant 4.2-2.5 Nlillion Y
fatur tht ws rognizd as an important diffrn twn aps and
popl y many, inlding Darwin \|872\. h fssil vidn f
Small nins ar signifiant.Darwin (|872) notd that whn human n. Sh lt brops t h di
stors first stood rt' thy frd thir hands and ould randish wpns yars ago thr appar
fo thrat nd fight, thus no longr rlying <rnlarg anins. Small nins cus nlnlnsls that pos
arry spial signifian,ut it is not too diffiult to imagin oth rasons ( L a k ye t a I . 1 9 9 5 , 1 9

that mak thm lss attrativ as
Hominida (rossin and Bn-
r anstorsvolvd ws inflund
rld limatsovr th past 7 million
hnk 1999; Potts 2007; Pikford
s of old tmpratursin high lati-
j tropis.Ths apprred at 16_13,
nning at 2.5 million yars ago' th
in highr lvations and latituds
:quatoriallands, spilly Afria.
of th frian aps and humans)
rd of hominids in th iddl and
Lndalmostnothing is known of th
rillas. Surprisingly, muh mor is
nh, th ominina.
nininais guidd by undstanding
tis. Th moiular lok rakts
.n 10 and 5 millin yars y most
:s,th arlistanstor of humans
s uniqu to th linag.This is o-
w s o f p r - H n n i g i a n o m p a r a t i v
an volutionarystudis.lf it looks
.W
l from th human rirlm' now
r aplik-onrirrg-humanlik s-
his squnlooks vry aplik. By
'uld xpt.
p i n r h s u a m i | yH o m i n i n a i s
2002). This prious fossil skull
lntaind faun linkd with datal
'to
6 million yars a]o.Its ovrall
stimatdbrain size slightly low
1). But v rh Iogi of ladistis, it
t dfinour linag.Th most im-
t h t o f p s n d 1 r r i m a t si n g n -
lwr prmolar. Tlris is a signifint
.tant diffrn twn aps and
t872) notd that whn human an-
ands and ould rndish wapons
lg on larg anins' Smll irnins
l diffiult to inragirrothr rasons
Humn uolution 26
for sltiontht favrs rdution to rliv th xpns of growing suh dag-
grs.Anothr hominin novlty sn in S. tbdensis is th forward plamnt
of th foramn magnum' In ovrall morphlogy (Zollikofr t al. 2005),
howvr, Sblthroprzs is aplik, as rfits a spis los to th origin of
th human linag.Its rain is minusul omprd with that of modrn h_
mans.It has normous brow ridgs that r vn largr than thos of modrn
aps or arlir mmrs of th gnus Homo whr row ridgs appar aftr
havingn lss dvlopd in arlir spisof our linag.
Th disvr of Shlthropus is du ttl the prsistn,skill, and lvr-
nssof Mihl Brunt and his tam' inludig Djimdoumaay Ahounta' th
hadian studntwho found th skull. Its loation 2'500 km wst of th st
Afrian Rift, whr almost ll th othr alist hominins wr found, r-
minds us to mindful of th limitd tnt of ou smpl.
Almost all of th vry arlist fossil rord of our volutionary history
oms from th Afrian Rift Vally. Baus of xtraordinary gologial
vnts' many lat Mion to Rnt trrstrial fauna ar prsrvd in sdi-
mnts layrd y volani ashs that an datd by th radioativ day of
naturallyourring potssium-40 to argon-4O.Fossils urid bnath layrs
of ash in thiopia, Knya, and Tanznia povid most of th vidn for
th arly pat of th stoy of human volution arrd th ontet in whih it
ourrd.
Th first haptr in this story ols from fossils nath ash layrs datd
to just lss than 6 million yars ago in th Tugn Hills of Knya (Snut t al.
2001; Pikford t al. 2002; Sawada t al. 2002). Namd rrorin tugesis,
this spisomind aplik tth and forlims with humanlik thighs that
appar to b adptd to ipdality (Pikford t al. 2002; Rihmond and
Jungrs2008).
Slightly latr in tim (5.8_4.4 million yas ago) is th olltion of th
thiopianspinrnsrfrrdto as rdipithcus (Hal|.Slssi2001; Wold-
G a r i l t a l . 2 0 0 1 ; H a i l - S l a s s i t l . 2 0 0 4 ; W h i t e t a l r .| 9 9 4 , t 9 9 5 ) . A s
wit rrori tt,tgsisand as ptd in n arly anstor' it is aplik in
most rspts,ut it dos shar rtain ky traits with latr hominins. Among
thsis a anin tooth that is lss projting and lss pointd than that of fos-
sil or modrn aps and a ranil s that is shortr. Its tos apparntly had
thaility to hyprtndlik thos in ipdal humans (Hail-Slassi2001).
It livd in a losd woodland nvironmnt (WoldGaril t l' 1994.200t\.
AstrIopithecusand othr Hominins
4.2_2.5illion Yars Ago
Th fossil vidn for hominin ipdality is hintd in intrprtations of
Shlthropustchdesis, orrorin, and rlipithus' ut 4.2_.9 million
yarsago thr appar in th fossil rord fossils attriutd to Austrlopithe-
usmensistht PoSsSsa shinon with nut-tlrousipdal spializations
( L a k y t a | . | 9 9 5 , 1 9 9 8 ; ! 7 a r d t a l . 2 0 0 1 ; W h i t t a L . 2 0 0 6 ) .T h i s s p i s
264 Hum uolutio
inisor roots ar no l
rtains numrous aplik haratristis,lik its prdssor'ut shrsr- A. frsis. Th rItir
tin volutionary novltiswith all latr hminins that a not psntin th angle ar ginning to l
more primitive Ardipithcus' ost onspiuous is th thikning of naml hominin xpt P. tbt
that oms haratristiof all ltr hominins. Its uppr anine has thik- h South Afrian s
nd enaml on th hk sid of its ap. Th molars r palrddfrom ontmpoary with . .
sid to sid, and th first and sond molars ar not markdly diffrntin som traits that appar
siz. Th r tu is mor humanlik. Its diduous lowr first molar is r. spis (Asfaw t al. 19
potd to intrmdiat twn thos of rdipitheus and A. frensisin mdial to th nasal ap,
siz and shap. In th rst of th sklton it is vry muh lik latr hominins nathism, rdud iniso
(fw dtails of rdipithus ate pulishd), spially its lbow, kn, and
faial ontour.
ankl.
Th sampl of . fresis (3,6_2.9 million yars ago) is st known from
Latoli (Tanzani) and Hdar (thiopia), is latr in tim than . mesi Th ..Robust,,Aus
and shars drivd haratristis with latr hominins suh as an ar tub that
is more roundd in outlin, anin with a shortr rown nd highr shoul- In 1938 Rort Broom
drs, a mor humanlik lowr first diduous molar and prmannt prmolar' volution y his dsri;
and a lss sloping hin. Also, on of th wrist ons (th apitat) is lssprim. Its teth, jws, and fa
itiv (although not fully modrn) (Johnsont al. 1982; Laky eta|'\995, Broom and susqunt
1998; Whit et a|. 2006; Almsgdt al. 2006; Kiml et aL.2006)' tion (Roinson 1972).
Kthropus pltops (3.5-3.3 million yars ago) ovrlaps in tim with shrs many drivd f
A. frsis ut appars to quit distintiv irr its mophologv (Laky my losly rlatd l
t al.2001). In som rsptsit is mo primitiv than its ontmporary.tr minins togethr as ..ro
xampl, it laks a ptrous rst on th tympani on and has a narrow e- ppropriat. Th arlis
trnal ar ttr with a small aprtu mor lik that in arlir homininsand (2.7_2. million yars a[
not lik that in , frnsis. In othr respts it rsmls muh ltr ho- usts'' ot omo that ar
minins' partiularly H. rudolfsis,in having a rlivlyflat fa,tall hks, massiv hk tth an
smll molars, and forward faing hkons. Ths lattr traits ar rltd haratristis of A. fr'
..suggsts
to mastiation that' aording to Laky t l. (2001,437), a dit-
P. robustus (2-1 million
drivn adaptiv radiation among horninins in this tim intval.''It is possil do shar apparnt volu
that th rsmlans twn H. rudolfesls and K. pltrlps ar du to tralopithins, suh as i
phylogntilly indpndnt vnts drivn y adaptations to similar situa- of th palat, rar or as
tions and ar, thry,on mor ampl of th prvasivnssof homoplasy' a v r i I t t l w a k a n i
It is qually possil that th rsmblansrprsnttru homology and im- and nlargd rain (Skl
ply a lad that links . pltops with H. rudolfsis' Laky t al. (2001)fa. Grin 2004).
vor th lattr viw, whih suggsts transfrring H. rudolfesls to th gnus
Kenthro1lus (i.e., K. rudolfsls). Although th f of Kthropus ap-
pars similar to that of H. rdolferz'sls,som or all of this rsmlanmay Origin of Homo
du to pnsion raks that hav distortd Kenthrops onsidraly
(Whit200).
on might dspai at th
Th nt-oldsthominin spisis . grbi (2.5 nrillion yars ago).It has genus Homo, ut from t
som shard drivd traits with latr and ontmporary hominin spisrl- traits' th fossil rord is
ativ to . frsis (Asfw et a|' 1999\. Rlativ to . frsls, ir shars du. In on spis, FI. l
..roust'' australopithins asolutly and rl-
with . frius and th to rmain lik that of
tivly largr hk tth. Its uppr third prmola is rportd to hav a mor and powrfully built sho
oval outlin nd n msioual lin etnsion. Its prmolars ar ginning th assoiation of the hir
to show som pansion (molarization), and its naml is thikr. Its uppr
 Hum t,olution 26''

inisor roots ar no longr latral to th nasal aprtur' as thy ar in

ik its prdssor,ut shars r.
. fresis.h rlativsizsof th postrior tmporalis nd paritomastoid
minins that ar not prsnt in th
anglar ginningto rdu, as thy do in all othr spiesof post-frsis
uous is th hikning of naml
hominin pt P' thiopius.
ninins. Its uppr anin has thik-
Th South Afrian spis'A. fricus (3_2 million yars ago), is roughly
. Th moirs ar pandd from
onteporary with . grhi and latr in tim than . frensis. It posssss
lars ar not nrarkdly diffrnt in
som traits that appar to drived and shard wit Homl rlativ to ths
diduouslowr fist molar is r-
spis(Asfaw et aI. 1999)' Ths inlud uppr latral inisor roots that ar
ff rdipithcus and A. frnsis in
mdialto th nasal aprturJ fltt livus ontour, rdud sunsal prog-
it is vry muh lik latr hominins
nathism'rdud inisor proumny' and rdud ipartit latral anteior
J), spiallyits lbow' kn, and
faiai ontour.

lion yrs ago) is st known from

is latr in tim than . mesis'
Th ..Roust''Australopithins
r homininssuh as at] ar tu that
a shortr rown and highr shoul-
In 1938 Rort Broom ognizda mostly unantiipatdhaptr in human
us moiar nd prmnntprmolar,
'istbons(thapitat)is lss prim_ volutiony his dsription of wht h a||dPrthrops (Boorn 193s).
Itstth,jaws, and fa wr massivly uilt (hnth nam robustus\, and
r n t a l . 1 9 8 2 ;L a k e t a | . 1 ' 9 9 5 ,
Broom and susquntauthors rgardd it s a sid ranh of human volu-
1 0 0 6 ;K i m l t l . 2 0 0 6 ) '
tion (Robinson 1972). Howvr, in many important rspts Prnthrops
)n yars ago) ovrlps in tim with
sharsmany drivd fturs wit Homo (Sklton and Hnr 1992) and
intiv in its rnorphology (Laky
'imitiv than its ontmporary. For may losly rlatd to ou gntls. It is omnron to lump lg_tooth ho-
mininstogthras ..roust'' australopithins,ut this grouping may in-
mpni on and has a narrow x_
. lik that in arlir hominins and appropiat.h arlist of th ..roust'' austlopithins,P. ethiopius
(2.7-2.million yars ago), sharsr.ryfw drivd haratrswith latr ..ro-
;pts it smbls muh latr lro-
usts''or Hom tlrat ar not ritd to hyprmastition(Grin 1988). It has
ing a rlativlyflat fa, tall hks,
massiv hk tth and hwing appratlrs ut rtains th primitiv skull
ns.Ths lattr trits r rlatd
haratristisof . irsis (!alkr et aI. 1986), Th latr ..roust'' spis,
t al. (2001, 437)' ..suggstsa dit-
P. robustus(2_1 million yars ago) arrdP' boisei (2'-I.4 million yars ago),
; in this tim intrvl.''It is possil
do shar apparnt volutionay novlties with oo rlativ to othr aus-
fensis ad K. pltops ar du to tralopithins, suh as a paraoli dntal arad shap, dp antrio depth
:n y adaptationsto similar situa-
of th palat' rar or absnt distma betwn th uppr inisors and anin,
of th prvasivnssof homoplasy.
a varilto wak nin jugum and fossa, rdud sr-rnasal pognthism'
s presnttru hornology and im-
and nlagdrain (Sklton and N{Hnr 199\ Strait t a|.7997i Strait and
"udolfensis.Laky t al. (2001) f- Grin 2004).
rring H. rudolfesis to th gnus
lugh th fa of ennthro1lus ap-
l or all of this rsmlanmay
Origin of Homo
ttd entbroptts onsidraly

on might dspair at th apparntly onfliting vidnfor th origin of th

,rhi (2.5 million years ago). It has
gels Homo, ut from th prsptiv of the umulation of shard drivd
ontmPoraryhminin spis rl-
traits'th fossil rord is lss prpling. Brains pand, and k tth r-
Rlativ to . frnsls, it shars
du'In on spis,H, hbilis (2._1.6 million yars ago), th ody appars
rlopithinsabsolutly and rla-
to rmain lik that of ltstrIopithecs,small with rltivly long forarms
rmolaris rportd to hv a mor
andpowrfully uilt shouldrs (Hauslr and Hnr 2007).If on aprs
lnsion'Its prmolrs ar ginning
thassoitionof th hindlimb fossils found at th sam sit and tim as th
rnd its naml is thikr. Its uppr
266 Humn uolutio

rniodntal rmains of H. rdolfesis (-1.9 million yars ago) as longig Th gologilly lt <
to that spis' thn the mor humanlik ody proportions and hip arhit- dmonstrats th fallay
tur first appar in this spis just aftr 2 million yars ago (MHnry and progrssive stags. Populi
offing 2000). Both of thsspisar transitional,with som primitiv and othr populations unlss
som drivd haratristis of latr Homo. Postranial assoiations ar rit-
ial hr aus ody siz appars to vry diffrnt. Homo hbilis was
vry small odid (35 kg), and H. rudolfensis was lrg (55 kg) (Hnry Arhai Homo to H
1'994|.Saling hk-toothsiz to ody wight shows that thy oth had r-
vrsd th trnd of vr-inrasinghk-tooth siz. Rlativ rain siz x- Larg-ind spistl
pandd, spially in H. bbilis' 600,000 yars ago and in
Asolut rairr siz pands furthr with th apparan ol 17.ergsterlr- Nandrtals dominat E
tus at last 1.8 million yars go, ut ody siz also inrass,so rltiv 3 0 0 ' 0 0 0 t o 4 0 , 0 0 0 y a r si
brain siz apparntly Was not dramatially pandd (l/lkr and Laky H' spis appars 6rst in
1'99).Th arly Afrian form of this spisis oftn rfrrdto as th spis 1999, 2000). Th arlis
H. rgster to ontrast it with th wll-known Asian sampl of H. eretus latr am th hallmark
(.!ood1991; Spoor et a|. 27|. Body size and spially hindlim lngth su-Saharan Afria for
.Walkr
rah modrn proportions in this spis (Ruff and 199). oth .}IBrartyand Brooks 20(
synapomorphis with |ater Homo inlud a furthr rdution in prog- spis sprad out of Afri
nathism, a rdution in th postglnoid pross that is fusd with th tym- gntially swamping lo
pani plat, and rdution in th asolut and rlativ siz of th hk tth. Gol 2007). Th spra
Brains ontinu to xpand, and hk tth get progrssivlysmall during rly (50,000-46,000 yar
th volution f th gnus Homo. aly longd to modrn l
old as 40'000 yars (Bowlt
from Asia who migratd
First Disprsal out of Afria Alaska prhaps 20,000 to
of most of th Amria
Th first apparan of human populations outsid Afria appars to hav Gol t al. 2008). Pop
ourrd soon ftr th apparan of arly Homo in Afria. Fossil Hoo r- thousand yars ago from tl
mains from th Dmanisi sit in th Rpuli of Gorgia ar most likly old .. 300 and Nw Zea|and
than 1'.7 million yars and ar vry muh lik thos found in Afria attrib. 2007; Hunt and Lipo 2006
utd to H' erts/bbilis (Gaunia and Vkua 1995; Gaunia t al. 2000; Thnologial dvlopm
Vkua et al. 2002; Lordkipanidze t a|. 200 5, 2007 ). omo rtus oupied pa, ut it rtainly shows
parts of tropial Asia prhaps s arly as 1.8 million yrs ago (Swishrt al' rud ston tools prsist fol
|994). Th at hronology is still unrtain, ut somtim aftr this and r muh mor rent. Hu]
bfor 0.5 million yars ago some populations hd adaptd to lif in th tm- l s st h n 0 . 2 5 " o f t h i r v
pat limati zon of northrn Asia (Ant6n and Swishr 2001). linge.
An arhai spisof Homo lft tools on th island of Flors, Indonsia,as Th historial distrtuti
arly as 800,000_700,000 yars ago (Brumm t al. 2006; Morwood t l. viopd only in th past 40,
1998)' and in surprisinglyyoung sdimnts(95,000-12,000 yars ago) a nw ezs quippd with Uppr
spis, H. floresisls, has n rovrd (Brown et a|. 2004; Mrwood Afrian homland, first to ,
et a|. 204,2005; orwood and vn Oostrzee 2007).It has a small ody urop (Klin 1999). This
(29 kg)' minut ndoranial volum (380 ), and arhai ody (Larson t al. genti homognity found
2007; Tohri t al. 2'007).It is most likly a dsndantof an arly disprsal with that of our own los
of arhai Homl that undrwnt insular dwarfism (Brown t al. 2004), al- dgr of gnti htrogn
though sm hallng this intrprtation (Argu et a|. 206; Jao t al. that in Homo spies (Ga1
2 0 0 6 ; H r s h k o v i t ze t a | . 2 0 7 ) ' tions to loal onditions ar

l million yars ago) as longing
dy proportions and hip arhit-
nillion yas ago (MHnry and
sitional,with som primitiv and
P o s t r n i al s s o i a t i o n s r
ry diffrnt. Homo hbilis was
sls was larg (55 kg) (Hnry
ight shows that thy oth had r-
) O t hs i Z .R l t i v r a i n s i z
l apparan of H. rgster/er-
dy siz als inrass,so rlativ
. pandd (Walkr and Laky
i is oftn rfrrd to as th spis
own Asian sampl of H. erets
and spially hindlim lngth
(Ruff and !alkr 1,99). othr
a furthr rduion in prog-
ssthat is fusd with th tym-
rd rlativsiz of th hk tth.
l get progrssivlysmallr during
s outsid Afria appars to hav
Homo in Afria. Fossil Hozao re-
of Gorgia ar most likly oldr
lik thos fund in Afria attrib-
kua 1995; Gaunia t al. 2000;
l5,27\.Homo eretusoupid
8 million yars ago (Swishr t al.
tain' ut somtim aftr this and
lns hd adaptdto lif in th tm-
i n a n d S w i s h r2 0 0 1 ) .
th island of Flors, Indonsia, as
mm t a|. 2006; Morwood t al.
(95,000_12,000yars ago) a nw
d (Brown et al. 2004; Morwood
;terzee207).It has a small dy
), and arhai od (Larson t al.
a dsndantof an arly disprsal
jwarfism (Brown t al. 2004), al-
L (Argu t a|, 20Q Jao t al.
Humn uolution 267
Th glogially lat ourrn of suh a primitiv spis of Homo
dmonstratsthe fallay of rgarding human volution as passing through
pogrssivstags.Populations adapt to loal onditions and divrg from
othrpopulations unlss hkd y gn flow.
r i t -
Arhai Horo to Horno spies
x - Largr-raind spis of arhai omo gin to appar in Afria y
600,000 yrs ago and in urasia somwhat latr (lark et a|. 1'994).The
Nandrtls dominat urop and wstrn Asia from approimatly
3 0 0 , 0 0 0t o 4 0 , 0 0 0 y a r s a g o ( K l i n 2 0 0 3 ; K u h n 2 0 0 7 ; K r a u s t a | . 2 0 0 7 ) .
. spiesappars first in Afria somtim for 100,000 yars ago (Klin
1999, 2000). Th arlist tras of Uppr Palolithi tool thnology that
latr am th hallmark of arlist mmrs of H, spies first ppr in
su-SaharanAfria for aout 90,000 yrs ago (Brooks et a|. 19951
Brarty and Brooks 2000). By aout 50,000 yrs ago populations of our
spissprad out of Afria to Asia and thn urop, apparntly rplaing or
gntiallyswamping loal arhaic Homo populations (Anikovih 2007;
Gol 2007). Th sprd Was apparntly rapid in Asia, as vidnd y
early (50,00046,000 yars old) arhaologialtras in Australia that pro-
aly blongdto modrn H. spis populations found thr in sdimntsas
old as 40,000 yars (Bowlr t al. 2003). Amria Was sttld y immigrants
from Asia who migratd aross a land ridg that onntd Siria and
Alaska prhaps 20,000 to 15'000 yars ago' ut th sussfulolonization
of most of th Amrias bgan aout 12,000 yars ago (Dillhay 2003;
Gol t al. 2008). Popl first rahd som of th Paifi islands svral
thousandyars ago from th ast, arriving at th arqusas Islands y aout
.o. 300 and Nw Zea|and y aout .. 1200 (Lum and ann 1998; Finny
2007; Hunt and Lipo 2006).
Thnologil dvlopmnt in human volution appars to rrari in
pa'ut it rtinly shows a pattrn of alration(Klin \999). Relativ|y
rudstntools prsist for ovr 1.5 million yars. Finly workd lad tools
ar muh mor rnt. umans hav had agriultur' itis, and writing for
lessthan 0.25% of thir volutionary dvlopmntas a sparatmammalian
linag.
Th historial distriution of human gnti varition pprs to hav d-
vlopdonly in th past 40'000 to 50,000 yars whn populations of H. spi-
s quippd with Uppr alllithi tool thnology sprad out of thir
Afrian homland' first to Asia and y at last 30,000 yars ago throughout
urop (Klin 1999). This rent olonization ounts fr th rmarkal
gntihomognity found among living mmrs of our spis ompard
with that of our own losst ilogial rlativs, P and Gorill' whr th
dgrof gntihtrognitywithin ah spisis man tims gratthan
that in Homo spil1s(Gagnu et a|. 1999; Ruvolo 1997). Rapid adapta-
tionsto loal onditions and th random snti hanes that n our in
268 Hum uolutio

small isolatd populations hav ld to svral onspiuous phnotypi mammalian avrag (th
hangs, suh as skin olor, hair form and olor, and ody proportions' ut t i o n s k p t u p w i h i nt
ths diffrns appar to rlatively rnt altrations. Th gographi sis (Brown et a\. 24),
distriution of skin olor is losely orrlatd with th intnsity of solar radi- ndowd with modrn r
ation suh that tropial popl in Afria, Asia, and th Amrias hav grat s u j o f r i n r o r g e n
dnsity of mlanin pigmnts in th skin for prottion gainst tissu and aly profound hangs t
mtaoli damag. Highr-latitud popl ar dpigmntdmost likly - othr uniquly human fa<
aus ultraviolt radiation from th sun atalyzs th synthsis of vitamin D ti hang that nald
in th skin and aus of othr fators (Jalonski and haplin 2000). D- haviors ginning aut ]
pigmntation of th skin in high-latitudpopls appars to tru of Nan-
drtals (ulotta 2071 Lalueza-Fo et a\. 2007). Body proportion varis
gographially aording to tmprtur' so that popl in hot limats hav volution of Humar
longr lims and lss gloular trunks to allow gratr hat dissipation
through th skin (Ruff 1994). ptions to this pattrn an oftn - Bipdalism is th first mz
plaind y th prsn of ultural daptations that allow prfrntial sur- first hominins wr mos
vivl and rprodutiv suss indpndnt of gnti daptations. Th first odis without th spi
H, spis in urop, fo xampl, had ody uilds haratristi of tropial mark of our linag (Tho
popl (thy am from Afria), ut thy had thnologis far suprior to possily uropan radiat
thos of th loal Nandrtls. Thnologial innovation is oftn th rason win's wods, ..to liv som
..a hang in its
for population xpansion and plays a pofound rol in dtrmining why r- mannr t
tain physial varitis of H' spiens oupy gographi rgions and displa ditions of its native oun
arlir indignous popl (Diamond 1997). th human lad proa
volutionary novltis. l
that podud th ipd
volution of th Human Brain dfin th human grad
transformations of th tr
Th aumulation of volutionary novltis through tim rvals th ontinu- fossil rord. It may t
ity of human volution. Th dmonstration of this ontinuity is a triumph of rats so that th lngth of
volutionary iology. It furthr onfirms Darwin,s viw of dsnt with mod- of th hallu. h xtin
ifiation. Th pansion of th rain in th hominin linag is on of th most pithus appars to hav l
dramati dmonstrations of this. Absolut and rlativ rain-siz xpansion big to (Kohlr and
follows tim rmarkaly wll, as shown in al 1. Th arlist speis for rmidus will hlp larify <
whih rain siz is known, . fresis' has an avrag ndornial volum ipdlity in our linag.
of 48 (asedon 4 spimns),ompard with modrn himps at 400 ' arly volutionists, rr
gorillas at 500 ' and H' spis at 1,350 . Th ndoranial volum for . akl (1868), prdit
grhi \1 spimn) is rportd to 450 , for . fricus (7 spimns)' rain. How' why, and i
4 5 2 , f o r P . b o i s i ( 1 s p i m n,)5 2 1 c c , f o r P ' r o b u s t s( 1 s p i m n ) 5' 3 0 uniqu ostur is a favor
, for H. hbilis (6 spimns), 6|2 c, for H. rudolfsis (1 spimn), 752 2004). It is possil tha
' and for . ergster ( spimns), 871 . Howvr' at last on spisof ipdality, givn th fat t
Homo (H. |loresislsl managd to surviv on a rmot island until 18'000 wih folim suspnsio
yars ago with n ndoranial volum of only 380 (Brown et a|.204). ipdal, this ody plan a
Brains epand during th Plistonfrom 914 twn1.8 and 1.2 mil- quadrupedal mammals
lion yars ago to 1,090 twn550,000 and 400,000 yars ago' 1,186 c (Rodman and Hnry l
btwn 300,000 and 200,000 yars ago' and aov 1,300 aftr 150,000 Thr has n a pul
yars ago (Ruff t aL. 1998). Rlativ to stimatd ody wight, th fossil primitiv traits rtaindin
spisgo from slightly aov th siz sn in modrn aps to 5.4 tims th On pol mphasizsth 1

svral onspiuous phnotypi
olor, and body propotions, but
nt altrations. Th gographi
d with th intnsity of sola adi-
\sia, and th Arrias hav grat
for prottion girrst tissu and
, ar dpigmntdmost likly e-
,talyzsth synthsis of vitamin D
lalonski and Chaplin 2000). D-
oplsapparsto tru of Nan-
I. 2007). Body proportion varis
;o that popl in hot limats hav
o llow gratr hat dissipation
j to this pattrn an oftn x-
ations that allow prfrntial sur-
rt of gnti adaptations. Th first
dy uilds haratristiof tropial
' had thnologisfa suprior to
ial innovation is oftn th rason
.oundro1in dtrminingwhy r-
ly gographirgions and displa
ls through tim rvals th ontinu-
n of this ontinuity is a triumph of
)winviw of dsntwith mod_
: hominin linagis on of th most
a n d r l t i v r i n . s i z x p a n s i o n
in Tal 1. Th arlist spis for
las an avragndoranial volum
dwith mdrn himps at 400 '
l . Th ndoranialvolum for .
, for , frins (7 spimns),
, for P, robustus (1 spimn)' 530
H' rudolfsis\I spimn), 752
. Howvr, at last on spis of
v on a rmot islnd until 18,000
only 380 (Brown t al. 2004).
> m 9 t 4 t w n1 . 8 a n d 1 . 2 m i l -
) 0 a n d 4 0 0 , 0 0 0 y a sa g o ' 1 , 1 8 6
, and abov 1,300 aftr 150,000
' stimatdody wight, th fossil
n in modrn ps to 5.4 tims th
Hum L'olutio 269
mammalianavrag (th olumn lald V in Tal 1). Not all plpula-
tinskpt up with rin pansion, s is dramirtiallyshown H. flrlrsien-
sls (Brown t al. 2004), ut all tant populations of H. spies ar fully
ndowdwith modrn airrs.Unfortunatly th fossil rord is mut ()n th
sujtof rain rorganiztion,whilr is pity usthr wr prsum-
aly profound hrrgsthat aompnid th dvlopmnt f sph and
othruniquly hurrranfaultis.A strong i1san mad for a major gn-
ti hangthat nald H. spis to spak nd prform modrn human -
haviorsginningaout 50,000 yrs ago (Klin and dgar 2002).
volution of Human Walking
Bipdalismis tlr first major grad shift in hominin volution, r.rtth vry
first hominins wr most likly gnralizd hominoids with rathr plik
odiswithout th spializationsfo ipdlism that latr am th hall-
mark of our linag (Thorp et a|. 2007). Thy wr part of th Afriirn and
possiblyuropan radiation of lat ion hominoids that am, in Dar-
..to liv
win'swords, somwhat lss on trs and mor on th ground'' du to
..a
hangin its mlrn of prouring susisttl'or to a hang in th on-
ditionsof its nativ ountry'' (Darwin 7872, p. l.]5)' h fist memrs of
th human lird proaly will rognizd in th fossil rord y sutl
volutionary novltis. Th major gnti/dvllpmntaltransformtions
that produd th ipdally adaptd trunk nd hindlinr morphology that
dfinth human grad proly appard latr. Th pis squn of
transformations of th trunk and hindlim may nvr known from th
fossilrord. It may that parts of th ipdal ody hangd at diffrnt
ratsso that th lngth of th plvi lads rdr'rdarlir than th addution
of th hallux. Th til-ltand distintl nonhonrinin ione ape oreo-
pithusappars to hv had a rdud plvi lngth but rtirrda divrgnt
ig to (Kohlr and Moyi-So|' 1997). lr full analysis oi rdipithus
rmiduswill hlp lrify our undrstandingof th natur of th traI-rsition to
bipdalityin our linag.
arly volutionists, inludirrg Lmark (1809)' Darwin (7872\, and
akl (1868)' prditd that ipdality prdd the pansion of th
brain. How, why, arrd in what haitat an aplik anstor adoptd this
uniqupostur is a favorit topi of spulation (Kingdon 2003; Mnr
2004)' It is possil that many aplik spis in th ion zrdoptd
bipdality'givn th ft that th ap bodv pln is daptd to nroving in trs
with forlim sr-tspnsion and not to f6int quadrupdal walkirrg. on
ipdal,this body plan an mov with an ngti ffiiny qual to that of
quadrupdalmammals and muh gratr than tht of quadrupdal apes
( R o d m a na n d n r y 1 9 8 0 ; S o k o l e t a L .2 7 | .
Thr has bn .r puliar polarization of viws aout th maning of th
primitivtrirs rtaindin th postrni of rly spisof ustrlrlpithus.
on pol mplrasizsth bipdal spilizations(Lovjoy 200.5,2005), and
270 um uoltio
ar uniquly Homo (Bra
th othr pol lls attntion t() tl.l many primitiv haratristis (Strn
tivly nlagd hip ioints
2000). Viwd in gand prsptiv,th divrgntintprtatil1s an sn
haps rlad ro long-dista
as diffrns irr mphasis. Both amps agr that all spis of AustrIo-
H' spies. Th disover
pitheus wr ipdal. hat implis tht thsspisdid not lim lik aps.
volution in that it has a ]
Thy did, howvr, rtain morphologia1 fatursssoiatdwith arboral.
rlativ forarm lngth (l
ism for at lst :r rillion yars nd a hip arhittudiffrnt frol that of
latr spisof Homl. That implis so diffrnfro-tmldrn humans in
gait and linring ility, ur thy did not walk :lnd lim lik ps.
Th volution of Hr
Th publishd hominin fossil rord dos not yt hav a tru intrmdiat
stag twn an aplik and a humanlik ody (Hnry and Jones 2006I.
xpt for rlativly sma
Th postrniur-lrof ustrll1lithesrtins mally primitiv aplik traits
lar to thos of many of th
ut is fundamntally rorganizdand highly spializdfor ipdality,unlik
of our sufamily livd il
that of any ap (Hnry 1986). hr is a profound altratiorriIl th gn-
thir dits wr simil l
ti tmplat that produd th short plvi lad, addutd hallu, and othr
limaxing with th appa
ipdal traits of this hominin.
ago), th hk teth 1
Th diffrn twn Astrlopithes'spositional and loomto -
Although Darwin and his
havior nd that sn in latr spisof Homo ;rmlarrwith th disov.
fossil rord would vnt
is of postranil fossils assoiatd wit . rgtstr/erets(]il/illrkrand
..StrppingYoth'' (KNM WT 15000). Thr minin nrgdonrsA . fri
Laky 199|, partiularly th
rlativ to ody siz. But
ar striking diffrnstwn th plvis and fmur of all spis of us-
molars with onomitnt
trIopithus and those of H. ergster/rets,with th laftr apparing
hks, and attahmnt :
muh mor humrrnlik (Mnrv and offing 2000). This proal,vrgistrs
proimat trend throug
a nrajor shift in adaptation twl-lstrlopitbttsand Homo. From this
hk-tooth siz from sn.
prsptiv'ustrlopithe,/sapprs t :l ipd with fr hirnds ftlr arry-
ntttsisnd . frcttsis
ing ut st daptd to short distans and rvith a halthy ppriation of
hug in P. boisei and P.
trs for safty, fding, and slping. Th longr fmora and mlr human-
so that from H. bbilis at
lik plvs tht appar 1.9 million yars a!]o wit omo mark th gin.
siz oms progrssiv
ning of an impotant hang.
l a t i v h k - t o o t hs i z
From th urrntly availal fossil vidn' thppar to e two grad
mgadorrtiaq l u o t i n ti s s
shifts in th postranial volution of th hurrrzrrrody. Th first Was th tran-
lar and first two molars
sition from gnralizdhominoid to hominin ipdalism that is doumntd
spis od wight. Ir is
in . msis' . fresis' . frius, and . grhi (Ward 2002). Th
rl indition of rlativ
diffrnsin postrnial morphology among thsspisar diffiult to as-
m g a d o n t i a t h o u g ht i m
sss usof tlr inompltnssof th fossil rrd.All shar ]rivdfa-
t o p r o s s g a t rq u a n t i
turs with H{lnll, ut onl1' . zlfresisarrd . frius ar omplt nough
rvrsl of th trnd in m
to mak dtild ompaisons. [n som rsptsthsspisar rmarkaly
to oth ditary hang (i.
similar rlativ to othr spis of Hominoid;r. For xampl, thy shar a
orporation of mr rna
similar mi of aplik, humanlik, and uniqu fatursin thir wrists, hips,
for mastiation.
and kns (MHnry 1986). Th,vapparntly diffr in th ratio of forlimto
hindlim joint sizs,howvr, with ' frinus ppaingto mo :rplike
vn though it is latr in tirrrnd mor Holo-|tke in raniodrrtalfaturs
(N4Hnry and Brgr 1998; Grn et a|.207). Both spisapp to shar
Rmaining NIystri
omination of spializdipdal traits, ut not xatly lik thos of mod-
Th divsity of living m
rn humans, in faturs of th forlim ssoiatd with liming (Strn
i s h d l a t i v t o w h a t i t r
2000). Th sod grad shift in postranial volution took pla out
it is lar tht thr wrr
1.9 milliln yars ago with tlr 21pparan lf rnr.rhmo humalik hips that
 Hum uolution 271

ar uniquly Homl (Braml and Lirma 2004)' Long fmoa and rla-
iy primitiv hartristis (Strn
tivlynlargdhip joints mark a signifianthang in loomotion tht is pr-
v r g n itn t r p r t t i o nsa n s n
haps rlatd to long-distan, ffiint striding mor lik that sn in modrn
rgrthat all spis of ustrIo-
H. spies. Th disovr of ' grhi revea|sth omplity of postranial
tsspisdid not lim like aps.
volutionin tht it has a humanlik ratio of fmur to humrus ut an aplik
fatursssoiatdwitlr aroral-
rlativforarr lngth (Asfw et a|, 1999).
arhitturdiffrnt from that of
diffrnfrom modrn humans in
walk an]lim lik aps.
s n()t ,ytlrav a tru intrmdiat
Th volution of Human astiation and Dit
ody (Hnry arrdJons 2006).
pt for rlativly small anin tth' th arlist hominins had tth simi-
ains mar-ryprimitiv aplik traits
lar to thos of many of th ion aps (Ungar 2007). Ths arly mmrs
ly spilizdfor ipdality, unlik
of our sr'rfamilylivd in woodland haitats (Pikford 2006). Prsumaly
a profound itration in th gn-
thirdits wr similar as wll. But ginning aout 4 million yrs ago and
: lad,ddutdh[lu, and othr
lima-ingwith th apparanof Prthropus boisi (2._1.4 million yars
ago), th hek tth pand through tim (onstantino and .il7ood 2007).
s's positionl and loomotor -
'zo anrlarr rvith th disov- AlthougDarwin and his ontmporarisprditdmuh of what th human
fossilrord would vntually rval,no on antiiptd th disovy of ho-
h H. rgstr/rttts(Walkr and
minin mgadonts. Afrian aps and modrn humans hav small hk tth
Youth'' (KNM $T 15000). Thr
rlativto ody siz. But rly hominins had rlativly hug molars and pr-
s and fmur of all spis of us-
'rtrrs'nith h lett pping molarswith ono-litantlygiganti jaws, alvolar bon, uttrssdfa and
hks'and atthmnt aras for th hwing muslatur. Thr is n p-
:fing2000). This proaly rgistrs
proimat trnd though tir in sussiv spis of inrsing rlativ
rlopithezusad Homo. From this
hk-toothsiz from small in rdipithecus rmidus to modrat i . -
a ipd rvith fr hands for arry-
mnsis and . frsis, ig in . fricnus, P. robustt'ts, and . grhi, and
nd with hlthy appriation of
hug in P. boisi and P. ethilpius. Th trnd rvrss in th Homo linag
llngr fmora anl mor human-
so tht from H. bbilis and H. rudolfnsis to modrn H. spis, th rlativ
s ago with Homo mark th gin-
siz oms progssivly smallr. On way to xprSs ths trnds is with
rlativhk-tooth siz, as is shown in th last olumn of al 1. Th
n'th:lpparto two grad
rngadontialqr-rotintis simply th asolut siz of th lowr sond pmo-
uman rrdy.Th first ws th trn-
lar and first two molars dividd y thir ptd siz drivd from th
inin bipdlismthat is doumntd
spisody wight. It is not intndd to pris ind ut mrly a gn-
zs' and . grhi (.!fard 2002). Th
ral indiation of rlativ siz. It is rasonalto assum that th inras in
ong thsspisar diffiult to as-
nrg;rdontia through tim is rlatd to ditay spializations(i..,th aility
fossiIrord. All slrar drivd i_
to pross gratr quantitis of low-nutrint and tough-txturd foods). Th
d . fricus ar omplt nough
rvrs;rlof th trnd in mgadontia in th volution of omo may rlatd
sprtsthsspisar rmarkly
to oth ditary hang (i..,lss lian on tough-tturdfoods nd th in-
noida. Fo ampl, thy shar
orpotionof ror mat in th dit) nd th prossingof foods with tools
niqu fatusin thi wrists, hips,
bfrmstition.
rtly diflr in th atio of forlim to
us apparingto mor aplik
H om o-|ik in raniodntal faturs
Z007).Both spisappar to sha
Rminingystis
i' ut not atly lik thos of mod-
h divrsityf living mms of th suprfamily ominoida is impovr-
assoitdwith liming (Strn
ishdlativto wht it was in th past. vn with th limitd fossil sampl,
ranial volution ttlok pla alut
ir is l that th Wre many kinds of aps and hunrns long ago. or
of muh mor humanlik hips that
272 umlt uolutio

sis await disovry. Thr wr proaly many volutionary xprimnts Argu' D., D. Donlon,
in th varid haitats of th Afin ion to liston (Suwa t al. irophali',g
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Arnason,U., A. Gullrg
pression that thr wr only a fw hominin linags (.!fhit t al. 2006), it
of pimatdivrgn
far mor likly that our family tr will turn out to quit ushy (Sarminto o f m t l d nh u m . r l l s
et a|.2007). Spisnams may nd to multiply to aommodat th divr. Asfw, B., . !7hit' o. I
sity, although a alan nds to maintaind twn xssiv splitting Austrlopithusgt
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Bgun,D. R. 1994. Rlt
It will b wondrful to rsolv th qustion of knukl-walking in human
intrprttions as
volution(Rihmond and Strait2000,200l; Rihmond t al.2001). Living
Athropolog37: 11
spis of th Afrian ld of Hominoid inlud thr knukl-walkrs B g u n ,D . R . , n d . V . \
and a ipd. Th odds ar that th first hominin Was also a knukl-walkr, nw iddl N{ion
ut th arlist hominins known so far ar ipds and limrs without sp- Bhrnsmyr' A. . 2006
ializations for knukl-wlking. Prhaps thr funtionl and dvlop- Bowlr,J. ., H.Johnst
nd N. A. Spoonr.i
mntal prosssthat pdispos larg-bodid aps to om knukl-walkrs
hangsat Lak ur
whn thy adapt to haitats that rquir mor trrstriality. If so, that appar- Braml,D. ., and D. E
ntly odd gait may have volvd in paralll irr two o more linags. of Hotl. Ntur 43
Thr may hv n svral pansios of populations out of Afria dur- Bromag,. G., and F. S
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am xtint. It is sdutivto ir-rragin Boks,A. s., D. . Hlg
only on triumphnt puls of mi.
R . G ' K l i n ,W . J . R i l
gration that sprad humanity for th first tim throughout urasia' Th
J . V n i s' n dJ . .
sprad of H. spies appars to hav happend in a short amount of gologi- sitswith on point
l tim' ut rhai spis of Hlmo wr poaly muh lss adaptabl B r o m ,R . l 9 . ] 8 . h l
and mor vulnral to th svr limati shifts that ourrd vywhr 77-379.
ovr th past svralmillion yars. Loal xtintions may hav bn ommon Brown,P., . Sutikna,.
R . A . D u .2 0 0 4 .A n
throughout th hanging gographi rang of arly hominins. Bfr 50,000
lors,Indonsi.Na
yas ago humirns Wr an insignifiantlysrall part of th vrteratfauna. B r u m m , 4 . ,F . A z i z , G . D .
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modifiation. It maks it vry unlikly that, for xampl, signifint n. A. Bauvilain,. Blor
J . D j x , . D n v s ,P
phaliztion gan for th volution f ipdalism in ou linag.n.
D. Graads,T. Lhm
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C h n ,F . - C . ' n d . W . - H L. i
lLIoGRAPY Hominoids and th {
and himanzees. m
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y m a n y v o l u t i o n a y P r i m n t s
n to Pliston (Suwa t al.
il hominins lads som to th im-
r lings(!7hit t al. 2006), it is
out to b quit ushy (Srminto
rltiplvto aommodat th divr_
aind twn ssiv splitting
orr of knukle-walking in human
1 ; R i h m o n d t a l . 2 0 0 1 ) . L i v i n g
a inlud thr knukl-walkrs
minin was also a knukl-walkr,
ipdsand limbrs without sp-
:hrr funtional and dvlop-
.d apsto om knukl-walkrs
lr trrstrility.If so, that appar_
in two or mor linags.
of populations or'rtof Afria dur-
ulatd pats of urasia and thn
nly on triumphant puls of mi-
it tim throughout urasia. Th
lndin a short amount of gologi.
r proaly rnuh lss adaptal
: shitts that ourd vrywhr
:tintionsmay hav bn ommon
of arly hminins. Bfor 50,000
mall pat of th vrtratfuna.
lf pssimismor optimism. Th ho-
t,ut it is also rih and onsis-
t Drwin's viw of dsnt with
hat, for ampl, signifiant n-
'f ipdalismin our linag. on.
, u t d
f t h s i n
rt ids ar groundd in aurat
is losi.
' D. Graads,D. Rd, and
t skltorrfrom Dikik, thiopi.
: in astrnurop and impliations
: 3 | 5 : 2 2 3 _ 2 2 4 .
ln lfrirni]apity in Asin Hono
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\Whit,T. D.' G. \foldGaril, B. Asfaw, S. Amros' Y. Byrr,R. L. Brnor,
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. F-.Howll, R. T. Kono' T. Lhmann, A' l,ouhart, O. . Lovjy, P. R.
Rnn, H. Sagusa'. S. Vrba' H..Wsslman,and G. Suwa.2006. Asa Issi,
Armis nd th origin of ustrlopitbs.Ntur 440: t]83_889.
WoldGaril' G., Y. Hail-Slassi,P. R. Rnn, W. K. art, S. ' Amros, B. Symptoms
Asfaw, G. Hikn, and T. Whit. 2001. Gology and palirot-ttlogy f th Lt
Mion iddl Awash vally, Afr rift, thiopi. Ntur 412 1'75_1'78.
A young oy in Kans;
!7oldGaril, G., T. D. Whit, G. Suwa, P. Rnn, J. d Hinzlin, W. K. Hart, nd
G. Hikn. 1994. ologial and tmporal lilmnt of arly Plion hominids rash with fvr, swolJ
at Aramis' thiopi' Ntur 371: 330_33. hk rvals tht th
Wolpoff' I\4.H. |971, tric Trds i Htlmiid Dtl ulutil. lvland: s (hikn po) and ha
lZstrn Rsrv Univrsity Prss. Th family sldom tr
1982. Th Arago dntal sampl in th ntt of hminid dntal volutiorr'
halthy' if slightly ov
In H. d Lumiy, ed., L'Homo rtus t l pl d l'homm d tul prmi
ls homiids flssils,389-4l0. olioqu Intrntionald ntr Ntional d
hait, howvr' of k
la Rhrh Sintifi.Ni: Louis-Jn Sintifiarrd l-itrary Pulitions. a young lak-taild
.Vood'
B. 199|. Koobi Frlr Rsrb Proit.o|' 4, Hlmiid ril Rmis. of groundhogs found ,
Oxford: larendon Prss. and prairi dogs ar k
Zollikofr, . P. ., M. S. Pon d Lon, D. . Lirmn, F. Guy, D. Pilam,
and tularmia (rai
A. Likius, . T. kay, P. Vignaud, and M. Brunt. 2005. Virtual rnial
nods that ould indi
rnstrution of ShlthroDs tchdsis. Ntttr 44: 7 5 5_7 59 .
a diffrnt dirtion.
As it happns' th
hiago, whr it
ulats in wild animal
a los rlativ of th .
a shipmnt of Gamiz
prairi dogs had n
link in th hain of i
forgd (D 200; G
though his lovd pl
halth workrs ar tr(
munity against monk
Th ase of this par
monkypox outrak
sons who am in on
pt faility (D 200:
loolltionof ust r lop ith us
PhsiIAthrrlpoklg 45

)l. tlpho|gof Austrlolitbeus

I1x..I()urlof l"-ttllutirl

:volution with artiular rfrn t.l

lplsiul o Qutitt it, B illog
volutionaryBiology of Disas
istorti<ln? Si 299:.|994_1997. and Darwinian diin
strlo1lithusrmids, a nw spis
t u r 7 1 : 0 6 _ | 2 .
.midus'a nw spis of arly
'5:88.
ihel F. ntolin
lbrs, Y. Bvne,R. L. Brnr,
il-Slssi, W. K. Hart, L. J. Hlrrsk,
Luhirrt,O. . Lvjoy, P. R.
Lan,antl G. Suwa' 2006. Asa Issi,
N t u r a4 4 0 : 8 8 3 - 8 8 9 .
,W . K . H a r t , S . H . A m b r o s ,B . Symptoms
ology nd palaontollgyof th Lat
' h i o p i a .N t u r 4 1 2 : | 7 5 - 1 7 8 '
A young oy in Kansas, in th midwstrn Unitd Stats,dvlops a svr
:rn,J. 1inzlin, W. . Hrt, nd
plamntof arl Plior.rhominils rash with fvr, swolln lymph nods, and approahing dlirium. A quik
hk rvals that th oy lrs survivd th typial hildhood inftions
;d Dtl urlltir. lvland: Cls (hiknpox) and has up-to-dat vainations and no history of allrgis.
Th family sldom travls far from hom, and all its mmrs appar to b
lonttof hominid dnt1volution.
halthy,if slightly ovrwight in th typial Amrian way. This family has a
1lletl l.hlmm d t'ttulpni
lntrrriltilnal
hait,howvr,f kping intrstingpts nd for his irthday gav th oy
d rrtrNationaI d
i l l t i f i a n d l , i t r a r v[ , u h l i i o n s . a young lak-:rildpraii dog, on of thos loval, smallr-sizedrlativs
/o|.4. Hrlmiid ril Rmts. of groundhogsfound on th Grat Plains. This pt was apturd in th wild,
and pairi dogs ar known to arry srious inftious pathogns lik plagu
Librman,F. Gu, D. Pilm, and tulrmi (rabit fvr). Th oy's symtoms inlud swolln lymph
JvI.Brtlnt.2005' Virtul ranil
nodsthat ould indiat plgu or tular-ti,ut th pstular rash points in
is.Ntur 44: 7 5 5-7 59 .
a diffrnt dirtion.
As it happns, th suspt prairi dog m from a pt faility nar
hiago,whr it ame th sour for morrkypo virus that natually ir_
ulatsin wild nimals half th world away in wstrn Afria. Th pathogn,
a los rlativ of th dradd smallpox Vriol, ntrd th Unitd Stats in
a shipmntof Gamian rats irnportd for th oti pt trade. Th rats and
prairi dogs had n housd togthr in a warhous, nd thr th final
link in th hain of inftin from Afri to th Arian idwst was
forgd (D 200; Guarnr t l. 2004), Aftr tratmnt th oy rovrs,
though his lovd prairi dog dos not. Th rst of th family nd loal
halthworks ar tratd with smallpo vin aus it stimulats im-
munity agair-rstronkypox as wll.
Th as of this prtiular family is prt fition ut is asd on an atual
monkypo utrak in six midwstrn statsin 2003 that inftd 7I per-
sonswho am in ontat with Gamian rats or priri dogs from th sam
ptfiiity (CD 2003). t |0% as fatlity, monkypo is not zlsdadly as

281
282 uolutilr Biolog of Disse d Dr,ii ediie a
smallpo, ut th rsmlanof th monkypo virus to its ousin, whih is mismth wn th
widly livd to hav n radiatd from th wild by a wrldwid vai- volvd and adaptd'
nation ampaign in th 1970s, is frightning.Th as also rvalshow mod- t w n h i g h f i n s si n t |
rn mdiin and publi halth us th framwork of volution to solv Ns 1991; Swyngh
potntial halth riss.This has n alld Darwinian mdiin (so) - hart disas (Swyngh
aus diagnosis and tratmnt dpnd on mthods of volutionary iology to and prosttanr ((
sluth th undrlying auss of immdiat symptoms (\/illiams and Ns fts (Glukman and
1991'; Stearns 1999).Inftion with monkypox was onfirmd y viral iso- hv nprimry r
lation' pidmiologial invstigation,and finally DNA analysis, and family Th rnt worldwir
rsmblans among po viruss trad this partiular virus k to Afria. a m p l t h t p r v i o u ss t
A similar story an told aout th oronavirus that auss svf aut gns that aus dia
rspiratory syndrom (SARS) and sparkd a puli halth pani in ality, howvr, fw a
2002_2004 whn inftions sprad from hospitals in Asia to North Amria. l a t i o n t o d i s s :t h
In a mr ight months gnti nalysis of th nwly mrgd SARS virus first g l o i n ' a n d m l a i r
ld to its idntifiation, thn to intrmdiats in ivt ats sold in wild-gam on, nd th story dl
markts in hin (Ruan t al. 200), and vntuallyto potntial rsrvoirsin populations (Hill and
a t s ( L i t a l . 2 0 0 5 ; . W . a n g t a | . 2 0 6 ) .o n a t r a g i a l l ym u h l a r g rs a l t h and inftious disas
human immunodfiiny viruss (IVs) that aus AIDS hav also n a s p t so t h n v i o
trad by gnti analysis to wild primat populations in Afria (ahn t al. immun rsponss .n
2000; Klet al. 2006). l 9 9 9 ; W i s sa n d M M
s o n 2 0 0 6 ) . F i n d i n gt h t
Although I aknowl
tiology tinary for tht |
m d i i n t h a t r q u i r
Darwinian mdiin sarhs for th volutionary origins of siknss-thos d p t i v i n h L r m n si
things that mak lif nasty, rutish, nd short. Darwinin mdiin suds off th world of m
y looking for disas origins in iologial adaptations of humans and th al sltion ut lso h
miros that plagu thm-thos traits that allow sm to surviv and r- of ommon dsnt fr
produ bttr than othrs. On asi tnt of Dawinian mdiin oms 1999iBuIl and !ih
from adaptationist thinking, that disassaris aus humans ist in a r m a I a d p t i v t o h L
Disas, howvr, is l
ial wold in whih a.
Darwinian mdiin dsribs n volutionary approah to aring for idal ut ar rlativ tr
the sik, and for publi halth in gnral.In this sns,Darwinian md- i t i s .T h i s W s t h p o i
iin looks to th adapttions' and maladaptations' of oth humans and y Gou|d and Lwon
th miros that aus diss.For miros in partiular' Darwtntan stan' W an undrs
mdiin must aount for rlativly rapid gnration tims of patho- sns, in th adaptiv
gni miros, and high rats of mutation and gnti rominatlon tm upright t th to
that lad inxoraly to novl adaptations and th mrgnof nw dis- movmnr that dos l
ass. Both pathogn and host, howvr, also must undrstood in why w hv svnvt
trms of volutionary history' homology, and phylognti onstraints Th answr ro rhis qu
on adaptation. Drwinian mdiin, thn, taks a dynami viw of oth and vrtbrats and in
inftious and gntidisass,whr svralvolutionary historison- inhritdfrom our an.
vrg within individal patints on th on hand, and in human popula- adaptiv nmr in th
tions on th othr. passdOn to Us as i l
and Lwnttn 7979).
 lttdii
:ypovirus to its ousin, whih is
th wild by a worldwid vai-
;.Th aslso rvls how mod-
ramwork of volution to solv
Dawinianmdiin (so) -
thodsof volutionry iology to
syptoms (\/illimsand Ns
ypo was <rnfirmdy virl iso-
inally DNA analysis, and family
s partiularvirus ak to Afria.
rnavirus that auss svr aut
kd a puli hlth pni in
'spitalsin Asia to North Arnria.
r nwly mrgd SARS virus fist
s irr ivt ats sold in wild_gm
n t u l l yt o p o r n t i lr s r v l i isn
a tragiallymuh lrgr sal th
tht us AIDS hv also n
looulationsin Afria (Hahn t al.
tionary origins of siknss-thos
ort. Drwinian mdiin suds
l adaptationsof humans and th
rt allow some to surviv and r-
lt of Dawinin mdiinolns
aris aushumans xist in a
ionary approah to aring for
. In this sns,Darwinian md-
laptations,<lfoth humans zrnd
: r o si n p a t i u l a r ,D a r w i n i n
pid gnrationtims of pato-
'ion and gttiromination
s and th mrgnof nw dis-
:r, lso mllst undstood in
y , a n d p h y | o g n t i o n s t r a i n t s
l, taks a dynami viw of oth
, l v o l u r i l n a rhy i s t o r i s o n -
n hand, and in human popula_
Llolutionry'Biologl, of Diss d Drii dii 28.]
mismath twn th urrnt world and th world in whih our spis
voivdand adptd, and that lrumans sr.rffrfrom sris of trd-offs -
twnhigh fitnss in th past and rdud fitnss irr th prsnt (.Williams arrd
Ns ]991; Swvnghdar'rw2004; Glukrn and Hanson 2006). Ositv'
hartdisas(Swynghdauw2004), tpe-2,diats (Dianrond 200), rst
and prostat anr (Gravs 2002\, goitr' iodin dfiiny, and irth d-
fts(GlLrkmnnd Hnson 2006), nd aging (\Williamsand Ns 1991)
havn primary targtsof this kind of thinking.
Th rntworldwid inrasin typ-2 diatsprovids a omplling -
mpi that prr,ious sltiorrflr tolrating starvzrtion rrrily hav sltd fo
gnsthirt aus diatsin a food-rih nvironmnt (Diamond 2003). In r-
ality, horvvr,fw ass doumnt ongoing natual sltion in humans tr-t
rlation tl disas:th s of sikl-ll anmizr,low-fur.rtioningo hmo-
gloin, and malria rsistanin su-Saharan Afria is ssntiallv th only
on,and th stor dos not hold up gnrallv outside a fw wll-doumntd
populatins(Hill and tulsky \999). As will showrr latr' oth gnti
and inftiousdisasshav multipl auss that inlud nutrition and othr
sptsof th nvionment' gnti sLlsptiility,mrvoni dviopmnt,
immun rsponss' and pathogrr divrsity (Burn and \/hit 19721 Stearns
1 9 9 9 ; s i sasn d M i h a l 2 0 0 4 ; G r a h a r n t a l ' 2 0 0 5 ; G l u k m a n a n d H a n -
son 2006). Firrding th aussof disasrmairrsa diffiult hallng.
Although I aknowldg that natural sltionis th most powrful volu-
tinary for that lds to adaptation, I t:rk a diffrnt tk for Dirrwinian
mdiirrthat rquirslss spulatilnaout whirt may or may not hv n
adaptivin humns in th past. vn so, volutionary iology has muh to
offrth lvorld of mdiin via undrstndingnot onlv adaptations y ntu-
ral sltionut also hrologis,traits tht ar shrd y organisrnsaus
of ommon dsntfrom thir anstors (Williams and Ns 1991; Starns
1999;Bu|Iand Wihmn 2001). larly oth gnti and irrttiousdisss
ar malildaptiv to humns in that moridity and mortality rdu fitnss.
Disas,lrowvr,is an insapalonsqunof an vr.hngingiolog-
ial world in whih adaptatior-rirrrdfitrrssr not asolr.rtly tund to sorn
idalut ar rlativ to what might our in a world of onstraind possiil-
itis.This Was th point of th farous ritiqu of th adaptationistPOgln
by Gould and l,wontin 0 yars ago (Gould and Lwontin 1979). For ir-r_
stan'w an undrstand th svn vrtra in our nks in a funtional
sns,in th daptiv tra.l-offs tlr:rtkp our orry skulls :rnd nrvous sys-
tm upright at th top (font) of our odis whil mainti.rining11iilityof
movmntthat dos not damag our spinal ord. But this dos not tll trs
why lv lrv svn vrr'rot six lr ight (Williars rrd Ns 1991).
Th answr to this qustion rsids in our volutionary history as hordts
and vrtratsatld in th dvlopmntalProgram nodd in th DNA w
inhritdfiom tlur anstors.Svn nk vrtr:rmay lrv rpresntdn
adaptivnurr in th past, ut this aus is lost in history, and th rsult is
passdOl1to us ils a plrylogntionstraint of our ody plans (s Gould
nd Lwontin |979\.
284 uolutkr Biolog of Diss d Drii diie t

Th viw of Darwinian mdiin I spluslnsidrsth volutirnof dis- A p r i m a r y x a m p | i

as as a srlsof ongoing vntsin populatins that xprinoth naturl immun rsponssvia l
sltionand rand<lmfors lik mutation within a framwork of phylogn- familiar with annuai fl
ti onstraints. In othr words, I wnt dotors to gin thinking lik popuia- Spanish flu of 1918-19
tiorr iologists, lik tlros of us who osss aout variabl populatilns of outbraks ou worldr
orgnisms, thy humans or thi miros, in an ologial sphr that vr-hanging surfaa
hngs in tim nd spa (Burnt and \/hit 19721Lederrg 2000; w. An- tions and nssitat
drson 2004; Purssll 2005). F.urthr,mdial sin will gratly nfit gson t al. 2003). In
frm thinking of popultions at svral lvls,from human populatilnsthat strands od for intr
inhait on pl or anthr to populations of miros that liv within on tin psul, polymra
tissu or anothr within patints (l,vin t al. 1999; Starnsand rt 2001; protins that at as an
Grnfll et aL.204).l gin y looking t two asptsof th urrnt human hmagglutinin (HA) for
ondition with rspt to disas:human plpulation siz and volutin of invasion into th host
patho!]ns,and human population siz rlativ to our own gntis and formd partils sap
mutations. ially pron to muttio
antigni subtyps witt
shor irds, and poultr
ADAPABlL|Y oF PAHoGNs
flunza A viruss, hav
Lik othr parts of ntur' humans rprsnt a rsour fo miroial et a|.2006). Th urr
growth. With th hlp of modrn mdiin th human popultion has grown H5N1.
fourfold in th pllst ntury' surpessing 6 illion in th past dad, and hu- \ithin ah subtyp
mans n()W onstitut a sustantial part of th plant's iomass and us a antigni drift in that l
lrg fratitln <lfth plant's ologial outpllt (Vitousk t al. 1986; Daily lss asily rognizd;
t al. ] 997). o a larg tnt' human populations hav pandd aus natd individuals (Bus
vainatiorr arrd antimiroial drugs prott us from th miroial world, tions for human halth
with th dirt nfits of lowrd infnt mortality and longr lifspan vains will rmain 1
(R. M. Andrson and May 1991; Lederrg 2000). Th othr sid of th mutations or antign sl
growth quation is agriultur and prottion of food. But add to this th hangs in viruln
grat divrsity of th miroial world and th provn aility of viruss,at- aly 2003; Kuiken
ria' protozoa, and singl-lldfungi to adapt to nW nvironmnts.Summd hmagglutinin will h
togthr' humns and thir ompanion animals and plants onstitut vast stan' y volving A
rsou with vr-inrasingmoility and ontt with wild rsrvoirsof our thfoats and noss.]
miroial pthogns. to inftd watrfowl o
Th outom is invital:pathognswill rgularly irumvnt our entimi- y dirt ontat btw
rtlil dfrrss,and w will olonizd again and again y miros that populations.
adapt thir transmissitln yls to ur haits and movmnts.For xampl,
rh rnt mrgnof SARS inluds gnti ir-rtmdiats twnoron-
GN|s oF M
avirussin ats, ivt ats in wild-gam markts in hina, and humans: it p.
pars that SARS jumpd to hurans aftr som volution in ivt ts' not Many wll-known hu
dirtly frlm ts tr humans (\Vang t al. 2006). uh of this an pr- Mutation is also invit
ditd frlm th frarwtlrk f Drwinian mdiin via th adaptiv potntial that for th most part k
of rirosnd th volutionary history of pathogns(l-drrg2000). Mi- our odis and mak or
ros, with muh shortr gnrationtims than ours, hav n shown tim som ar nvironmntl
and agin to adapt to novl nvironmnts y natural sltion and povid Individual mutations o
slm of our st modls for primntl studis of volution (rt 1998; in favor of muttions .
Lan 200.5). ommon gntidisas
'i diilt
rsonsidrsth volutiorr of dis-
tins tht prinotlr natural
within a framwork of phlogn-
ors to ginthinking lik popula.
:ss out varial populations o{
ls, in arr ologial sphr that
tte 1972 L1rrg 2000; \/'An-
:diur|sin will grtl enfit
ls' frlmhuman popultior-rsthat
s of mirosthat liv within on
a I . 1 9 9 9 1S t a r n sa n d r t 2 0 0 1 ;
two asptsof th urrnt lruman
populatilnsiz and volution of
l t i v t o o u r o w n g n t i s a n d
)rsnta rsour for miroil
th hurnanpopulation has grown
l i l l i o n i n t p a s t d a d ,a n . hl u -
f tlr plant's iorn:rssand us a
t p r - r(tV i t l u s k t a l . 1 9 8 6 ; D i l y
pultionshav pandd aus
t us from th miroial world,
-rt mortality and longr lifspan
rg 2000). Th othr sid of th
tion of food. But dd to this tlr
h provnaility <lfr,iruss,t_
tt to nw nvironmnrs.Summd
imls and plants onstitr.rta vast
l otat with wild rsrvoirs of
I r g u l Iyi r u m v n t u r l l l l t i m i -
again nl again y miros tht
its and lnovmnts.F<lr-rpl,
ti intrmdiatstwn oron-
k r si n h i n a . n d h u m . t r l si:t p -
som volution in ivt ats' not
2006). uh of this an pr-
ldiinvia th adaptiv potntial
pathogns(Ldrrg2000). i-
i tha ous, hv n shown tim
by rrturalsltion nd provid
str.rdis of vlution (t 1998;
uolutilnr Biolog,of Dises d Drluiin dicine 285
A primary ampl is th shitling antigns (surfaprotins that stimulat
immun rspollssvia spifi antiodis) of influnza A viruss. \V r all
familiar with anrrual flu outraks' though prhaps not with th pandmi
Spanishflu of 1918-|919 tat killd mor than 40 million individuals. Flu
outraksour worldwid ah yar On a lss dramti sal as th viruss'
vr-hangingsurfa antigns avoid immunity stimulatd v prvious inf-
tions and nssitatonstant updting of flu vains (Bush et al.1999;Fer-
guson t al.200). lnf]unz viruss r rmarkly sirrrpl:ight RNA
strndsod for intrnal protins tht surround th RNA, an trnal pro.
tin apsul, polymrass (nzyms)fo viral rpliation, and 3 protruding
protinsthat at as antigns' 1 1 protins in all. Th 3 surfa protins ar
hmagglutinin(HA) fo inding to host lls, protin hannls that aid RNA
invasion into th host ll (M2), and nurminidas (NA) to hlp nwly
formd Partils saPth host ll. Gs oding for HA and NA ar sp-
ially prorr t() muttion' and otlr A arrd NA ar rprsntdy multipl
antignisutyps within grorrps of rltd hosts. For instn, watrfowl,
shor irds, and poultry, th main rsvoir and spillovr hosts for th in-
flunzaA viruss, hav at last 16 HA and 9 NA antigni sutyps (olsn
et a|, 2006). Th urrnt virulnt avian influnza sutyp is thus dsignatd
H5N1.
lfithin ah sutyp, howvr, HA nd NA gns ar subjt to rgular
antignidrift in that flu viruss umult nugh muttions to bom
lssasily rogr-rizdy th imnrun systmsof prviously infted or vai-
natd individuals (Bush t' a|. 7999: Frguson t al. 200). Th implia-
tions for human halth ar normous, oth in trms of prditing how long
vainswill rmain fftiv (D. J. Smith 2006) and in th possiility of
mutations or antign shuffling twI-lvirus sutyps,whih ould lad to
hangs in viruln and transmission twn hosts (Baignt and .
uley 200; Kuikn t l. 2006\. h grt far, r>follrs' is that th H5
hmagglutinin will hang influnza's tnsmissiility in humans, for in-
stan)y volving HA affinity frorn lls dp in th lungs t lls that lin
our throats and noss. Rathr than sttrdass assoiatdwith posur
to inftdwatrfowl or poultry, a nw form of HSN1 ould transmittd
y dirt ontt twn popl, lading to disas outraks in human
populations.
GNls oF MUAloN
Many wll-known human disass rsult from mutarions in our gnoms.
utation is also invital, a onsqun of mistaks in opying th DNA
that for th most part kps th rod straight during ll yls that rnw
our odis and mak our ggs and sprrr.Sot mutations :rspontanous'
somar nvironmt-ttally aused,ut st ;rssurdthat mutation nvr sleps.
Individualmllttions ur at rltivly low ats,ut th numbrs ar tiltd
in favor of muttions apparing in high absolut frqr.rny.Prvaln of
ommon gntidisasslik diats,musular dystrophis'and ansan
286 uolutior Biolog of Diss d Drtaii dii

undrstood in th framwork of volution vi thos inoral mutations al. 2005). Rgardls

and thir inhritanwithin familis and thni groups (Frank 2004; Tishkoff sist, and it appars t
and Kidd 2004). At th as lvl muttions ris in somewhr twn 1 in nti drift, provids l
10'000 and 1 in 100,000,000 DNA rpliations.Givn human population allls (Blanpain t
of mor than 6 illion prsons' ah with 30,000 gns and 6 illion s 2005; Sabti t al. 2(
pairs of DNA' novl mutations will aris ah gnration y th thousands,if Finally, inftions
not millions. Susquntly,novl mutation n trad and prditd within tints' and Darwinia
familis y pdigrs ut an also om to diffr in frquny btwn plpu- on human halth.
lations with highr or lowr proaility of inhritd disas. of thir gntis an(
Th vst majority of mutations will disrupt gns and harmful. on- prdisposd to inft
sidr phnylktonuri (PKU), th gnti diss ausd y th inaility to on inftion sts il
mtaoliz th amino aid phnylalanin aus of mutations in th PH For instan' som a
gn (Srivr and !atrs 7999). Th disas has varial phnotyps' of ptibility to a whol
whih mntal rtrdation is on of th wrst, ut it an avoidd if phny- rosy, turulosis, m
lalanin is xludd from th dit of afftd individuals (arfully rad th 2002). vn mor p
lals of your low-alori soft drinks). Mor than 400 mutant PH allls lik dngu' influnz
that nod malfuntioning nzyms hav n idntifid in various human damag from th pa
populations. Routin nonatal srning for unusually high phnylalanin tm whil attaking 1
lvls and dftivPAH nzym now provids a rapid tst for diagnosisand disas ar omplx
tratmnt for th most svrsymptoms dvlop. and immun rspons
On th othr hand, som positiv surpriss ar hiddn in th human
gnom as wll. A numr of individuals of northrn urlpan dsntin-
ftd with HIV-1 hav nvr dvlopd AIDS, and many of thm ar ho-
mozygous (hav th sam alll on oth hromosoms)for a mutation in th Diagnosis
ytokin rptor R5 on th surf of D4+T lls of th immu sys-
tm (Dan et a|. 1996). HIV.1 uss th R5 rptor to ntr D4+T lls Wht aspts of ar
and vntully ovrwhlms th population of hlpr T lls that nrmally r- winian approah to
spond to invadrs in th body, lading to a ollaps of th immun systm eyond individual p
and AIDS. This mutation is alld R5-^2 eaus of a dltion of 32 thir onntions wit
as pairs in th gn, lading to a nonfuntional protin that prvnts lutionary history. Th
HIV-1 from inding and invading D4+T lls. R5-A2 is found in 5% mnt to allviat pai
to 15Y" of northrn uropans ut is ssntially asnt from ast Asian' rapid diagnosis, tra
Afrian, and Nativ Amrian populations (Dn t aL.2,002). mnt of fvr and iro
Baus it is impossil that AIDS sltd for inrasd frquny of ths may rprsnt
growth of atria (l
R5-^32 within a singl gnration' it has n suggstdthat high fr.
quny of R5-A32 rsultd from strlng sltion in th past, spially Trating th symptor
from inftion y plagu or smallpox, whih wr oth ommon and ddly that inftions prsist
in uropan ppulations during mdival tims and until th lat 1700s (Gal- Th follwing di
vani and Slatkin 200). As is oftn th as with surmising sltionin th and hopfully an la
past' othr vidndos not natly fit th plagu or smallpox hypothss.In th asi symptoms z
mi inftion y th plagu atrium is as lmmon in R5-dfiint standing of disas:
s t r a i n sa s i n R 5 * s t r a i n s ( M s a s t a l . 2 0 0 4 ; l v i n t a l . 2 0 0 4 ) ' i n h u m a n 1. Is this an infti
populations svralothr nonfuntional alllsof th R5 rptor also - 2' If it is an infti
ist in high frquny (Blnpain t al. 2000), and th R5-^32 mutant lll a. If othrwis ul
has n rovrdfrom Bronz Ag (2,900 yars ago) skltonsfrom svral nvironmnt (
pls in urop long for plagu is thought to hav invadd (Humml t hazards)?
 dici
n via thos inxol mutations
ni gror'rps (Frank 2004; Tishkoff
arisin somwlrrbtwn 1 in
ior-rs. Givn a humn popultion
0,000 gns and 6 billion s
h grrrationy th thousands, if
an trd and prdited within
iffr in fr1ur-rY twn popu_
rhritddiss.
pt gnsand hmful. on-
issausd y th inaility to
21us of mutarions in th PH
as hs varial phnotyps, of
t, ut it n r,oiddif phny-
:d individuals (afully rd th
r than 400 rnr.rtntPH llls
ln idntifidin various human
lr unusually high phnylalanin
cs a rpid tst for diagnosis and
dvlop.
'riss r hiddn in th huntn
f n < l r t h rn u o p nd s
IDS, nd many of thm ar ho-
omosms)flr a trrutationin th
]D4+T lls of th immurr sys_
'5 rptorto ntr D4+T lls
lf hlprT lls that normally r-
r lllapsof th ir-nmunsystll1
\32 ausof dltion of 2
ulltional protin that prvnts
lls. R5-A32 is found in 5"%
:ntially asnt from st Asit-t,
Dnt l. 2002)'
td for inrsd frqunv of
as n suggstdthat high fr-
sltionin th past, spially
n wr oth omr-rronand da1l
ns and until th lat 1700s (Gl_
rvith surmising sltion in th
laguor smallpo hypothss.In
as ommon in CR5-dfiint
) 0 4 ; l v i n t a l . 2 0 0 4 ) , i n h L r m a n
ls of th R5 rptor also x-
nd th R5-^32 mutant alll
yasgo) skltonsfrom svl
ght to hav invdd (Humml t
uolutioltr Biolog of Diss d Drtuii diine 287
al. 2005). Rgardlss,svralothr muttions in th R5 rptor also pr-
sist,and it appars that sltionvia rsistanto inftions, rathr thn g-
ntidrift' provids plausil xPlanation for th high frquny of mutant
allls(Blanpain t al. 2000; Galvani and Slatkin 2003; Novmbr t al.
2 0 0 5 ;S a t i t a l ' 2 0 0 5 ) .
Finally, inftions and gnti disassoftn o-our within th sam pa-
tints,and Darwinian mdiin an also nompass this rodr prsptiv
on human halth. Patints may siknd y pathogns, ut othr aspts
of thir gntis and volutionary history (thniity) an mak individuals
prdisposdto inftion or to gratr pathlgnsisand mor svr disas
on inftion sts in (ook and Hill 2001; \Wissand MN4ihael 2004).
For instan,snr llls oi HLA gns of th immun systm inras sus.
ptiilityto a whol varity of pathogns, inluding thos that aus lp-
rosy, tubrulosis,malaria' and AIDS (Hill and Motulsky 7999; Dan t al.
2002)' vn mor profound is that diss fom ommon human maladis
lik dngu'influnz,malaria, and turttlosissr'rltsnot only from dit
danag from th pthogns bt also fror ovrrtion of th immun sys.
tm whil attaking thos pathogns (Graham t al. 2005). Thus th roots of
disasar ompl and in many instans grow from dp in our gntis
nd irmun rsponssto inftions,ppropiat or not.
n t i n -
Diagnsis
What asptsof ar for th sik and for publi halth distinguish th Dar.
winian approah to disas?Th largst distintion is tnding mdial ar
yondindividual patints to onsidr human populations in th ontet of
thir onntionswith th miroial world nd th patints' gnti and vo-
lutionary history. This an b don without ompronrising immdit trat-
mnt to allviat pain and sLrffringand in many ass an provid mor
rapid diagnosis, tatmnt' and rovry. A typial anrpl aiss ln trat.
mnt of fvr and iron dfiinvin patints rvith inftions,ausoth of
ths may rprsnt part of an adaptiv dfnsiv rspons to slow th
growth of batri (Williams and Ns 1991; Nse and.!7illiams 7994).
Trating th symptoms of inftd patints ould hav th undsid fft
that inftions prsist longr, with gratr potntial for tansmission.
Th following dision tr slrows atwys through this roadr ontt
and hopfully an lad physiians not only to diagnos and trat disson
th asi symptoms and auss ar known ut also toward gloal undr-
standingof disas:
1. Is this an inftion or a rrgnitlondition?
2. If it is an inftion, is this prson othrwis hlthy?
a. If othrwis unhalthy, is th patint ompromisd y poor
nvironmnt (.g.'poo dit and nutrition, smoking, or oupational
h a z a r d s )?
288 uoltior Biolog of Dises d Drtuini dii
aids (lysin, hydro
. If othrwis unhalthy ut with a good nvironmnt,ould th
triian, ut in his l
patint b gntillyprdisposdto inftion (.g.,poor immun
first dtt GA1 via
systm)?
a gnti tst asd
If
. othrwis halthy, should symptoms tratd (.g.,nausa,fvr,
whr th Amish or
nasal ongstion)'or should this inftion allowd to run its
of uropan dsn
ours with minimal intrvntion?
rdud among Am
3. If it is an inftion' is this pathogn uniqu andlor ommon in humans,
Rognizing this
or is th pathogn's sour from othr animals (alld zoooti dises;
.$olfe in. In part this kir
s th rviw y et a|. 207|.
.\ihr sionals undr th st
a. did this patint ontt th inftious sour' and how was
th silvr_ulltya
th pathogn transmittd (..g., y dirt ontat or y an arthropod
in th 1930s and 1j
vtor lik a mosquito, fla, or tik)?
rdisovry of this t
b. How ontagious is th pathogn:should patints isolatd until
thrapis and th r
thy ar n longr inftious?
losis that wr thol
. Do w know th inftious miro's losst rlativs?l/ill thraois
Andrson 1999)'
to fight thos inftions hlp this patint as wll?
d. Aftr w idntify whthr th pathogn is a atrium, a virus, a
protozoan' or a fungus, dos th inftious agnt hav rsistanto
Tratmnt and.
antimiroials?
4. If this is a ongnital disordr' what is this prson'sfamily history and
Darwinian mdiint
thniity?
th patint in front
a. Is th individual from a small population with a history of marriags
prditiv and prr
btwn los rlativs (.g.,first ousins or losr)?
th mdial world,
. Is this disashighly prvalnt in othr ppulations with th sam
mbra oth natur
thni origins?
th pross that tr
5. !ill a thrapy or intrvntion hav an fft yond this patint?
.!7ill translation humans
a. thrapy slt for highr pathogniity or viruln in th
ntions to th natu
miros (.g.,fftsof imprft vains)?
.lfhat sin of volution
. kind of sletion prssur will antimiroials rat, and what
dvloping tratmr
will th trad-off twn intrvning with a drug now and
sn latr, som .
failur of thrapisin th futur if drug rsistanvolvs?
th ost f ignoring
ass.
fftiv us of this dision tre prsupposs a thnologial world whr
laoratory diagnostis asd on modrn iohmistry and gnomis are
availal for rapid srning.lThis tr also prsums that mdial and ADAPA|oN
halth profssionals larn th naturl history nd ology lf pathognsand
If on onsidrs mi
parasits in th ara whr thy prati, and that thy undrstand th thni
aus rsistan to i
divrsity of th population thy srv.
rmrgn of pr.
Th work of Dr. D. Holms Morton mong th Amish of Pnnsylvania
Andrson 19991 L<
and Illinois provids lar onntion twn iomdial rsarh and
multipl-drug rsist
linial prati (Blkin 2005). orton and ollagus first diagnosd and
y aquiring drug r
thn idntifid th undrlying auss of prtzl syndrom, a form of rral
This is tru for Nels
palsy with Svrmntal rtardtion and thn arly dath in fftd hildrn
gonorrha and dir
(Strausst al. 2003). Similar t PKU, prtzl syndrom is glutari aiduri
Similarly' multidug
(GA1)' ausd y m'-rtationsin th dgradation pathwys of thr amrno
 uolutionr Biolog oi Dises nd Duini diine 289
dii
ids (lysirr,hydoylysin, and tryptophan). Morton is primarily a pdia-
od nvironmnt,ould rh
triian,but in his lini in ntral Pnnsylvarriah pursud th thnology to
inftion(.g.,poor immun
firstdttGA1 via nondgrded protins in rin sampls and thn dvlop
a gntitst sd on mirohips. Th mutation has n trad to urop,
ns b tratd(.g.'nausa, fvr'
whrth Amish originatd, and ours at low frquny within populations
tin allowd o ttn is
of uroparrdsnt. \With srning and dit, disas prvaln has en
rdudamong Amish to a third of th prtratmnt rat.
iqu and/orommon in humans,
Rognizing this rodr ontt is an intgral part of Darwinian mdi-
animals(lldzoooti disese;
in' In part this kind of training was at on tim providd to halth profs-
sionalsundr th sujt of gographi mdiin, ut it was swpt asid in
ftioussour' nd how ws
th silvr-ulltyars immditly aftr th disovry of antimiroial drugs
rtontat r y an arthropod
i n t h 1 9 3 0 s a n d 1 9 4 0 s ( B u r n ta n d W h i t | 9 7 2 ; W . A n d s o n 2 4 ) . h e
rdisovryof this ranh of mdiin oms on th hls of failurs of drug
luld patints isolatd until
thrpisand th rmrgnof inftious diseasslik polio and turu-
losis tlrt wr thought to hav n onqurd (Grrtt 19941 Lvin and
; losstrlativs?Will thrapies
Andrson 1'999).
int as wll?
3nis a batrium,a virus, a
)tiousagnthav rsistanto
Tratmntand Thrapy
this prsorr'sfamily history and
Dwinian mdiin asks hlth profssiorrlsto look yond symptoms of
th patint in front of thm and to prati mdiin in a way that is oth
r t i o nw i t h a h i s r o yo f m i g s
prditivand pmptiv(Zrhouni 2005; ulliton 2006)' Th hallng to
t s i n so r l o s e ) ?
th mdial world, inluding iomdial rsrh and drug disovry' is to
rr popr-rlatinswitlr th s-t
mrotlr rratural sltiorrand ommon dsnt (horn<llogy)as prt of
th pross tht translats iomdial rsh to linial prati. In this
fft ynd this patint?
translation humirns annot st thmslvs apart from thir volutionary on.
;niityor viruln in th
ntions to th natural world, inluding thir own gntis and iology. Th
rins)?
siof volution is highly pditiv,and it is unthial for thos who ar
antimiroilsat,and what
dvlopingtri1tmntsto ignor knowldg of volutionarv iology. s will
ning with a drug now and
.ugrsistanvolvs? b snlatr, som of ths pitfalls an only disovrdin retrospt,ut
th ost of ignoring volutionry iology in mdiin has n high in som
aSs.
ossa thnologialworld whr
i o h n r i s t r a n d g n o l n i s a
rlso prsurnsthat mdil and ADAPA|oN
:y and ology of pathogns and
If on nsidrsmiros' th ida of natural sltionis asily aptd-
d that thy undrstandth thni
us rslstanto antimiroial drugs has apidly volvd and rsultd in
rmrgIlof prviously ontroild pathogns (Grrtt |994; Levin and
long th Amish of Pnnsylvnia
Andrson 1999; Lederrg 2000). In som ass atria hav volvd
]twn iomdial rsarh and
multipl-drugrsistan,partly ausof novl mutations ut just as oftn
d ollagusfirst diagnosed and
y aquiring drug rsistanvi DNA hng from othr atrial spis.
zl syndrom, a form of rral
This is tru fo |'Jisseri,th atium rsponsil for sually transmittd
n arly dath in fftd hildrn
g<lnorrha rrd dirtly trarrsrnittdatrial mningitis (Snydr t al. 2005).
ll syndrom is glutari aiduria
Similarly, multidrug rsistan has volvd in th flsh-ating Stpblocous
l a t i o n p a r h w y so f t h r a m i n o
2 9 0 uolutionr Biollg of Diss nd Drtuii edii

urus' whih is oftn aquird in hospitals. In its nw and imprvious form ar not yt availal
this ommon skin atrium auss svr disas of skin and soft tissus' simulation studis
somtimslading to dath (Dikma t al. 2001; ady and ov 2003). gl drug until rsi
on th othr hand, it is implausil that humans will volv nw mha- gratst halth r
nisms of dissrsistan,at last in th asnof som kind of rgular and th sam ptints
atastrophi hallng that would svrly rdu fitnss of thos who lak with th lowst tr
gnti prdisposition for fighting inftions. Again' this is not to say that son 1.999;'Bull an<
humans hav always n unafftd y th powr of natural sltion- Again, possil ou
rnt gnomi analyssshow ongoing sltionin rain siz (Mkl-Borov ontrolld human
t al. 2005) and rnt sltion for adlt latos mtaolism in uropan provid dfinitiv l
and Afrian populations (Tishkoff et a|. 207). But th disparity of gnra- As notd for inf
tion tims of miros nd hmans spurs natural sltion in favor of mi ult, although sv
ros, whih will xprin sltion at multipl lvls: within individual u s ( . g . ,m a s l s
hosts, twn human hosts, and twn humans and pathogn rsrvolrs possiility that 5
in ommnsal or wild populations of othr animals. human pathogn' :
An intriguing possiility is that our immun systms ar urrntly undf for widsprad usi
sltion via immunopathology, disas usd y ovrrsponsy our im. in will provid s
mun systms to What ould othrwis mild inftions (Graham t al. fortuntly' this
2005). our immun systms ould sn as a sris of trad-offs twn sltion within ht
rognition of slf and nonslf, dfns against invading pathogns, and th high ts of path
potntial damag to noninftd tissus whn laring invadrs from inftd ally' imprft v
tissus. In this as sltion would toward an intrmdiat phnotyp mission to sap ]
that alans trad-offs. Gnomi analyss of immun-systm gns' om- individuals (Gand
ind with primnts in modl systms that furthr dmonstrat how ths halth impliationl
kinds of trad-offs funtion, will provid an ativ ara of rsarh for yars must tak into a
to om (Graham t al. 2005). and tht within-l
Thus modrn mdiin still finds itslf on a shifting landsap, and w on- (Starns and rt
tinu to writ th haptr in human history suggstd y Joshua Ldrrg loal outraks or
(2000)' whih h alls ..our !its vrsusThir Gns.''volution in miros pathogns in th ft
will nvr stop, ut w ar ginning to apply stratgis spifially dsignd Finally, muh ht
to halt thir transmission, slow rsistan to antimiroials, and improv mnt that pathog
vains. Th first issu is prhaps th simplst: hygin an halt pathogn slows. transmissio
transmission. This was immdiatly apparnt from th famous pidmiologi- viruln id om.
al sluthing by Dr. John Snow in London, who in 1854 rmovd th handl uropan raits i
of a puli watr pump on Broad Stret in Soho and provd that a loal the virus was intro
holra pidmi am from a ontaminatd wtr supply (!7ills 1996). Sim. sap. Lowrd vir
ilarly, plagu and th Blak Dath' artsd y th batrium Ysii pstis is sond ampl
and rsponsil for som of th most infamous human pandmis, hav sis that alans vi
largly wand as human disassin aras whr rodnts and thir flas ar r- nition of viruln
movd from human living quartrs (Gag and Kosoy 2005). In trms of vo. within th host, to>
lutionary iology, hygin irumvnts th opportunity for sltion for host, and, finally,
pathogns by simply rmoving hains of inftion and prvnting transmis- Neisseri meigitt
sion (Galvani2003). transmission is fror
Slowing volution of antimiroial rsistan is on of th largst publi ally asymptomati.
halth hallngs w fa, ut most work in this ara is hypothtial aus mission and is atu
ritial data that ould distinguish whih stratgis would most fftiv optimal viruln
 diie
. In its rrwand imprvious form
dissof skin and soft tissus,
20I;adyand ov 200).
humans will volv nw mha-
snof som kind of rgula and
rdu fitnss of thos who lak
rs. Again, this is not to say that
h powr of natural sltion-
:tionin rain siz (kl-Borov
latosmtaolism in uropan
t07).But th disparity of gnra-
natural sltion in favor of mi.
multipl lvls:within individual
h u m n s n d p a r h o g nr s v o i r s
animals.
lun systms ar urrntly ndr
" r s d y o v r r s p o n s y o u r i m -
: mild inftions (Graham t al.
as a srisof trad-offs etwn
inst invading pathogns,and th
:n laring invadrs from irrftd
rd an intrmdit phnotyp
; of immun.systmgns' om-
at furthr dmonstrat how ths
l ativ ara of rsrh for yars
a shifting landsp' and w on-
y suggstdy Joshua Ldrrg
ir Gns.''volution in mirobs
lly stratgisspifiallydsignd
to antimiroials, and improv
plst: hygin an halt pthogn
nt from th famous pidmiologi-
who in 1854 rmovd th handl
Ln Soho and provd tht a loal
l watr supply (Wills 1996). Sim-
r y th trium Ysini pstis
[amous human pandmis, hav
rr rodnts and thir fls ar r-
nd Kosoy 2005). In trrs of vo.
r opportunity for sltion for
. f t i o na n d p r v n t i n gt r a n s m i s -
:anis on of th largst puli
Lthis ara is hypothtial aus
tratgiswould most fftiv
F,uolutionrBiolrlg of Dises nd Drtuii ediin 291
ar not yt avirilal (Lipsitlr 200l). Rgardiss,mathmatil modls and
simulation studis indiat th;rt in liu of attirking ah mio with a sin-
gl drug until sistanrahs a lvl that ompromiss puli halth, th
gratsthalth nfitswill gaind by dploying omintions of drugs in
th sam ptints at t sm tim or targting for ontrol thos pathogns
rvith th lowst transmissi.lt-t rzrts(Bonhoftrt a|. |997: Lvin and Andr_
soo1,999 B u l l n d W i h m a r r2 0 0 1 ; L i p s i t h 2 0 0 1 ; D . L . S m i t h t a l . 2 0 0 5 ) .
gin, possi1Outomslravnugh ontingnisthat orrly tim, arfully
ontrolldhumarrtrials, and laoratory pimntswith modl systmswi1l
povidedfinitiv 11llswrs.
As notd for influnza A' kping fftiv vins orr hand an diffi_
ult' although svral vains hav rmaind fftivvn aftr dads of
us (.g.,masl.svins). T.his qustion has arisn again baus of th
possiilitytht H5N1 avian influnza ould mrg as a dirtly transmittd
hurnpthogtr,ird it is not f:rsilto provid prftly matd vains
for widspr:rdus. Thus th hop is tht vn an imprftly mathd va-
in will provid som lvl of prottion nd slow virl transmission. Un-
fortunatly, this dos not tak into aount that pathogns undrgo
sltionwithin lrosts that tluld favor mor virulnt pthogns (dfined y
high rats of pathogn rplition alising ll r tissu damag). Spifi-
ally' imprfr vains mav alrs pathogns to volv oth fastr trans-
rnissionto sp prottd irrdividuals and high viruln in unprottd
i n d i v i d u a l s ( G r - r d o n t a l . 2 0 0 1 ; v a n B o v n t a l . 2 0 0 5 ) . A g a i n , p u l i
halth implitiotls of using virinsthat only Partially potet individuals
must tak int() ount that sltion or-rperthognsoLlrs at svral lvls,
and that wirlrin-host volution gnrlly Slts for gratr viruln
(Starnsand rt 2001; Galvani 2003). Rlasing partial vins to stm
loal outaksluld hav th unintndd risk of sltingfor mor virulnt
pthognsin th futur.
Finally, nrlrhas n nrd f th volutiln of viruln and th argu-
mnt that pathogns should volv lowr viruln ausdamag to lrosts
slows transmission (wald 1994). mpirial support for th optinral-
virulnida o-tsfrom a frv xampls lik th rdud dath rat among
}-uopanrits ilrftdwith nryoma virus in Austli in tlr yars aftr
th virus was introdud t.lontrol th r:ritsthat wr rvaging th land-
sap.Lowrd viruln in th trium that auss diphthria in humans
is a sond anrpl (brt and Bull 200). I}utagain, th trad-off hypoth-
sis that alnsviruln and trnsmission assumsa rlativly simpl dfi-
rrition of virln, wlrilr is tually a fuIrtion of pathogn rpliation
within th host' tiity of rh pathogn to th host' immun rsponsof th
host, and, finally, whih tissus ar afftd. For instarr' th atrium
|'lisserimenigitidis auss mningitis whn it infts spirrl tissus, ut its
transmissionis from inftions of th uppr sPiratory trt, whih ar usu-
ally asympt<lmati.hus virr.rlnin N. migitidis is r-rnlatdto trans_
missionand is lly a spillovr f inftions within hosts. From th ida of
optimal viruln oms th notion that disasould avoidd if pathogn
2 9 2 uolutionr Biolog of Diss d Drini dii

viruln ould ontrolld. Th qustion thn ariss: wht will th

targt for sltion?t nd Bull (2003) suggsttlrat sltion should tar- Prognosis
gt viruln dirtly y attaking pathogll toxiity; vains against diph-
thria and prtussis hav n dsignd to do just that, with th rsult that !7hr dos Darwir
surviving Strainsar lss pathogni. gratsthllngis
to hanging nviror
th largst and rn<
ot",lMoN DEsN
doomsday slrrio
Idas of olnmon dsntand homology may hav gratr diffiulty in ing th possiility woul
aptdin mdial prati for th sam rasonsthat th notion of ommon that ould rdu di
dsnt gnratsognitiv dissonan in th gnral population. Th on- of volution hs m:
pt of homology, that traits in diffrnt spishav ommorr volutionary Givn variation ir-r1
origins' is ritial to dvlopmnt of pharnrautials,whih ar first tstd fitnss arrd with inl
on mammals lik mi, rats, nd dogs. Promising andidats may thn tion is a rtairlty.I
tstd on primats for linial trials with humn voluntrs. Thus io. intuitiv outoms
mdial rsarh advans y undrstanding that spis phylogntiaily lvls. Th possiil
losst to hrrnranshav th most similar gntiS'physiology, :rnd immun most imortant on
systemsto hurnns nd thus prov to rlial rodls for rsarhand d- iiri (\/illiamsrrd
vlopmnt of tratmnt and thrapy. ft vains luring
How badly an things go if mmon dsrrtis disgardd?on poignant potntially alarmin1
as is that of Bay Fa, an infant orn with an undrdvlpdhrt who whil th vain s
rivd th hart of a aoon at Loma Linda Univrsity Mdial ntr in 1orsvr disas
1984 (Bily t al. 1985). Th aoon donor wzrson of fiv that had bn aross multipl lv
tstdfor irnmunologial sinlilarity on th lrasisof thr HL gns.Th a- transmissiolt, l.lnirt
oon with lowst rativity to th infant's illmun lls (lymphoyts)was Ns 1991; lvld
hosn as donor. Bay Fa survivd for 20 days aftr surgry with th hlp of Th prognosis for
th immunosuppssiv drug ylosporin ut vntually did from rjtion -ranpopulations is
of th transplantd hart and othr organ failur. thial nd produral man that ]ntis
qustions asid, a trouling part of th story is that svlmistaks dirtly triizrrrswith infonr
rsultd from a rjtion of volutionary thinking. lik PU and GA1,
Lonard Bily' th lad sugon for Bay Fa's opration, admittd to ould don with l
ing a fundamntalist hristian during a radio intrviw for th Aus. batd, spially for
tralian Brodasting orporation.2 Baily hos aoons as donors us fo inftions, and rl
thir harts wr availal nd th right siz. Askd why harts of him- thzlt us gnti di
panzs' humans' losst living primat rlativ,wr not onsidrd,Bail,v ut gain, tlris is a m
answrd, ..Th sintists that ar kn on the volutionary onpt' that Progrss in grth
w atually dvlopd srially from suhuman primats to humans, with mi- patints to ur thi
tohondrial DNA dating and that sort of thing, th diffrns hav to do trial in Philadlphia
with millions of yars. That blgglsmy mind sornhow.I don't undrstandit provd iothnolog
wll' and I'm not sur tht it mans a grat dal in trms of tissu homology.'' nti disashng
Infant transplants y Bailv.s group at Lora Linda susquntlyusd only larg populati<lt-tso
human donors. Th prospt of using othr spisfor transplantationon- ertainty that ugl
tinus to qustiond ausfor most organs th ross-spisimmunity luld radiat gn
arrir is high, vn with immunosupprssivdrugs, and trsof th risk y than it did a nt
that novl pathogns ould transfrrd to hr:nransfrom th dorror animls Tlr prolms th:
(Dshamps t al' 200.5)' volution fasintin
i diie
on thn ariss: what will th
suggsttht sltion should tar-
,n toxiity; vains against diph_
do just that' with th rsult that
y hav gratr diffiulty in ing
)asonsthat th notion of ommon
h gnralpopulation. Th on-
ishav ommon volutionary
nautials'whih ar first tstd
.omisingandidats may thn
ith human voluntrs.Thus io-
ing that spis phylogntially
ntis' physiology, and immun
ial modls for rsarh and d-
:ntis disrgadd?On poignant
ith an undrdvlopdhart who
nda Univrsity dial ntr in
tor Was on of fiv that had n
lasisof thrHL gns.Th a-
immun lls (lymphoyts) was
lays aftr surgy with th hlp of
' u t v n t u | ldyid from rjetion
r failur. thial and produral
y is that svralmistaks dirtly
inking.
tby Fa,s opration' admitted to
a radio interviw for th Aus.
:hsabons as donors aus
;iz. Askd why harts of him-
ltiv' Wrnot onsidrd, Baily
n th volutionary onpt' that
ranprimatsto humans, with mi-
thing, th diffrns hav to do
d somhow.I don't undrstand it
lal in tmsof tissu homology.''
na Linda susquntlyusd only
s p i sf o r t r a n s p l a n t a t i o n o n .
r g a n st h r o s s - s p i ism m u n i t y
v drugs,and ausof th risk
l humans from th donor animals
uolutionr Bkiog of Diss nd Drtuii dii 2L).]
Prognosis
.!7hr
dos Darwinian mdiin tak us in th futur? Fist and formost, th
gratsthallngis that miroshav an almost unlimitd apity to dapt
to hngingnvironmnts,and that humans in th modrn world ar on of
th largst and most onsistnt nvironmnts vilal to thm. But r
doomsdaysnariosof disaspandmis nessary?Not rally, ut to ignor
th possiilitywould also foolish' and many intrvntionsan adoptd
thatould rdu disassvrity(Purssll2005). Th issu is thatth pross
of volution has many fas,with hang in th futur as th only rtainty.
Givn variation in pathogn populations, with som traits onfrring highr
fitnssand with inhritanof trit diffrns,adaptation y natural sl-
tion is a rtainty.Rsponssto sltion,howvr, somtimsh:lv oLlntr-
intuitiv outoms aus sltion ts in diffnt dirtions at diffrnt
lvls.Th possiility that volution ours at multipl sals is among th
most important ontriutionsvolutionary iology has mad to mdrn md-
iin (williams and Ns 1991; Starns and rt 2001). Dploying impr-
ftvainsduring puli halth mrgnis,as disussdaov, provids a
potntiallyalarming xampl, whr pidmis ould slowd rgionally
whil th vainsltsfor highr virulnamong surviving pathogns,with
mor svrdisasin susquntoutraks (Gndon t al. 2001). Trad-offs
arossmultipl lvls lay at th hart of th volution of pathogn viruln,
transmission,and rations of our immun systmsto invadrs (Willim and
N s 1 9 9 1 ; w a l d 1 9 9 4 ; e r ta n d B u l l 2 0 0 ; G a h a m t l . 2 0 0 5 ) .
Th prognosis fo gnti hang that would rdu disas inidn in hu-
man populations is mid. Rapid advans in gnomis and gn dislvry
man that gnti srening,prnatlly or soon aftr irth, will provid pdia-
triins with informatiln ndd to trat vn mor of th mtaoli disass,
lik PKU and GA1, ausdy mutations.Gnti srningof adults,nd wha
ould don with that information, is a lnattr of this tht is still ing d-
atd,spiallyfor th risk of gntidisas,th prdispsitionof irrdividuals
for inftions, and rats hargd for halth insuran. Srning for nlutatit-'rls
that aus gnti disasould hlp potntial parnts did how ttl prod,
ut gain, this is a mattr of privay and this lutsidth sinof volution.
Progrssin gn thrapy-rintroduing funtional gns into th gnoms of
patintsto ur thir disas st bak aftr a fatality in a gn thrapy
tril in Philadlphia in 1999 (Thompson 2000) ut is ing rvisitd with im-
provd iothnologis.But nothing in th study of modrn gnomis and g.
nti disashangs th fat of human population siz' and that mutations in
larg populations our t high asolut rats. Thus W an still prdit with
rtintytht ugnis,th disardd ntltion that sltivrding of humans
ouldradiatgntidisas,would do vn lssnow to allviathuman mts.
ry than it did a ntury go whn ugnisprograms wr first institd.r
Th prolms that fa Darwinian mdiin ar th sam ons that ak
volutionfasinatins to som and frustrating to othrs. \/ithinth rno rtain
294 uolutior\, Billll| lf Diss d Drtuii dii

framwork of inhritarrand natural sltion, volution is inflund by Andrson,\i.2004'

tWntith-nt
han vnts lik muttions, han nountrs with an individual who
Btrignt,S. J., andJ.
brings nw inftion into ommunit' or a rapidly altrd nvironmnt. irds:Dtrmin
On th on hand' som find omfort in th ida from Darwinian mdiin Biosss25:6
..prditd''
that sonr disassan from tlr maladaptati-ln lf humns to Bily,L. L., S. L. N
.Willims
thir urrnt r,vrld(Ns nd 1994\. Orr th othr lrarrd,rv rvould Boon-tl-hirm
fl ttraout Darwilrian mdiin and looming plgusof mldrntimsif Amrin edi
Blkin, L. 2005. A d
th idntitisof th nt pandmi-ausingpathognsWr mor prdital
Blanpain, ., B. L.
(many would say that viruss lik HSN1 or ola r most likly; s lav- V. Wittmr, G
land t a|.2007)..W.an tak gratromfort, howvr,from gographipi- nonfuntional a
dmiology, muh as Darwin rnay hav found omfort though his intrstin 96: |68-1645
gographi distriutions of populations nd spis(Burnt nd Whit 1972; Bonhoffr, S., . L
to prvnt anit
Ldrrg 2000; W. Ardrson 2004; Tishkoff and Kidd 2004). Thus w
Sies US 9,
know that th historis of populations will influn th prvaln of
Bull, J. J., and H. A.
ongnital disordrs, and w know that disas organisms hav gographi d Sstmti:
origins, zrsdo th resrvoirs for disas and potntial inst vtors for trans- Burnt, N4., and D. t
mission. By :rknowldgingtlrat humans liv :rspart of th worldwid olgy amridg: :
of inftious disas rndthat voiutiorof pathogns is th invital out- Bush, R. ., . A. I
Prditing th
om' w n vn mo prpard th nxt tim th monkypo visits
ntrs for Disas
Kansas. outrak of mt
.Wisonsin,
20(
642-646.
AKNoWLDGNs
l v l a n d ,s . , D .
.!(lr
mrgn:
Mv rsrh is supportd v th National Sitr-rFundation: olgy f Inftious
irobiolo91,
Disss Program (F-027052) and th Short Grss Stpp Long Tm ologial
olk, G. S., and A
Rsarh projt (DB 02|7631). I thank th ditors and anonymous rviwrs,along
disas.Nture
with Irn kstrand, Normn Johnson, ary Plss,and my physiian ark Paulsn,
ulliton, B. J. 2006
for omnrnts. I also thnk Dr. Pulsn for his strit attntion t mv iIrftionsarrd
Zerouni. t
gntls.
Dailv. G. ., S. All
H. A. N4oony
1997. osyst
Nos osystms.1ss
Dan, M., M. arri
1. avt must mird hr:disasdsribsth physial damag to patint'
by gnti vrsr
whih is n individual phnrnon.SrningIr idntify individuls with partiu-
Humn Geti
lar gnotv1.ts or spifi inftions, ut ths individuals will nt nssrily
Dn, ., M. Cri
dvlop dis.rs.Nonthlss' srningan pdit diss'nd disirsrisk an
undrstotldat th lvl of plpulationswhn posursto pthognsnd/O J. J. Godrt' S
Vlahov, R. Kas
f r q u r n i s f g r l t i m u t . r t i n r r h i g , h .
Gnti rstri1
2. original rdio rodst on Jun 3, 198.5,during th progr:lm Hlth Report.
alll of th l
3. S Haldan (1964I for an amusing nd radalssay tht dfnds
Dshamps, J. Y.' F
mathmatial thoris of population gntis,inlr'rdingrfrnto tl-rlan
notransplan
twn Inuttion nd sltion,nd som lovly r.rsaout his 11,.n Iifg.'..|i.o
Diarond' J. 2003.
fight with olortl nr.
Dikma' D. J.' }4.
N4. Bah, and
|BLIoGRAPlJY to Stphloco
susptiility
Andrson, R. M., nd R. . ay. |991. Iftlus Disessof Hu"tlts:Dmis urp, and th
d trll, oflrd: Ofrd Univrsit Prss. Program, 199i
i dii
] t i o n ' v o l u t i o n i s i n f l u n
:ountrswith an individual who
r a p i d l y l t r d n v i r o n m n t .
r ida from Darwirliilr-r mdiil-l
r th rnaladaptatiorrof humans to
94). on th othr hnd, w would
loming plagusof modrn tims if
pathognsWr mor prdital
ol:rr most likly; se lirv-
)rt' howvr'from gographi pi-
-rdorfort
through his intrst in
i spis(Burntand Whit 1972;
,rkoff nd Kidd 2004)'
Ths w
will influn th prvaln of
Lsasorganisms hav gographi
potrntil instvtors for trans-
] as part of th rvorldwid ology
. pathognsis
th invital ut-
nt rim rh monkvptlx visits
: Foundtion: olog of Inftious
rass StlpLong Trm ologial
tos trd nonylolls rviwrs, alot.tg
lss, nd my physiiirn ark Pulsn,
.it attntilnto m inftions
nd
sth physial damag t() patlnt,
n idntifr.individuls u,ith prtirr-
lividLralswill not nssrily
Litdisas,and dis:rsrisk n
Osursto pthognsand/or
Juing th IO!.rm Lllth Rplrt.
Ldlssaythat ]fnds
;luding rfrnto th alan
i vrsbout his wn lif-nding
lus Disassof umtls: Dltlis
lss.
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 edii
us rsistn
rvisitd:ommunity-
s urus-a rrrrgingprolm for
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Reuituf ,llog,uolution, d
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Vood,J. M. Daly, J. A. Mumford,
triologiland volutionary
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Rviw of ats and SARS. mrgig Iftious Disss 12: 1834-184'
.Wiss, plaining th difft
R. A., nd A. J. Mihae|. 24' Slialand nvironmntal risk fatsin
th mrgn of inftious disass.Nture dii 10: S70-S76. ontribution to th
.!illims,
G. ., and R. . Ns' 199]r'T dawn of Darwinin mdiin. Thr is a tnd
Qurtrl Ruieul lf Bbklg 66: 1'_22. an gnratd
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inftious disass.Nture 447 : 279-283. sis of two typs (
Zrhouni, . A. 2005. US imdial rsarh:Basi, translational, and linial spis morpholo1
sins.Jourl of th mri edil Asslitil 294: 13.52-13.58. into distint, not (
onstraints on tht
som forms n
amin in dtail
obsrvational, ant
m p l . A l l t h r a
tion of ontinuou!
 dii

Torasz. |999.T dvlopmnt

at Stp h loous
:thiillin-rsist
:dii340:5\7-523'
; in influza
orrtrtll.
Sie12:

1005.Stgi inrritions
istane.Prodigslf th
in
Byondth Darwinian
_1.5 8.
| .S a u n d 2s 0. 0 5 ' o l l l p a r i v
:strvlt thptlrognr
Paradigm:Undrstanding
1 - 218 .
Dises.Nw Yrk: oford Biologial Forms
:althnddiss:
Work in progrss.
Brin Goodtlli
r' andD. H. ortorr.2003.Typ I
7 ptints.
AmriJourlof

\cthirurgiBIgi104:

lssons' highhops.FD
;/faturs/2000/5 00-gn.html. Th qustion that I shll min in this ssay is th rlation twn form
ls of iogography of l-ruman
, Centics36: S21_S27' and funtion in volution' and how th rigin of diffrnt forrs is undr-
ght'. C. Bitt.J. S. Silvrmn' stod. Form hs vo aspts:slrpor strutur' as in tlr morphologv of or.
osr:rn, . Irahim,S. A. omar, ganisms; and pattrn in tinl tlr lrirvilr,suh s th movmlrts of
ad,J. K' Prithard,G. A. !ary, individual orgarrisms(swimming' walking, flying) or th spatial pattrns of
o no f h t r m a n
l a t a s p r s i s t n i n
olltivs,suh as floks of irds or faging olonis of soial insts.Ths
+0. htrtristis of spisr oftn rfrdto olltivly as hratrs' and
. . d l k r ,a n d .
thy hav dirptiv, funtinl signifi:.rn for tlr survivl of tlr spis.
Jr irnprftvaination:
RI Soit of Lolldo, Series B Howvr, thr is a distintion twn undrstandingth origin <lfa form
(i..,how th form an ris i organisms during thir volution) anl th pro-
d P . . N 4 t s o n 1. 9 8 6 . H u t r l ssrvhrynatttrl sltiorrats t hang th rlativ frqunisof th
sis.iSic6: 68-373. diftntforrrrsin poprrli.rtinausof thir survir,lvaIu.N21tulsl-
D a s z a k ,a n d B . T . t o n . 2 0 0 6 .
tiorr annot xplain how any form origir-rats, and it n usd <lnlyin x-
s Disss|2: |84-1840.
plaining th diffrntil aundan of haratrs' onntd with tlrir
t n d n ri o n m n t a rl i s k f t o r s i n
ediill l0: 570_576. ontriution to th fitnssof ganisms'
l n o t ) ritn i n l r l d i i n . hr is tndnyam()ngvolutionirrvbiologiststO ssumtht Ily ti)rnl
an gnratdas a rsult of random r,ltriationin tlr gns,rsulting in lr
luoltio of Popl d indfinitdivrsity of possil forms for volution tl work with. This is th
assumptionthat I shall suggstis mistkrras a gnrl prinipl, ol-rth a-
}07.origins of mjor uman
sis of two t,vps of vidn.Tlr first f. tlrs tlms for osrr,tion lf
isi' translational,zrndlinial spismorphology, whih shlwsthat flrtnsgnratdduring volution fall
:soitiln 29 4: 1 52-1 5 8. into distint, not ontinuous' lasss.An impliation of this is that thr ar
onstraintson th possil forms of organisms and tlrir havilr,so that
sonrforms n gnratd,,lril orhrs annot. TIr as tht I sll _
trrrrinin dtil ms forrrplant morphollgy, sin w hav rimntal'
obsrvational,nd thorti:rlvidnflr distint sts lf pttrns irr this -
m1.ll. All thr ar rqtrirdto mak a onvining s against th assump-
tior.rof ontinuous vriation as a asi prinipl in volutiln.Thr r mn
2L)9
.00 Bod t|l Driin Prdigtn

xampls that an usd forn animl mophoiogy alld olltive havior

h
pattrns' som of whih I disuss, ut th plnt ampl mks th grrral
poits larly and illustrats how th argumnt tat rlats to morphology
an pursud in otlrr :rses.
Tlr sond typ lf vidn that I prsnt flr onstraints ln tlr possibl
\
forms availal for volution oms from t ntur ilf th prossSthat
gnrt th forms. Ths ar th dynari ativitis whry diffrnt mor-
phologis and hviors ar gnrtedy intrationswithin orgnismsdur-
ing thir dvlopmnt and twrr.thrnin prdr.ringhirvior pattrns.
This involvs som lmnt of thortil modling, showing what typsof
prossS or'rld giv ris to t flrs rrd rr,lry hs sult in disrt,nol
ontinuous' variation in haratrs.
Th viwplint psntdh in no sl.lshallr.rgs t irr-rportano{ nat-
ural sltionirr volution. Whthr th flrms f organisms ar initilly gn-
ratd as distint pssiilitisor as a ontinuum of variations, sltionstill Figur 1. Th tr
in fushia 1middl)
oprats to influn th frquny of th diffrnt forms <lrdingt the fit-
nss that thy onfr on th individual mnrrslf a spis.Th important
point nrd is that tlr gnrativorigins of iologial strrrtursad pttrns
ar not to undrstood in th sarWay as thir diffrntial aundanin
diffrnt l,raitats.
.W.
ar still t from hilving anything lik a orrrpltdsription of th 90.. Tlr tlrird pt
prosssin organisms that undrli th origin of thir distintivharatr. ah nod nd a
istis. Hwvr, I liv tlrat w undrstnd suffiintl,vwll th asiprin- fom h prdin
ipls of ths gnrativ ativitis to draw som importnt lnlusions th growing tip oI
aout oth th distintions and th rlatilnshipstwnform and funtion turs y growth al
in iology. on rh sis of th xampls prsntd:rnd th gr-rrl prinipls
that govrn nonlinar dynamis, I shall agu that th prosssinvolvd
oRPHoGl
hav th onsqunthat som ftrrrs plssil,whil othrs are not. Un-
oF MoRP
drstanding th origins of th forms that dfin diffrnt spis thrforr-
quirs thrtw ltndstadth dynmi prosssthat gnrttlrm,whih A omrlrnsiv
r not th<lsof natr.rralsltion. Douady nd lrrd
a modl that sho
damntal ida is tl
Phylltais as a Slf-organizing Growth Pross mordium lvill arir
egir-rfalls low
A lassi ampl f form in flowring plants onrns th arrangmntof sourof inhiitio
lavsalorrgth stn-r(phyllotaxis).It has n rognizedfor lnany yarsthat ing. Howvr' th
P t t r n s r. h o u g ht h r r I s os o m i n -
t h r a o l l l y r h r i m a 1 ..( ) l n m ( ) n that as lavs gro.
frquntly found arrangmnts. Th rnajor arrangmnts r shown in Figure an initiatd. It
1. Th first of ths has th forr of a spirl whll on looks dorvn on th that rsults in th .
plant, illustratd hr y yua. Sussivlavs ou1.lyp()sitionsrotatd mristm and its g
throrrgh wll.dfind avrag angl (l37.5o) rlativ to th prvious laf. terns of laf arrar
Th sond pattrn is alld wlrorld, in rvhil-rtwo Or tnor lvsarlst ondary ons. Tlrt
ah nod. h lavsat ah sussivnod ar rotatd so tht th lavsar pattrning poss
loatd ov tlr gps lf tlr prvilusrvhorl. Th xampl slrown is fushia' l trms, a of
whih has two lavs at ah nod' with rotation of sussivrvhorls through dynami prossl

)rphologyand olltiv hvior
plant xampl maks th gnral
mnt that rits to rnorphology
nt for onstaintsorr th possil
th natur tlf th prosss that
ativitiswhrv diffrnt mor-
ltrationswithirr oganisms dur-
in prodtring hirvior attrns.
nodling, showing rvhat typs of
why thsrsult ir-rdisrt, not
hllngsth imp<lrtrr
m s o f o r g n i s m sa i n i t i a l l y g n -
ruum of variations' sltion still
ffrnt forms aoding to th fit-
l so f e s p i s
. h impotan
iologialstrutursand pattrns
.tsthir diffrntialattndan irr
<a omplt dsription of th
gin of thir distintivharatr-
d s u f f i i n t Iryv l lt h
lW som importl-lf onlusions
ships twnform and funtion
:sntdand th gnral prinipls
]gu that th prosss involvd
ossil,whil othrs i1rnot. Un-
tirldiffrntspisth|.llr_
sssthl1tgratthm, whih
,owthPross
nts onrnsth arrangmnt of
ln rognizdfor mn-:rrsthat
r r s . h o u r . ht h : t I s s r t l r i t r -
rrangmntsar slrown in F.igur
al whn on looks down on th
lavsoupy positions rotatd
5") rlativto th prvious laf.
hihtwo ()r mor llvs is at
] ar rotatd so tht tlr lvs ar
l. Th ampl sho,rr is fushia,
rion of sussivwhorls thrugh
Bed th DriLl Prdigm 01
of nat_
igur1. Th thrr.. typsof phyllotais:spial' as in yua (/e/;drrsst'
as
in fuslri (midrlla);and distihlus,as in r,,liz(right).
90.. Th third pattrn is knorvn lls listihtrs,in r,l,lrilrth
is a singl laf at
ah nod lld irh sussivlf is loirtdon th opposit sid of th sfm
a s i p r i n - from th prding ln. Th gnratolf all ths pattrns is tlr lnristm,
th growing tip of th plant' wh th lirvsar produd as distint stru-
tursy growth and diffrntiationlf Ilsin intration with on anothr.
MoRPHoGENE| FILDs As GNRAoRs
oF oRPHoLoGY
A omprlrtrsivxaminatitln of tll gnrativpross in th rristmld
Douady nd oudr (1996) to synthsizth omponnts of this pross intl
a modl that slrtlws how th thr typs of phyllotis ould our. Th fun-
damntlida is that th mristm is a dolain of grwth in whih a laf pri-
mordium will aris only if th onntration of n inhiitory fill in this
rgion falls l<lwll thrshold valr-r.ah rrw laf that dos form ats as a
sourof inhiition to nighoringtissu' prvntinganothr lf from fom*
ing. Howvr, th irrlriit<lr influn drass,ith inrasing distrr,so
that as lvs erlrv irnd mov aprt' th ar rgions whr laf formation
an irrititd.]t is th pattn of intrlrtionsin this slf-orgnizingpross
tht rsultsin th rrangmntof lavs.Dpnding on th initial stat of th
mristmand its gomtry' th pross gllertson of th thr n-re jor pat-
trns of laf arragmnt as th primary pattrn and othr pattrns as s-
ondary ons. Th trm usd in iology ro dsri this typ of sptial
pattrningpross irr dvloping organisms is lttorpbogtifild. In thni-
al trms' h of th liffrntpattns that ariss from th slf_orgirrrizing
drramipross ls alld an ttrL1t()r, rvhil-ris th stal soluti-lllto th
02 Beod the Drluiin Prdigm

morphognti fild for partiular initial and oundary onditions thatpro-

dus a partiular form.
What an this modl rval aout distintionsamong th thr majorphyl.
lotais atgoris?It is known that spiral pattrns our in just ovr 80%of
high plant spis. Th initial laf pattrn in th sdlings of all ths
spis is ithr on storag laf, as in monootyldons, or two' as in di.
.lfhat
otyldons. mrgs from th work of Douady and oudr is that with
ths initial onditions on th fild organization of th mristm, spiral phyl-
lotais is th pattrn that is most assile(it is th attrator that lisonth
main ranh of th dynami gnrator).Furthrmor' th divrgnangl
btwn sussivlavsonvrgSon th osrvd angl of 137.5. onthis
ranh of th pattrn-gnrationpross.Th othr spiral pattrnsosrvd
infrquntly, with diffrnt divrgn angls' ar assil,ut thy rquir
that paramtrs in th modl hav spifi rangs' as do rar whorld at-
trns. Diffrnt plant spis an thrfor hav diffrnt phyllotais pattrns
as a rsult of gnti influnson initial onditions and paramtrvalusin
th mristm, ut thr is an ovrll ias in th dirtion of spirals,as ob.
svdin ntur-

oRPHoGN| FlELDS PRoDU FoRMs WlH

MAHEA|AL RGULAR|ls

Plants with spiral phyllotais hav a haratristi mathmatial ordrin

th spatial rlationships of thir lavs.Som mpls of ths ar givnin
Figur 2, whih shows th spiral arrangmnts of lavs in ruri els,
a mmr of th onke Puzz|e family. Th numrs on th lavsrfrto
th squn in whih thy wr gnratd, 1 ing th yungst laf or th
last to gnratd,2 te prvious laf, and so on. Th spirals in this figur
ar drawn through lavs that mak dirt ontat with ah othr. Now look
at th diffrn twn th numrs of sussiv lavs along any spiral.
In Figur 2ate diffrnsar 8 along th spirals dfind y dottd linsand
13 for diffrns along spirals with solid lins. This dfins what is knownas
(8' 13) spiral phyllotaxis. Th othr two xampls in th figur hav (5,8)
and (, 5) spiral phyllotais (Figurs2 and 2' rsptivly).Th numrsin
ths pairs,3' 5, 8, and 13, long to a wll-known mathematialsrisd-
find as i * 1= n + _| ah sussivnumr of th srisis th sum of th
two prding numrs. Th ptiulr numrs of th sris dpnd on th
two numrs that start it. If w gin with th numrs 1, 1' th srisis 1,1,
2, , 5,8, 13,21,34, 55, and so on. Most plants with spiral phyllotais
long to spirals dfind by sussiv numrs in this sris, as w sawln
Figur 2 for th onky Puzz|e tr. Douady nd oudr (1996| showd
that ths ar th spirals that aris on th main ranh of th gnrativpro-
ss in thir mdl. owvr, spirals with numr diffrnslongingto Figure 2. Leaf arrang
o t h r F i o n a i s r i sa r a l s o f o u n d i n n a t u r ' s u h a s ( 2 , 2 , 4 , 6 , 1 0 , 1 6 , Aruri ls. See
2 6 , . . . ) , ( 2 , 5 , 7 , 1 2 , 1 9 , 4 \ , 6 8 , . . . ) , n d ( , 4 , 7 , 1 0 , 1 7 , 2 7 , 4 4 ,. , , ) .
Ths an aris from th dominant spiral ranh y spifying partiular initil
 Bond the Drtuilti Prdi,qm 0
tl nd oundary ondititlns that pro-

intions rnong th rhr rrlajor phyl-

al pttrrls<rurin just ovr 8" >|
)attrn in th sdlings of ll ths
t nlnoltyidorrs, or two' s in di-
k of Douadv and Coudr is that with
niztionof th mistr,spiral phyl-
il (it is th attrator tht lis on th
) . F u r l r r m r t'h d i v g n n g I
th lsrvdangl of |7.5" on tis
;s.Th oth spiral pattrns osrvd
ngls,r ssil,ut th},rquir
:ifi rangs,as do rar whorld pat-
lr hv diffrntphyllotxis pattrns
rl onditionsnd paramtr valus in
ras in th dirtion oi spirals, as o-

)DUFoRs WlH
s
:haratristimthmatial ordr in
S o m a m p l so f t h s r g i v n i n
]mntsof lvsin ruri ls,
,. Th numrs on th
lvs fr to
rtd' 1 irrgth ylungstlaf or th
',
and so on. Th sirals in this figur
,tontwith
ah othr. Now look
of sussivlavslolrg any spiral.
.hspiralsdfindy dottd lins and
c lins.This dfinswhar is knor,vnas
'o ampls
in th figur hv (5, 8)
rnd 2' rsptivly).Th nLrmrsin
wll.known mthmatial sris d.
lunrrof th sisis th sum of th
numrsof th sris dpnd on th
:hth nunrbrs1, 1, th sris is 1, 1'
ost plants with siral phyllotais
t u n r r si t l l i s s r i s ' s w s w i n
) o u a d n d o u d r ( 1 9 9 6 ) s h o w d
: m i n b r n n ho f h g n t i v p o -
th nlrrnr diffrns longirrg to
Figure2. Laf arrarrgmntson ranhs
n a t u r 's r r ha s ( 2 , 2 , 4 , 6 , 7 0 , 7 6 , of spisof onky Puzzl tr,
ruriecls.S tt for disr.lssiorr.
a n d ( 3 ,4 , 7 , 1 0 , 1 7 , 2 7 , 4 4 , . . . ) .
lranhy spifyingpartiular initial
101 Bod th Dt'ii Prdigm

onditions and pramtr vlus in th mdl of Douady and Coudr. Th and Coudr. Th |at
rar pttrns <lfr,vlrorldlaf arrangmntstht ar distintiv to partiular morphogn from la
spis ar lso gnrtd y th rndl lvlrrrspifi oundry orrlitions nw laf to form t a
and paramt vls:rr spifid, f<lringtlrs mods tl ppar' though vidn prsntd y
tlr a lss stal tl-ranth dtlllrirlnt ons in trms of pkig dnsitof dution of primordiur
l:rvs. auin that gtstransp
mation in its nighr
is thus rplad y rr
RLA|VE sABIL|Y oF DlFFRN PHYLLoAI FoRs
inhiitory tion y
A l l t h p a t t r n sg n r a t di n t h m o d l < l fD o u a d y a n d o u d r a r s t a l vtd so that instad .
flr srm rang ()i Lroundary and initial orrditions on th morphognti laf formation in a sl
fild in th tnristm, l)tlt som ar lss snsitiv to hangs in thsthan phogn tht indus l.
othrs' Agin, th ]onrinan t a t t r n s f F i g u r l a r t h o n s t h a t a r l a s t nw laf and its boun
snsitivt viritilnsin th gnrativonditilns in th modl, ad among nw laf inititation irr
ths th spial pattrns ar th most roust. This fild modl of th dy- a fild of inhiition
nmis of laf podution in th mristm thus provids us with a possibl rns whr th paks o
planation of how all th diffrnt laf pattrns an gnratdin highr port of th morphogn
plants; tht is, it givs us a plausil pittrr of how th diffrnt pattrns Th lgnt studi
ar possil forms of th sam dynami pross undr variation in th gn. mophognti filds iI
rativ onditi<lnstht an rlatd to diffrrttgn arivitis and ound- tions. larly thy d r
l l r v o n d i t i o n s l n t h n r r i s t m( m < l n o l tasn d d i t s ) .I t i s n o t n s s a r y o r modl of th whol
appropiirt tO tls th onpt tlf ntual sltion or diffrntil {itnss of dirtional-fild ompt
th harats to plain how th osvd rang of forms ould hvorig- tial loation of uin 1
i n a t d i n p l a n t s . W h t t h m o d l t l l s u s i s t h a t w i t h i n s o m p l a l r tl i n a g that is loalizd to th
lr linags thr volvd within th mristrn a dynami pross with th t r a n s p o r t i n ga u i n p i
gnrtiv haratristis dsid. This modl gnrats thos arrdonly vasular lls of th d
thos pattns lsrvdin known plant s-lis. Thr ar many pattrns n that undrlis th
tat w n inragirrtht might produ1,suh as a singl laf at a nod, to explaind in trm
trvo rotatd at 90" tO this ilt th nt nod, tlrr at th nxt rlttdrlativ morphognti fild. lV
to ths t.uvo,irnlthn rpat of this s]un.Ths ar nvr osrvd. aross ll walls, ll d
Biologial foms ar not sltd fror ny imaginabl st or i] ontinuum m i s t mt o p o d u t l
of possiilitis. Nonlinar dynami prosssof th typs that or'rrin d- osrvd in laf phyll
vlopmnt always gnrata disrt st Of forms assilto th dynami sk to provid n int
prosssinvolvl.It is not natural sltion that dfinsths pssibilitis, Pross. Clarly any r
ut th intrinsi l,vnamisof th gnrtiv pross in th dvlopingor. ahiv th sam rangr
ganism. Undrstnding ths rnorphognti prosssis what providsus nd thi r transt.ormar
,ith n undrstnding of th origins lf spis s distint forms assibl
w i t i n a l i n a g ( G l o l w i n 1 9 9 4 ) . DvLoPMN
WN oRl(
F|LDs AND MoLULAR GNRAoRs
I now turn to th issu
A rnodl suh as that of Douady and oulr shows what priniplsarr. plaining aspts of phy
quird t<lprlvid a andidat xplanatiln <lfphyllotais, ut it oprts t usd to dfin th ada
quit a hig lvl oi stration.Rnt stulisy Rinhardt t al. (2003)in- ism funtions in a par
diat th ntur lt th rrrlulsinvolvd in grrratingphyllltatipattrns with othr nrmrs o{
nd rv| fiId 1r.trirtrri
diffnt in dtil frorrrtlrat essumd y Douady t o t h h a r r e r so f n

modl of Doudy and oudr. Th
n t st h l t l r t ] i s i l l t i v t t l p r t i u I r
I whn spifi oundary onditions
.ingths
mods to ppa, though
otls i trrs of paking drrsity of
jRN PHYLLoA|
FoRs
:l of Drludyand oudr r stal
al nditions on th rlrplrognti
is snsirivt hng.sin ths than
f Figur 1 ar th ons tht a lst
ondititlnsi th rnodi, arld along
roust. This lild modl of th dy-
mthus provids rrs with possil
f pttrnsan grrrirtdin higlrr
riturof how th diffrnt pattrns
j p r o s su n d r v a r i a t i o n
l diffrrrtgn ativitis nd ound-
ltsand diots).It is nlt nssary or
ral sltionr diffrentil fitnss of
lvd rng of fortns ouid hav orig-
us is that within som plant linag
lristm dvnai
oss with th
his modl gnrts tlros nd oniy
rrt spis. h ar many pattrns
l L r ds,u h s a s i n g l l a f a t n o d '
ld, thrt th nt rottd rlativ
squn.Tlrs ar nvr osrvd.
a n v i r n l g i n l | \ t o r i l o l l t i n u u m
rsssof tlr typs tht our in d-
:t of flrmsssilto th dynmi
,tionthat dfins
ths possiilitis,
]tlvpossin th dvlopirrg or-
]ntiprosssis wlrt provids us
. spis
s distintfors ssibl
RAoRs
udr slrows what prinipls ar r-
-'n ot phyllotaxis, ut it operts at
tudis1.Rinlrardt t l. (2003) in-
d i n g r l r l t i n gp h y l l o t
tail from that assuld y Duady
Be'odtb Dru,,iittPrdigm 0s
andoudr. Th lattr assumd an inhibitory
influn,possily arrid y a
morphognfrom a laf primordium, that
drasswith distan,allowing a
nw lf to form t ritial distn from
a dvloping lf. Howvr' th
vidnprsntdy Rinhardt and ollagus
is that th influn is an in-
dution of primordium formation y auin.
A nw lf ts s a sink for
auin tt gts tanspotd away from t sit,
thus prvntin8 nw laf for-
mationin its nighhood.Th irrhiitio.'
Dou,jy and Coudr
isthus rplad y rmoval of an indur from ",.u,n.dy
th nw laf rathr than dirt
inhiitory ation y a morphogn. The fild
rul is thrfor fftiviy in-
vrtdso that instd of prmissiv rgions
of low inhiition that all<rwnw
laf fomation in a salar fiId, w hav lomins
f lvatd auin, a mor-
phognthat indus laf initiation, and
transport of uin way fom th
n w l f n d i t s o u n d r i si n t h n r i s m
s r h l a f d v l o p , .p . . u . n . i n 8
new laf inititation in its nighorhood. Th
dynami. i, -o.. .omplx than
a fild of inhiition austhr is now
dirtionl olnponnt that gov-
rnswhr th pks of auin ur in th fild,
arisirrg from polarizd trans-
port f th rorphogn.
Th lgant studis of Rinhardt nd ollgus
tk th qustion of
in th gn- morphogntifilds in plnts in sonre vry intrsting
and lrallnging dir-
tions.larly thy do not plain phyllotaxis,
whih quirs oth ohrnt
modl of th whol and an xprimntally
valid moulr dynami. h
dirtionl-fildomponnt tht undrlis uin
flow is govnd y th spa-
tial loatin of auxin pumps in th ll mmran,,u.
u, PIN1, a protin
thtis loalizd to th apial sid of lls in
th orrtrlayrs of th mrsttn,
trnsortingauxin apilly' ut orintatd for
asa1transport in dvloping
vasularils of th dvloping laf. hus
thr is fild of ..g,,l,to.y influ-
nthat undrlisth prodution and loalization
of PIN1 in lls and nds
to xlaind in trms of molular orgaizing
fors in ordr to modi this
mophogntifild. Modling th los oupling
twnorintd pumplng
arossll walls, ll diffrntiation,and ,.'"in
flo- attrns througout th
mristmto prdu th ohrnt marosopi
pattrns of lf arragmnts
obsrvdin laf phyllottrxis prsnts n intrsting
hallrrgfor thos who
sk to provid an intgratd-r'rndrstanding
of ti, .nbu,]t morphognti
pross. larly n1, modl that sks
ro plain ths phnomn ,nust
ah,ivth sam rang of intgratd undrstanding
of phyllotati pattrns
andthitransformationsas th modl of
Douady nd Corrdr does.
DEvLoPMN AND voLUloN: H
RELAIoNsHlP
BWN oRlGINs oF FoRt"ls AND HlR
FRaUNlEs
I now turn to th issu of natural sltion
and what rol it might play rn -
pliningasptsof phyllotais. Fitnss t
hartat is th onpt that Darwin
usdto dfin th adativ.fatursof organisms,
that is, how wll a organ-
ism-funtions in a partiular nvirnm.
t i p t t r n s ,o that it survivs t rprodu
with othr mmrs of its spisand lav
offspring. Fitnss an applied
to thharatrsof n organism (.g.,pattn
of lvson th stm of hihr
Bod the Drtuii Prdigm

plants' pigmntation pattrns and shaps of lavs)and to th lif yl as a it is a disadvantg

whol. Natural sletion is gnally takn to dsri th pross whry trmins liffrnti;
organisms with hirratrsadaptd to thir haitat aris in th ours of vo-
lution: organisms with diffrnt hrditary haratrs aris as a rsult of
gnti variation' and th rnor adaptiv (fittr) haratrs inras in fr- olltiv Bh
qrrny in th population rlativ to thos haratrs that ar lss fit. Th
haratrs osrvd most frquntly in spisar thrfor rgrdd as having Th hvio piltt
gratr funtional valr.lto th organism' so that th diffrntial aundan aspt of thir vo|
of a haratr is takn as a masur of its adaptiv valu to th organism.If sts to ftrri.rgfrr
w apply this rsoning to th frquny of phyllotaxis pattrns in th lavs lavs in t autum
of highr plants, W may onlud that spiral phyllotais is tlr pattrn with Just as th ar di
highst fitnss or utility to th lif yls of highr plant spis, sin this is ar diffrns in t]
th most aundant pattrn' takn to stalishd historilly y natural signifiant irr]i
sltion. s p i s .I s h l l n o w
What is th sltiv valu of laf arrngmntsin plants? A rasonabl trns s iolrlgial
suggstionis that lvssrv th funtion of athing light for photosynth- nami ordr in an l
sis: th diffrnt pattrns of lavs on a stm hav diffrns in th ffiiny dvlopmnt ar k
with whih thy ath photons' whih dpnds n how muh thy ovrlap In this hmr th
and shad on anothr. Studis hav shown that th spiral pattrn illustratd young' fdirlg and
y yua in Figur 1a has th last mutual shading whn light is oming from and kping tlr rt
dirtly ovrhad' sin lavs that a arrangd in rows nssrily shad ind olonis kpt i
on anothf (Figurs 1 and 1). This ould b a asis fo undrstandingth ativity pttrn am
diffrntial aundan of laf phyllots in tms of natural sltionand of ants. Whn ind
funtion. Howvr, whn othr fators ar onsidrd, this proposal lossits thir fllows or at
.sfhn
strngth. avrgd ovr th ours of a day' with th sun ouping all thm' it was showl
angular positions rlativ to th plant' th diffrns twn diffrnt laf of th trm (Glik
arrangmnts with rgard to shading om minimal. Furthrmor, obsr- rhythm throughout
.sh
vation of spis in diffrnt haitats shws that thr is no orrlation als? dos th
twn shad plants and spiral phyllotaxis, as might xptd if maxi- Nigl Fanks rrn
mization of light aPtur orrlatdwitlr spiral laf arrangmnt.Plant adap- in a olony of ants
tation to shad is largly through vriation in laf shap nd pigmntation; th young in t
th lattr inrass asorption of th light frqunis availal. Th ovrall ativity-inativity
vidn is that phllotati pattrn is sltivly nutral in plants, sin all about 10 minuts
th diffrnt pattrns ar osrvd in various light haitats. In this as dif- individually priod
frntil aundan, th prdominan of spiral phyllotaxis in lrighr plants, hamr ths }'l
ds not appar to b xplaind y natural sltionof a funtional hara- whol hiits :r r
tr. If th onlusion is ort that all phyllotati pattrSwork adquatly in th flaslring of fi
for th photosynthti nds of plants, thn spirals prdominat austhy full moon. <rwv
ar dvlopmntallymor assiland stal. Thr is nothing surprising ants from oltlnis <
or prolmati aout this onlusion. It mans simply that w nd to tak (7990), whih lso
ount of th rlativ dvlopmntal assibility and staility of diffrnt found that thir a
morphologial pattrns, whih will afft th frquny of hara.trs in spis hiitd dtminist
and nruryhav a primry influn in ass whr th hratrs ar sltivly thn, dos rgul
'W
nutral. annot simply assum that diffrntial aundn is always- viduals?
plaind y natural sltin.Of ours, it is lways assumed that if a hra- This pro[.llmwal
tr gnratd in a spis is not usful' it will not inras in frquny, andif modl of th intra

of lavs)and to th lif yl as a
n to dsrith pross whry
ir haitat aris in th ours of vo-
ary haratrs aris as a rsult of
(fittr)haratrs inras in fr-
ls haratrs that af lss fit. Th
is ar thrforrgardd as having
, so that th diffrntial aundan
s adaptivvalu to th organism. If
of phyllotais pattrns in th lavs
piral phyllotais is th pattrn with
of highr plant spis,sin this is
stalishdhistorially y natural
angmntsin plants? A rasonabl
n of athing light for phtosynth.
m hav diffrns in th ffiiny
lpnds on how muh thy ovrlap
rnthat th spiral pattrn illustratd
l shading whn light is oming from
arrangd in rows nssarily shad
uld a sis for undrstanding th
s in trms of natural sltion and
r onsidrd,this proposal loss its
of a day, with th sun oupying all
r diffrnstwn diffrnt laf
om minimal. Furthrmor, osr-
shows that thr is no orrlation
axis, as might ptd if mai-
spiral laf arrangmnt.Plant adap-
i o n i n l a f s h p a n d p i g m n t a t i o n ;
ht frqunisavailabl. Th ovrall
ltivly nutral in plants, sin all
rious light haitats.In this as dif-
lf spiral phyllotais in highr plants,
rral sltin of a funtional hara-
hyllotati pattrns work adquatly
rnspiralsprdominate aus thy
l stlT . hr is noting suprising
' mans simply that w nd to tak
ssiility and staility of diffrnt
th frqunyof haratrsin speis
:s whr th haratrs ar sltivly
diffrntial aundan is always x-
it is always assumdthat if a hara-
will not inrasin frquny' and if
Bod th Drinin Prdigm 07
it is a disadvantag,it will nlt surviv. Howvr, it may not utility that d-
trminsdiffrntial aundan.
olltivBhavior in Ant Colonis
Th havior pattrns of orgarrismsar larly an important and intrsting
asptof thir volution. Th abilitis of fish to swim, of irds to fly, of in-
ststo forag for food' and of plants in tmprat limats to drop thir
lavsin th autumn all ontriut signifiantlyto thir apaity for survival.
Just as thr ar diffrnstwn spisin thir morphologis, so thr
ar diffrnsin thir havior pattrns that allow us to rogniz thm as
signifiant indiators of th distintiv lifstyls and haits of diffrnt
spis.I shall now tak a look at th way W an undrstand havioral pat-
trnsas iologial forms. For this I us an xampl from th study of dy-
nami ordr in an ant olony. Th qun and th young ants at all stagsof
dvlopmntar kpt in a sp within th olony alld th rood hamr.
In this hambr thr a many workr ants that tnd th qun and th
young' fding and laning thm, moving thm out as th young matur'
and kping th rood hamr fr of dris. Rsarhrswho hav am-
indolonis kpt in laoratrishav osrvdth ourrnof a rhythmi
ativitypattrn among th workrs in th rood hamr of rtain spis
of ants. Whn individual workrs Wr osrvd, ithr in isolation from
thir fllows or at low dnsity so that thr wr fw intrations twn
thm, it was shown that thir havior was haoti in th thnial sns
of th trm (Glik 1998). Th qustion ariss:how dos a rgular priodi
rhythmthroughout a olony aris from th intrations of haoti individu-
als?Whr dos th ordr om from?
Nigl Franks and ollagus(Franks et,a|' 1990) oservdthat th workrs
in a olony of ants f th gnus L|ltothor who wr tnding th qun and
th young in th rood hamr xhibitd a wll-dfind rhythm in thir
ativity-inativityyls. Roughly spaking, th workrs wr inativ for
about 10 minuts vry half hour. Th first assumption ws that ah ant is
individually priodi in its ativity-intivity yl, and that in th rood
hamrths yls om synhronizd through intrations so that th
whol hiits a rgular priodi yl. Suh synhronization is wll known
in th flashing of firflisor in fmal monkys oming into strus with th
full mon. Howvr, whn ol (1991r\xamind th havior of individual
ntsfrom olonis of a spislosly rlatd to that studid y Franks t al.
(1990)'whih also hibitd rhythmi hvior in th rood hamr, h
found that thir ativity-inativity yls wr thnially haoti: thy -
hiitddtrministi haos, without any sign of a rgular rhythm. How'
thn,dos a rgular priodiity merg frlm th intration of hoti indi-
viduals?
This prolm was studid y Sol t l. (199) y onstruting a omputr
modlof th intration of ants in a rood hamr. ah individual modl
Bod th Drii Prdigm

ant Ws dsribd in its ativity-inativitybhavilras haoti, in ardan som mryosr lar

with th primntal vidn, and th modl ants intratd with ah dnt of how th worl
othr by itation,again in aordanwith osrvation:if an ativ ant in- S o l ( l 9 9 9 ) h e v s h o
trats with an ativ or an inativ ant, oth ass rsult in inrasdativ- olony ativity, pri
ity. Th modl Was onstrutd as a kind of moil nural ntwork in whih r thn horipa
ants playd th rol of itl nurons that ould mov arollnd on a grid mi yls of ativit
that rprsntdth rood hmr. lt Was not possibl to prdit th out- ltd: thy enhan
om of th modl, sin thr Was no rason to assumthat haoti individ- whn thy ar dvl<
uals that intratd y xitation would dvlop a ohrnt rhythm. p r o a l y w h y r h s l
Howvr, that is prisly what ourrd. }.urthrmor,th modl also on- Flowvr, it is l
formd to osrvations that wr not uilt into th modl: a rgular prio- of th rhythmi ati
diity mrgs in th ant olony only whn th dnsity of ants dsa t i o n o f h h y r h mp l
ritil lvl, as osrvd y ol (199|) in his primntlstudis.Th p- undrstnding of hov
riodiity mrgsquit suddnly aov th ritial dnsity;in thnialtrms, w i t h t h p r o p r t i s(
it has th haratristisof a phas transition, lik th suddn ondnsation orgiztio i Biol'
of a gas to a liquid at a ritial tmperatur,so that on kind of ordr sud- rvlations of slf-or
dnly rplasanothr. It is intrstingto ask what kind of planation of th srvd in natur dos
mrgntproprty this modl givs us. aris from th rpat
units.'' A4odling th :
sKlNG AN EPLANAIoN FoR H FoR o f t h p o s s i i I i t yo f r }
shows that th dnsit
Th mrgntpriodiity in th rood hamr annot rdud to th b- of mximum dnsiy
havior of th individual ants and thir intrations in any spifi ausal a wid rng of i
sns.That is, knowing aout havior of individual ants and th natur of nomnon to aris in
thir intrations is not suffiint to prdit th priodi havior of th m u s t o u s i n h r
whol olony. Nvrthlss,this priodiity is a onsistnt proprty lf th tr valus in th mod
modl aov a ritial dnsity of ants. Hr is a as of onsistnywithout t o w i d a n g f g
ausl rdution, whih is found in many assof mrgnin omplx sys- fat tht th rhythm
tms. It my wll that mathmatial thorms will disovrdthat al- that it is not sltd
low on to mk prditins aout suh systms.Howvr' ths wr not t h g r a d u a I i s tv i w l
.What
known at th tim th modl Was onstrutd. th modl maks laris all iologii poprt
th nssary proprtis for suh an mrgnt pross. Th mathmatial trnsitions. ut omp
funtions usd in th modl dsri ths proprtis without plaining fatur of th way i
what thir onsqunsar. This is typial of sintifi produr: models nd Goodwin 2000).
oftn dsriwithout giving ausal xplanations of th phnomna.
Suh modls ar vry usful in larifying varius asptsof mergntprop.
MRGN..o
rtis in omplx volving systms.In partiular, W an ask what rol natu-
ral sltion might play in th rnrgnof rhythmi ativity of workrsin It is ths spontano
th rood hamr. To xplor this' w nd to ask what funtion rhythmi l 9 9 5 ) a I l d . . o r d fr.
ativity has for th olony and how it srvsits survival. Franks t al. (1990) mi morphology or
suggstdwhat this might . H and othrs hav osrvd that if an ativ in omplx living syst
ant nountrs an mryo r a larva that is alrady gtting attntion from ditions aris y a rand
othr workrs, th ativ workr will go to anothr mrnr of th rood. d i v r s i f i a t i o nw i t h i n .
Thrfor, if workrs in th rood hamr ar tiv at th sm tim, thy inluding th ourrt
proaly distriut thir ar lvr th rood and th qun so that thris f r o m d i v r s i f i t i o no 1
littl dupliation of ttntion. If thir ativity pattrns wr haoti, thn vriation of ondition

hirvilr as hoti' in aordan
: nlodl nts intrtd with ah
v i t ho s r v a t i o l li:i a n t i v
rlthassrsult in inrasd ativ-
of moil nural ntwork in whih
that ould mov round on a grid
yas not possil to prdit th out-
lson to assumthi.rthaoti individ-
,uld dvlop a ohrnt rhythnr.
'. }.urthrmr,th model also on-
rilt into th modl: a rgular prio_
,.hrrth dnsity lf arrts xds
in his pimntlsrudis.Th p_
ritialdrrsit1'; ir-rthnil tms'
;ition,lik th suddn ondnsation
:ur'so that tln kind of ordr sud-
sk what kind of planation of th
R H FoRM
ramrannot rdud to th b_
intrationsin any spifi ausal
of individual nts nd th ntu of
dit th priodi havior of tlr
iity is a onsistnt proprty of th
[ris a as of onsistny without
l ssof mrgnin omplx sys_
thortrrs will b disovrd that l_
1 sstms.Horvvroths wr not
rtd.What th modl maks lar is
nrgnt pross. Th mathmatial
:hsproprtis witlrout plaining
il of sintifiprodur: modls
laniltitrnsof th phnomna.
ng various asptsof mrgent prop-
trtiular,W an sk what rol natu-
of rhythmi ativity of wokrs in
ndto ask what furrtion rhythmi
rvsits sr:rvivl.Fnks t al. (] 990)
thrshav osrvd tht if an ativ
r e t i s a l r a d yg t t i I t g t r n t i o n f o m
o to anothr mmr of th rood.
rrr ativ at th sam tim, thy
lrood irnd th qun so that thr is
tivity pattrns rvr lroti, thn
Bond th Drii Prdigm
sonrmryosor lrva might gt mor attntion than othrs simply y ei-
dntof how th workrs w distriutd in th ood hamr' Dlgado and
n t i n - 5o16(1999)hav shown that whn this is modld y omputr simulation of
olony ativity, priodi pattrns do indd produ mor vnly distriutd
aIthan haoti pattrns. So w lrav a possil planation of why rhyth-
mi yls of ativity in th rood hmr ar bnfiial nd may s-
ltd:thy nhan th survivl hans of th yourrg y virtu of good a
whn thy ar dvloping, and sr inras th fitnss lf th olony. This is
proalywhy ths ativity yls ar ommonly osrvd in suh olonis.
Howvr, it is lar that natural sltion in no sns plains th origin
of th rhythmi ativity pattrn in th rood hamr. Th possil fun-
tin of th rhythm plyd no rl in dsigning th modl' whih provids an
undrstandingof how rhythri ativity pattl]sa11mrg in n nt olony
with th proptis osrvd. As statd in th omprhnsiv volum Seff-
..On of th
orgiztio i Biologil Sstms (amazin t al. 2001, 89):
rvlationsof slf.organization studis is that th rihnss of struturs o.
srvdin natur dos not rquir a omparal rihnss in th gnom ut an
risfrom th rptdappliation f sirnpl ruls y larg numrs of su-
units,''odling th ant olony as omplx systlll provids a dmonsttion
of th possiilityf rhythmi havior in th rood hamr. Furtlrrmor,it
showsthat th dnsity rquird for rhythmi ativity is quit low (aout 20%
of maimum dnsity)' as osrvd xprimntally,.rrrdth rhythm riss for
a wid rang of xitaility of th irrrs.This is what is rquird for ph.
nomnonto aris in volution: it must lur fairly radily ( assil)and
must roust in th snsthat th should a orrsidralrang of param'
trvlus in th rodl ovr whilr th phnomnon ariss. This orsponds
to a wid rang of gnotypsin whih th proprty an aris. In addition, th
fatthat th rhythnr riss suddnly as a phas transition in th olony means
that it is not sltdgradually, with progrssivlyinrsing dfiniion, s
.W
thgadualistviw of volution would hav it. rtinly annot say that
all iologialproprtisaris in this suddn, unxptdmannr' as do phas
transitions,ut omplity thory is dmonstrating that this is a ommon
faturof th way iologial organization mrgs(sGoodwin 19941 So1
ndGoodwin 2000).
MRGN,.oRDR FoR FR''
It is ths spontanOus'roust mrgnt proprtis that Kauffman (1'993,
1995)alld ..ordr for fr'' in volr-rtion:
th prolution of som orgnis-
mi morphology or lravior mad lssily th ourrn of onditions
in omplliving systmstht allow th proprty to gnrated.Ths on-
ditionsris y a random sarh of possiilitisthat rsult from spontanous
divrsifiationwithin organisms oth of gns and of dynami possiilitis,
inludingth ourrn of haoti ativity' as w hav sn in ants' and
fromdivrsifiationof nvironmntl lnditions outsid th organism. Th
variationof on]itionswithin organisms and irr tlrir nvirol-nntsis what
t 1 0 Bttr]th Drtuii Prdigt

Dzrrwin ssurlt() gllrat hrlitirrl'!'arity within popr-rltionsnd th ( 1 9 5 2 ) , A 4 i t h i s o n( l 9

hbitt vrity t<lrvlril.rthy an adapt. N:rtual sltion mav plain th tion ar disr-rssdin (
diffrntial frqunismong spis of various forms that hv risnin tionary dvloprnnt
volution, suh as th ourrn of rhythmi tivity pattrns in th rood Gilrt et a|. (1996)' )
hanrrof various spisof ant. owvr, thr ar situatins in whih dif- thial prsptiv, r1
frntial aundan mav l.'plaind in tms of th dynamis of tlr gnr- of onvntional sin
ativ plssand th rllrtiv stailitis of th diffrnt forms produd.In tn Nture's De: Hel
ord to urrdrsndth phtronrnof volution, w nd to urrdrstand
oth th gnrativ1yrrmiprossstht produ th distintivharatr-
istis of spisin th 6rst pla (tlrigins)rrd th fftsof nattrral sltion BIBLIOGRAPH
Olr th frqunisof tl-rvarious hratrs.Th intgration of thsarasof
Arthur, \7.2002. Th
sttrdy, dvlopmnt an.lvllution, into a tlhrnt dynami pitur of th dvlopmntal iol
volutinry pross is l-lowlld volrrtionardvlopnrntaliologv. amazin'S.,J.-L. Dn
B o b a u2. 0 0 1 .S r
PrintonUnivrsi
o l , B . I . 1 9 9| . I s n i
Summary and nlusions Prodigsof tb l
Dlgado,J., and R. V. S
Form and funtion hilv always n ntral onpts usd in iology to olonis.Jourlof
'h
urldrstnd orgnisms ar-rdthir volution. partiular thor of volu- Douady, S., nd Y. ou
tion dvllpdy Drwin inludd oth, ut thr has n a tndnyto and III.Jourlof1
Franks,N. R., S. Bva
rplrasizth rol of ftrr-rtion in dsriing holv organisms ar sapdand
bhaviourwithin th
thir lrvior is mlldd lry l.tturalsltiln.A lrasi assumption lras n
tbtil Bill
tlrt biologial fort-rrsvilry ontinLlousl,virs ll rsult of random vrition in Gilrt,s. F.' J. lvI.opit
gns,arrlit is naturl sltiontht ..dids''whih partiular torms surviv dvlopmntal iol
nd ar favord. Any dislntinuitisof flrm twnxisting spisar thn G l i k ,J . 1 9 9 8 . h o s :
rgarddas a rsult of this sltionpross'whih thn omst sulptor Vintg.
G o o d w i n ,B . . | 9 9 4 ' h
in volution.
and Niolson.
T viwplint dsridhr diffrs fr<lmthis y prsntingvidnthat 207. Ntrc's]
tl.rrtlphologv and llvior pattrns >ftlrgarrismsoftn aris rsdistint Books.
f<lrmsrrd do not llrrgto a ontinuLlrnlf pssiilitis.Furtlrrln'th in- G o r d o n ,D . . | 9 9 9 , l
rrinsi frqunisnd stilitisof th fornrs podud an ontriutto th York: Fr Prss.
rlativ frquny of th diffrnt forms' s() tht natural sltion is not th Jan, R.V. 1994. Phllot
ambridg:amr
only fator involvd in plaining thir diffrntialaundan.This vidn
Kuffmn,S. A. 1993.
()l11sfrom osrvation of morphology, fr>-t xprimntal study, nd from uolution. Nw Yor
mtlilsthat dsri hlrv diffrnt morpht>logisand havior pattrnsn 1995.At Hrlt
ais ftlt dynami prosssrvithin nd trvIrtlrtanisms. ynard Smith,J.' R. B
hr is nl diffitrltyirr omining nttlrl sltionwirlr a disrtstof R. Lar-rd, D. Rup,
plssiilitisfor volLrtionto wok with. Distint forms still nd to tstd volution. Qurtrl
inlli, t. 20.Th D'
io thi survival valr"r,nd disrimination garding funtion iln md uoltio.mrid
among whatvr forms ar gnratd,rsultingin diffrntfrqunisof th ithison, G. 1972. P
frms orrlating with diffrnt haitats. llowvr, undrstanding th pri- Rinhardt,D., ..R. Ps
milr,vgnrativorigins of thsforms ndst distinguishdfrom sond- J. Traas,J. Frimi, r
,vsltion pr()sss.Ftlr edrs who wirnt to pursu th issus furthr, uln trnsPort.N.7
Salazar-Ciudd,I.,.}.Jr
god ovrviw .listtssilns t al. (200l), Arthur (2002),
inlud irt.nzrzitr formatinin dvlo
Wilkins (2002), anl N1inlli(2003). A nl,-lssidisussionof onstrintsis Sly,T. D. |995. Th
ayrrard Smith t al. (l985). Phyllotais is disussd in Snow and Snw Prss.

Varitywithin populations and th
Natural sltion may plain th
f v n r i u sf r l r l n s h r h v
:hmiativity pattrns in th rood
rr'thrar situations in whih dif-
trmsof th dynamis of th !]nr-
of th diffrnt forms produd. In
voluti()n'w nd to undrstand
at produth distintivharatr-
r nd th fftsof natural sltion
lrs. Th intgrationof ths aras of
a ohrnt dynami pitur of th
ionry dvlopmntliology.
ntral onpts usd in iology to
n. Th partiular thory of volu-
t, ut thr has n a tndny to
ling how organisms ar shpd and
tion. A asi assumptirnhas n
s e s u | ot f r n d o m v a r i
ids''whih partiular forms surviv
rm twnisting spis ar thn
lss,whih thn omsth sulptor
.om this y prsntingvidn that
f organismsoftn aris as distint
of possiilitis.Furthrmor, th in-
lrms prldudan ontriut to th
, so that natural sltion is not th
iffrntialaundan.This vidn
from primntlstudy, and from
lhologisar-rdhavior pattrns an
twnorganisms.
uralsitionwith a disrt st of
)istint foms stiil nd to tstd
on rgardingfuntion an mad
ulting in diffrnt frqunis of th
i. FIoWvr,undrstanding th pri-
dsto distinguishdfrom sond-
I Want to pursu th issus furthr,
a z i n t a l . ( 2 0 0 1 ) ,A r t h u r ( 2 0 2 ) ,
w-lssidisussilnlf onstrints is
lis is disussdin Snow and Snow
Beld th Druii Prdigm 11
(\952), ithison (\972), and Jan (1994), Soial insts and thir organiza-
tion ar disussdin ol (|99|), Sly (1995)' and Godon (1999). volu-
r i s n i n tionary dvlopmnt (also olloquially known as vo-dvo) is th topi of
Gilrt et aL' (1996). I disuss ths issus, trying to put thm into a roadr,
thialprsptiv,arguing for a holisti prsptiv that rquirs a rthinking
of onvntional sin t ahiv tru ologial harmony and sustainaility,
inNtre's Due: Iig ur Frgmeted ulture (Goodwin 2007).
lBLIoGRAPHY
Arthur,w, 2002' Th mrgingonptualframworkof volutionary
dvlopmntal iolgy.Ntur 415z 757-764.
Camazin, S.'J.-L. Dnubourg, N. R. Franks,J. Snyd,G. Thraulaz,and .
Boabau.201. Self-orgiztioni BiologiISstms.Printon,NJ:
PrintonUnivrsityPrss.
ol,B. J.199\.Is animalhviourhaoti?vidnfrom th ativityof ants.
Proedingsof th Rol Soitof Lodo, SerisB 244:25-259,
Dlgado'J.'and R. V. Sol.1999.Slf-synhroniztion and task fulfilmntin ant
olonis.lourl of TheoretiIBklklg 25: 433441..
Douady,S.,and Y. oudr.1996.Phyllotaxisas a slf-organising pross.PartsI, II,
and III.Jourl of TheoreticlBilllg1'78:25 5-312.
Franks,N. R., S. Bryant,R. Driffith,and L. Hrnrik.1990.Synhronisation of
haviourwithin th nstsof th ant Lptothorruorum.Bullti of
thmtilBiolog 52 597_612.
i o n i n Gilbrt,S. F.' J. M. Opitz, and R.A. Raff. 1996.Rsnthsizing volutionaryand
dvlopmntal iology.DulopmtI Biolog \7 : 357_372.
Glik,J. 1998. hos:b mzig Sielf the Uprditbl.London:
Vintag.
.!idnfld
Goodwin,B. . 1994.Holu the Leoprd hgdlrs Spors'London:
andNiolsn.
2007. Ntur'sDu: Hlig r Frgmtdultur. dinurgh:Floris
Books.
Gordon,D.NI. \999. ts t Work:Hotu lstolo ls rgized.New
York: FrPrss.
Jan,R.V. 1994,Phllotis: SstmtiStud i PItorphogesis.
mridg:amridgUnivrsityPrss.
Kauffman,s. A. 1993. rigis of rder: Self-rgniztil d Sltkli
uolutio.Nw York: Oford UnivrsityPrss.
1995.t Hom i the Uiurs.Nw York: Oxford UnivrsityPrss.
MaynardSmith,J., R. Burian,S. Kauffman,P. Alrh'J. mpll,B. Goodwin,
R. Land,D. Raup, and L..!olprt. 1985.Dvlopmntal nstraintsand
vlution.Qurtrl Ruitaof Biolog 60 265-287.
inlli,A. 2003. The Deulopmtof iml Form: tlge,orpbolog, d
uolutio.amridg:mridgUnivrsityPrss.
Mithison, G. 1972.Phyllotaxisand th Fibonaiseries.Sinc,196:27-275.
Rinhardt, D., .-R. so,P. Stigr'T. andl, K. Baltnsprgr, . Bnntt'
J. Traas,J.Frimi' and . Kuhlmir.2003. Rgulationof phyllotaxisy polar
auintranspoft.Ntur426:255_260'
Salazar-Ciudad' I.,J. Jrnvall'and S. A. Nwn.rn.2003. hnismsof pattrn
formationin dvlopmntand volution.Deulopmt1'30 2027-207.
Sly,T. D. 1995. Th Wisdom of the Hiu. amridg,A: arvard Univrsity
Prss.
12 Bod the Drluini Prdigm

Snow' ., and R. Snow. 1935. primnts in phyllotais: Diagonal siits through

dussat apies. Philosophicl Trsti-lsof th Rol Soiet of Lodo,
SriesB 225: 63_94.
1952. Minimum ra and laf dtrmintion. Proceedigs of the Rol
Socit of I,odo, Sris B 139: 545-.566.
5o16,R. V.' and B. . Goodwin. 2000. Sigs of Lif: Hotu ompleit Pruds
Biolog. Nw York: Basi Books.
Sol, R. V., o. \4iramonts, and B. . Goodwin. 199. Osillations and haos in ant Philoso
Though
soitis.Jourl of Thoreticl Biolog 16I: 343_357.
.Wstr'
G. ., and B. . Goodwin. 1997. Form d Trsformtioz. mridg:
amridg Univrsity Prss.
-h
Wilkins, A. S. 2002. uoltio of Dulopmtl Ptbtus. Sundrland, MA:
Sinaur Asslis.
Kim Stre

Th dvlopmnt of v
l t a l a h i v m n t st l
major hallngs:how
t i m t o p l a i n d 'l
ganism and nvironm
ths hallngs. Thrt
iota (spiallyif w t
thr ar tainl mi
Donoghu 2004; Dwl
somtims rathtakin1
1'996). vn so, w no\&
divrsity and adaptati
of organismsin popu
diffr, it drivs diffr
ariss in th population
portant part of any org
of diffrntiation. Diff
mntal htrognity.
volutionary ilog1
onrn planations
to xtintion y a mt
.!as
thir tintion a
m a l s ? S o m o n r n r
r s d m o r o | s ss
whn it stilizd or v
tant has mass tintio
nisms that gnrat ad
that produ vriation?
volutionary hang?F
i t t h a t s u h l m n t s.
phvllotais:Diagonal slits through
s <lith Rol Soiet lf Lldo,

oation' Prodigs of th Rol

,l-if: Hotu omplit Prudes

n' 1993. osilItins and haos in ant Philosophyof volutionar

Thought
61 34-357.
m d Trsformltio. amridg:

||tltIPtll s. Sundrland. lr4A:

Kim Sterln

Th dvlopmntof volutionary iology sin 1858 is on of th grat intl-

Itualahivmnts of sin. Th living world prsnts sin with two
major hallngs:how is its immns divrsity aross tim and at any givn
rim to plind, and how is th traodinry adaptiv fit twn or-
ganismand nvironmnt to explaind? Thr is no douting th sal of
thshallngs. Thr ar no rlial masurs of th divrsity of today's
iota(spillyif w tak into aount th divrsity of miroognisms),u
thr ar rtainly millions of tant spis of organisms (raaft and
Donoghu2004; Dawkins 2004). Likwis, th adaptivnssof organisms is
somtimsrathtaking in its omplexity and its xquisit prision (Dawkins
\996),ven so' w now hav at hand, in road outlin, an planation of oth
divrsitynd adptation. In rutal summary, sltiondrivs th daptation
of organismsin a population to thir nvironment,and usnvironmnts
diffr, it drivs diffrntiation too, taking advantag of whatvr variation
arissin th population. orovr' sin th iologial nvironmnt is n im.
potantpat of any organism's world, thr is positiv fdk in th pross
of diffrntiation.Diffrntiation rds diffrntiation y inrasing nviron-
mntalhtogneity'
volutionary iology has mny important opn qustions. Som of ths
ollrn xplanations of partiular vnts. !r th dinosurs rally drivn
to tintion by a mtor impt, or Wr thy on tlrir way out anyway?
Was thir xtintion a nssary prondition for th volution of th mam-
mals?Som onrn road pattrns in th volution of lif' as divsity in-
asdmor or lss srnoothly ovr tim, or hav thr n long priods
whnit stailizdor vn droppd? Som onrn mhanisms.ow impor-
tant has mss xtintion n in th history of lif? Ar th sltivmha_
nismsthat 8nratadaptation importantly onstraind y th mhanisms
thatprodu varitin? What ar th rlativ rols of sltionand han in
volutionaryhng? How important ar slfishgenti lmnts,and how is
it that suh lmnts ar usually nutralizd in ganisms' gnotyps? As

t1.t
 1 4 P h ilo soph of u oltionr Th o ught

othr essaswili hav mad iar' thsar important and diffiuit qustions' also in sltiorr.Tlr fi
ut thy sm to mpirial qustions. Sin philosphs do nt ollt sussful, but thy ar
data, what ould philosophrs hav to say aout volutionary iology tht puzzls aout th ntu
hlps us undrstand it? ow ould irlswrirrgsulr qustiot1sas ths rl- synthesis dpn1don :
vant to philosophrs'o proits? My prlit in this ssay is to ?lnswr yt it was r-rnlarjust h
ths qustions.I gin with th rlvanof philosophy tO volutionayi- hs th synthsis o]
ology; I orrludwith th rlvanof volutionary biology to philosophy. Nonthlss, th on
mor intratal irr th
within iologv. Th ml
Frmnt in volutionary Bilogy: onptual individualism, to th id
InnonLost ual organisms. For a
(1976) agud that sl
.Wiilianrs
h no-Dawinin synthsis w:rs dvlpd y th Ronald A. Fishr nd Dr'vkins
( 1 9 3 0 ) - J . B . S . H l d a n ( 1 9 3 1 ) _ S w lW
l r i g l r t ( \ 9 | , 1 9 3 2 ) t r i u m v i r a t i n mins its indir,idualism
th arly 1930s and was ompltd in Amria y Thodosius Dozhansky th fomidal wapon
( 1 9 3 7 ) , r n s t M a y r ( \ 9 4 2 ) , G o r g G . S i m p s o n ( 1 9 4 4 , 1 9 5 3 ) , a n d G . L d - plagi vrtrats-ar
yrd Stbins (1950) and in Britain y . B. Ford and his shooi (.g.,Fishr tion builds orplx si's
and Ford |947; ain and Shppard 1950, 1'952, 1954 Kttlwll t955, th form th pltform f
1956) for, during, and aftr \/orld.!arII. It bam th rivd viw of tion must linagsol
volutionay iology. lt was individualist, mirovolutionary,gradualist,and tags b prsrvd-tha.
sltiorrist.Th synthsisfousd on individual orgnisnrs:omptirivin. Yt organisnrs dtl not
trations tak plae etwnindividual organisms in loal popultions,not would sh vn if I
twn groups or spis. Individual organisms ar mor or lss fit; A lon of m would n
individuals prospr or fil. Th synthsiswas mirovolutionafy: th lrg- and ths mrks do not
sal hngs visil irr th fossil rord ar nothing ut th aumultionof dnt, I hl, itrst Oll }'
small-salehangs in loal popul:rtions.Th synthsisws gradualist:volu- thousands of gns in
tionary hangs in loal populations ar slow and inrmntl. orss lost of th gns I arry ar
thir tos and grw thi tth littl y littl and ovr long priods. And it was opid' that is, pliat
sltiorrist.han vnts piay :-toie in evolutionary hang. A potntilly ags rflt th diffrin1
favoral mutation might simply just nt happn, or it might happn to an mny gnrations and
organism that is fit ut unluky. But aording to th synthsis,natural sl- short and thin. }.or th l
tion is th lominant atlsal fator that drivs th volutionary dynamis of not itlst rpliators, th
populations. f]urrthi prospts
Th synthsislft fomidal thnial prolms unsolvd, ut th thory odis, th vlrilsin
of volution onivd this way smd rlativly onptually unprolm- tory of lif is tl s it as
ati. This onptual lirrity was not omplt. For mpl' undrstanding ing to sur th rsot
th natur of spis smd a prtiuiarly prssing prolm, for th vry Dawkins 1c76. 1982).
gradualisnr of th synthsis smd to srr!]gstthat th apparan of wll- his lin of thought l
dfind spis might an illusion gnratd y th lngth of our lif spans. to muh ontmporary
Thy sm stal and wll dfind only ausw do not liv long nough g n s I i t l na g n u i
to se linagshang and diffrntit.If w had n around to s th an- Thr sm ro ll t ls
stral at population gradually hang and diffrntiat into its array of gn sltiorr hav n
modrn forms, prhaps w would not think of lopards, jaguars, panthrs' l:rnstrspons wlul
and snow lopards as distint, namal units of natur. Thr wr othr is- tint and suprior altr
sus too. for xampl, whthr volution is gnuinly indtrministipro. prov thir rplition
ss ausof th rol of han. not iust in th snration of variation ut uild a wll-adptd

importantand diffiult qustions'
Sin philosophrs do not ollt
Y aout volutionary iology that
ring suh qllestiollss thse e rl-
projt in this ssay is to answr
: of philosoph,vto volutionary bi-
llutionary iology to philosophy.
onptual
.lopd y th Ronald A. Fishr
Tright(I91, 1932) triumvirat in
nriay Thodosius Dozhansky
mpson (\944,195), and G. Ld-
B. Ford nd his shool (.g.,Fishr
i0, 1952, 1954; Kttlwll 1955,
r II. It am th rivd viw of
mirovo1utilnay, gradualist' and
i v i d u a l o g e t r i s n r s:o m p t i t i v i n -
lrganismsin loal populations' not
organisms ar mor or lss fit;
was mirvolutionary:th larg-
] nothing ut th irumulation of
.h
synthsiswas gradualist: volu-
slow and inrmntal.Horss lost
and ovr long priods. And it was
volutionaryhange. A ptntially
happn, o it r-niglrthappn to an
ding to th synthsis,natural sl-
ivsth volutiorrarydynamis of
prolmsunsolvd, ut th thory
.lativlyonptually unprolm-
plt.For xampl, undrstanding
ly prssingprolnr, for th vry
ggstthat th irpparan of wll-
rtd by th lngth of our lif spans.
ausw do not liv long nough
w had n around to s th an-
and diffrntiat into its array of
ink of lopards, jaguars, panthrs'
nits f natur.The Wr othr is-
i s a g n u i n l yi n d t r m i n i s t i p r o -
t in th gnrationlf variation ut
P h iklslph of ullutitlr h ought ll.t
.h
also in sltion. fittr organisms in a population ar mor likly to
sussful,ut thy r not rtain to mor sussful.Thus thr ',vr
puzzlsaout th ll:ltur of fitnss nd its ausurlrll in volutiln. flr th
synthsisrlpnldon distintion tr,rr tlrl nd petd fitnss,and
ytit Was unlar just how this distintlOnWas to drwn (So 1984).
Thus th synthsisof volutilnarythory did pos onptual qtlstions.
Nonthlss,th onptualgogrphy lf volutionary iology atlrmuh
mor intratal in th 1970s and 1980s as th synthsis am undr fir
within iology. Th most systmatihllngto th synthsishas brrto its
individualism,to th ida tht the targtsof sltionar, lusivly' in1ivid-
ual organisms. For x:rmpl,G. . Willils (|966| and Rihd Dirr'"'kirrs
(1976)argud ht sltiondos not rll at rr irdividualorgenisr-ns t all.
\ililliams nd Dawkins laimd that tlr gradualisnr of th snthsis rtnd-
mins its individrrlism. ompl iologil adaptations-sllsoy systms'
th formidal wpons of prdatlrs,th powr nd stranrliningof grat
plagivrtrats-ar uilt inrmntally' n small stp at a tim. Sl-
tion uilds omplx systmsumulativly v prsrvingsmll improvmnts
that form th platforrn for furthr small improvmnts. So th targts of sl-
tion must lingsof nar-prftopis-flr only thus an smll 1vlrn-
tgs prsrvd-thilt prsistthrlughmany vls <lfsltion.
Yt orgtrnisnlsltrnt>tform linags'l' daughtr is not i1oP}.tlf m, nor
would sh b vl'l if I wr n asul rtur rproduing [r Ioning.
A lon of m wlrrldnot a nar-prft()py' for lif's aidnts rark m,
and thsmarks do not rappar on a lon' As a rsult of a hildhood ai-
dnt,I hav just ln y: a llnwould hav two. In ontrast' though' mny
thousandsof gns irr my daughtr ar opis of gns in m; likwis' rost
of th gnsI arry r opis of parrrtalgns.Gns, unlik orgnisnls,.tr
opid,tht is, rpIiatl,and thy do forrn prsisting linags.Ths lirr-
agsrfltth diffring susslf gs:s()rtar dp ni tlshy. lasting
manygnationsirnd isting in man'vo1rjg5in ah gnratiotl.othrs ar
shortand thin. Fo t most prt' thsdiffrnsar no aidnt:glls r
not just rpliatos,thy ar ativ rpliators. Thir own h:rratristisin-
flunthir prospts for repliation, normally through thir influrrln
bodis,th vhils in whih thy rid. Th ight Wy to think aout tlr his-
tory f lif is to s it :ls a giant struggle tw linagslf gns,ah try-
i n g t o s u r t h r s o u r sn d d f t l r r p l i a t i o n ( W i l l i a m s 1 9 6 6 , | 9 9 2 1
D w k i n s| 9 7 6 . | 9 8 2 \ .
his lin of thorrghthas aisd v,vdiffiu|t issus indd that r ntral
to muh ontnrPorarphilosophy of volutionary iology. In wht snsis
gnsltilna gnuin altrnativ ttl th individualism of th s1'nthsis?
Thrsmto at last thr answrs to this qusti()n'and th dfnlrsof
gn sltion hv nvr rally pikd thir rspns. h simplsr and
lnstrsponswtltrld to insist tht gn sltion is an mpirillr' dis-
tintand suprior altrrrtivto individualism. Thor-rghgns normally irrr-
provthir rpliation prospts y hlping tlr lth gns in an rryo
uilda wil-daptd ody that will r thm all intl th nt .r.ltion'
)1() P h ilo soph of u olutior h ough t

thy do not always rpliat so ooprativly.For ampl, som gnsan think of th iologial u
gt to th nxt gnration only if thy ar in an organism of th right sx. Th O n s u h p h n o m n < li
gns in th mitohondri in my sprm lls ar doomd, for mitohondrial linags mostly rpliat
gnsdsndonly in th fmale lin. y darrght'smitohondril gnsr ls? Givn tt thy do.
all opis of gnsin hr mothr. In many irumstnsorganisms ar fittst of organisms, in partt
if th s ratio of thir offspring is 50-50 or thrabouts. But gns that d- singl-lld stg? Dw
snd via only on sx hav no intrstin th ody of whih thy ar a part yls, and, indd, s<;m
having offspring of th othr sx, and ths gns ar undr sltion to dis- is possil to formulat t
tort th sx ratio in thir favor, vn though this is maladaptiv for th or- haratristi llf yl fr
ganism of whih thy ar a part. Many suh sx-ratio distortrs ar known sltion maks that qu
(Burt and Trivrs 2005). S-ratio gns and othr so-alld gnti outlaws xampls of th advanta
prompt th following rspons: gn sltion is prditiul differt from Th issu of gn sl
th synthsis. Gn sltion prdits that phnotyps that prss th fittst pt of th dat aout
gns will om mor ommon ovr tim; th synthsisprdits that ph. losophy of volutionary
notyps tht hratriz th fittst organisms om more ommon ovr that oopration is ntl
tim. Although ths prditions ovrlap sustantivly, thy ar not idntial. al systms ar ompos
any hav rspondd y apting that whn gns ar outlaws' W to hav in ways tht
should indd s thm as targts of sltion,ut thy tak ths to x- qustion, thn, is how n
ptional ass that do not undrmin th gnral validity of individualism. tion on th individuals tl
Sltion dos t on gns' ut only in vry spial onditions. So a sond David Wilson' StphnJ
way of dfnding gn sltionis to apt that it is, frrth most part' pr. at on individuals, ut t}
ditivly idntial to th synthsis, ut that it appals to distint undrlying spis (Sor and \rVils
ausal struturs. Twl thoris an lad to th sam prditions aout o- vidualism of th synths
srvaloutoms vn whn thy posit distint undrlying ausl prosss. gadualism. Arding t
lliott Sor and David \7ilson, in thir hallng to th synthsis, apt lif ar just aggrgation
that individualist viws of th volution of oopration an mak th right srv volution in tto
prditions, ut thy argu that ths misidentify its ausal asis. oopra. rabit immun systmsl
tion, thy say, volvs only whn and ausooprativ ats nhn th oming lrgr in dry s
fitnss of th groups that ontain ooprativ agnts (Sor and Wilson all th mhnisms of
1998). Likwis, with rgard to gn sltion, fit gns ar fit aus thy that ar hanging in sm
t on thir nvironmnt in ways that nhan thir proaility of rplia- and lots of tim to au
tion. Th usual mthod gnshav of nhaning thir rpliation prosptsis tns of th fossil rord
by taming up with othr gns to produ vhils, whih in turn rprodu' In a famous 1972 ess
rpliating th gnsthy host. Most of thsvhilsar organisms,ut pr- fossil rord dos not st
haps they an hivs, nsts, and symioti allians too. Fit gns mak thy maintind that th
wll-adaptd vhils, whih in turn rpliat thir uildrs. Th fat that trpolationism to tru
gns typially adopt this indirt Stratgy of rpliation rings th prdi- spis do nlt hang
tions of gn sltion into lin with thos of th synthsis,ut th various spis om into ist
piturs of th ausal strutur of sltion (gn' individual organism) r- thy split into daughtr
main distint. dos dsri th typla
A third lin is to rgu that gn sltionhs huristi advantagsovr th r o | | | 9 9 l ) . B u r i | : l d r
synthsis.Stritly spaking,it is rrot a diffrntviw of th ausal struturo{ xtrapolationism. If v
volutionary pross;instad, it nals us to s possiilitisand prolms hangs in loal popula
that th synthsisonals. Dawkins took this lin in Th teded Pho- than long piods witho
tpe (7982), his most important dfnsof gn sltion.H gav many x- is tru that thr ar l
ampls of volutionary phnomn that ar simply mor salint whn w ass; poptrlations an

ly. For xampl, som gns an
r an lrganismof tlr right sx. Th
ls ar doomd, fo mitohondrial
tlgtr'smitohondrial gns are
lirltmstans organisms ar fittst
tlrrabouts.But gns that d-
th od1,of whih thy ar a part
g'nsar undr sltion to dis-
g this is maladaptiv for th or-
h s-ratio distortrs ar known
nd othr so-alldgnti outlaws
tion is prditiul diffrent from
pnotypsthat prss th fittst
; th synthsisprdits that ph-
isms om mor ommon ovr
rstantivly, thy ar not idntial.
rat Whn gns ar outlaws' w
'in, ut th tk ths t -
gnralvalidity of individualisrn.
r 1 s p i a | o n d i t i o n s .S o s
)t that it is, for th most part' pr-
rt it ppalsto distirrtundrlying
o tlr sm prditions aout o-
stintundrlyingausal proSss.
l r | I n gro t h s y r l t h s i s 'a
lf ooprationan mak th right
idntify its ausal basis. oopra-
us ooprtivats nhan th
:rtiv tlgnts (Sor and Wilsn
:tion, fit gns ar fit aus thy
hn thir proability of rplia-
n i n gt h i r r p l i r i o np o s p
vl-rils, whih in turn rprodu,
:svhilsar rgatrisms,ut pr-
oti allianstoo. Fit gns mk
iat thir uildrs' Th fat that
y of rpliation rings th prdi-
of th synthsis,ut th various
n (gn,individual organism) r_
n has huristiadvantagsovr th
rntviW of th ausal strutre of
s to spossiilitisand problms
' this lin in Th teded Pbeo-
f gnslti<ln' H gav many -
a simply mo salint whn w
Philosoph1'of ullutior Thought )t/
tlrirrko{ th iolgizrltlnivrsas n nsml of ompting gn lings.
on suh phnomnon is th volr-rtionof th organism itslf. Why do gn
linagsmostly rpliat indirtl,v,via th ooprtiv onstrution of vhi-
ls?Givn that thy do' why do vhiles hv th distintiv haatristis
o[ organisms, in partiular, a lif yl that involvs rprodutiotr via a
s i n g l - l l sdt g ?D a w k i n s p o i n t s O u t t h a t i t i s a s 1 ,t o i m a g i n o t h r l i f
yls,lld, indd, sonr of ths ltrntivsirr al, rhough unommon. It
is possiltl forulatthis qustion out th oigin of organisms and thir
haratristilif yl from withirl th prsptivof th synthsis,ut gn
sltionmaks that qustion insapal, and Dawkins givs marry othr
xamplsof th dvntagsof th gn-sltionprsptiv.
Tlr issu of g sltionand its rlation to individulism is iust On as-
ptof th dt aout th units of sltionthat is now ntral to th phi-
losophy of evolutionary iology. F,volutionary iologists hav onr to s
that loperationis ntral to th volution of mplxity. ompl biologi-
l systmsar onlposd of individuals, and ths :rroftn undr sltion
to havin wys tlrilt undrmin th intgration of th olltiv. A ky
qustiol1'thn, is lrow nw lvls tlf oganizatiolrvolv in th fa of sle_
tion on th individuals tht mak tlrm up to go thir own way. lliott Sor'
o n d David lfilson, StphnJay Gould, and othrs hav agrdthat sltiondos
at on individuals' ut thy hav argud that it ats as wll on groups and on
spis(Sorand !ilson 1998; GoLrld 2.002).Furthrmor' iust as tlr indi-
vidulismof th syntlrsishas isd diffiult npt:rlissus,so too hs its
gradualism.Aording to th syntlrsis,lag-salpttrns in th history of
p t lif ar just aggrgationslf small-salhangs. Biologists an atually ob-
srvvolution in ation in loal populations: thy an osrv hangs in
ritimmun systmsas rits ar posd to a nw virus, or finh aks
bonringlarg in drv sasons.Aording to th graduzrlismof th syr-rthsis,
all th rhanisnrstlf volutionar1.hang ar rtrvork in loal popultions
tat ar hanging in small Ways ovr a fw gnrrtilns. ! just nd spa
and lots of tim to aumulat th rsults of ths mhanisms into th pat-
t si s trnsof th fossil rord.
In a famous 1972 essaNils ldrlgeand Stphn Gould argud tlrt th
..trirpolationist''
fossilrord dos not sLlpport this itr.tr'In partiulr,
thymaintaindthirt th tr of lif dos not hang nough ovr til for -
trapoltionismto tru (ldrdgand Gould 197z|. Thy argud tht most
spisdo not hang signifintly ovr most of thir lif spans. Instad,
spisom into xistnrlativl,vquikly and tlrn do not hng until
th1'splitinto dughtr spisor om xtint. Tlr tnt to whih this
dosdsritlr typil lif histtlr,vof a spisis still t>pnto dt (ar-
rolr|1997|.But if ldrdg and Gould ar right, tlrir pitur dos hallng
trapolationisrn.If volutionary pilttrns r nothing ut th aggrgat of
hangsin lol populations, w w<luldpt tO s gradual hang rathr
thanlong priods without hang intrruptdy sudlnspisformtion. It
is trr-r that thr ar loal prlsssth:lt n rsult in stasis in partir.rlr
ass;populations an trk tlrir lritat v shitiing glgraphillyrathr
 r
I

I
!

1 8 Phibsopb of uoltionr hought

than adapting in pla; adaptations an havioral rathr than morpho- volvd in rthinking
logial and hn n invisil to us. vn so, if th fossil rord is nothing tual issus. Th nt
ut population-lvl volution summd ovr larg quantitis of spa and tant prolm aout t
tim, w would not xpt stsis trl its dominnt pttrn (ldrdg 199.5; biology.
Gould 2002).
Gould has dvlopd a nror radial ritiqu of xtrapolationism that r-
jts oth its individulism and its gradualism. In a lassi artil Lwonin volution within
dfind th ida of a Drii popultio: a populatin of individuals that
diffr on from anothr and xprin diffrntial rprodutiv suss, In th past dad oI
and in whih dsndants inhrit thir anstors' distintiv traits. Suh intgration of voluti
populations volv undr natural sltion (Lwontin 1970). Gould argud multifatd. It aris
that spis groups form Darwinian populations (Gould 2002; Gould nd onnts vnts in lo
LIod 1999). Spis hav dtrminat origins and ndpoints. Dughtr spis. It ariss in dt
spisrsml thir anstors:ronstrutingphylogny would e impos- playd littl rol in l
sil if this wr not so. Spis ar diffrntially sussful.\Within a r- twntith ntury. As
gional iota som spis linags will diffrntiat rihly, radiating into tal iology rmains hl
many nihs' whras othrs will rmain rstritd in numrs and logi- ar oth volvd anir
al pntration. Gould dos not pt spis-lvlsltionto plain th logial sins and t
adaptations of organisms, for ampl, thir prptual systms, ut h thy nd somhw
thinks that sltion will plain haratristisof spisthmslvs:suh about human agn
proprtis s gnti variation, population struture' nd gographi rang I shall rifly introdu'
.shat
ar visil to spis-lvlsltion and hn n th targt of spis- is gn? Tl
lvl sltion. twn th lassial g
Gould's idas aout spissltionfa srious prolms. For on thing' and molular biolog
thr ar no onvining as studis of spissltionin ation. Morovr, sial gn of th frui
thr ar srious thortial hallngs to his idas. It is y no mans lar fft and was trans
that th inhritan ondition on Darwinian populations is satisfid.It is quns with oth
tru that th organisms in a daughtr spis rsmlth organisms in th DNA squn-hor,r
parental spis.But th spis-lvlproprtis of th dughtr-its distri- snsitiv to the loal
bution and population strutur-may not rsmlr thos of its parnt at off and an vn
all. Furthrmor, in thinking of spis sussand failu, w my not asn of rprssor
abl to distinguish drift from sltion.Th sussor failur of indiuidul rgions nd not ;
orgnisms is not always th rsult of sltion.Som mortality is th rsult not passd on wh
of unhappy aidnt rathr than ill dsign. Som rganisms ar fund y Numann-Held 199\
luk, dspit thir dsign flaws. Hn ralizd fitnssdiffrs frorrr ptd strat away from thi
fitnss.Sussdu to systmatifatursof th population's nvironmntis and instad think of 5
sussin virtu of petd fitnss,as whn a swallow dodgs a hawk. Su- itan: unlss sus
ss du to frak aidnts-ing lown y a storm to frightr that is that mak thm su
hading in th right dirtion-is drift. But th distintion twnptd adaptivly ovr tim i
and frak auss of mortality dpnds on th istn of a population of flow of auratly o1
vnts that is larg nough for thr to pilttrns in thos vnts.Howvr, tionary iologists oft
spis population sizs ar so small and yls of sltin r s fw that formation aout th
W may not al to draw th distintitln twnsltivand aidntal gns is ditd y nat
mortality. gnftion and unsui
I hop tht ths ampls hav gun to xplain th rlvanlf philoso- w an think of th 1
phy to volutionary thoy. It is rlvant us tlr mpirial issus in- aot organisms an<

bhaviorrrlrathr than mlrpho-
:n so, if th fossil rord is nothing
vr lrg qllantitis of spa and
c o m i n n tp t t r n( l d d g 1 9 9 5 ;
tiqu of trapolationism that r-
,lism.In a lassi artil Lwontin
: a population of individuals that
diffrntil rprodutiv sLlss'
l111stos' distintiv taits. Suh
n (Lworrtirr1970). Gorrld argud
l t i o n s( G o u l d 2 0 0 2 ; G o u l d a n d
origins nd ndpoints. Dughtr
lting phyl<lgrry would irnpos-
rntillr,stressful.Witlrin r-
]iffrntiatrihly, radillting into
rstritdin numrs and ologi-
lis-lvls.ltionto xplain th
thir pptul systms, ut h
:rististlf spisthmslvs: suh
n strutur'and gographi rang
ln n th targt of spis-
sriousprllrns.F.or on thing,
:issltionin ation. Moreovr'
l his idas. It is y no mi1nslar
nian populations is satisfid. It is
:isrsmlth organisnrs in th
)prtisof t dughtr-its distri
tot rsmltos of its parnt at
iussand failur, W may not
h susso failur of idiuidul
]tion.som rrrortalityis th rsult
4n. Sor lgnistnsar fund by
alizd fitnssdiffrs fom xptd
. of th population's nvirlnmnt is
[rna swallorvdodgs a harvk. Su-
n y a storm to a frightr that is
ut th distintion twt-rptd
ln th istnof population of
l pttrnsin thos vnts. Ilowvr,
[ ylsof sltion ar so fw that
on twnsltivnd aidntal
to xplin th rlvn of philoso-
nt austh mpirial issus in.
P h iloso1lbof uolutior Th ought 19
volvdin rthinking th synthsisar not larly sepaatdfrom th onp-
tual issus.Th nxt stion makes th sam point in disussing on impor-
tant prolm aout th rlationship of volutionary thory to dvlopmntal
biology.
volution within th Lif Sins
In th past dad or so a furthr issu has om inrasingly salint: th
intgrationof volutionry iology with the rst of iology. Th prlm is
multifatd.It riss with ology in idntifying th transmission lt tht
onntsvnts in loal populations with th volution and diffrntiation of
spis.It ariss in dvlopmntal iology, a domin of th lif sins that
plyd littl rol in th onstrution of th synthsis in th nriddl of th
twntithntury. As a onsqun'th natur of volutionary deviopmn-
tal biology rmains highly ontstd. It arises for humn iology, for humns
ar oth volvd animals and nulturd agnts: th prsptiv of th io-
logial sins and th prsptiv of th soil sins ar both valid, and
thy nd somhow to part of a singl unifid framwork for thinking
aout human a.ny(Strlny 2003). It riss for molular iology, and
I shall rifly introdu on aspt of this as.
!0hat is a gn? This qustion is hallnging aus th rlationship -
twnth lassil gntisof ndl and his twntith-ntury sussors
nd molular iology has turnd out to vry ompl indd. Th las-
sial gn of th fruit-fly primnts oth lrad a wll-dfind phnotypi
fftand was transmittd as a singl unit. Thr may b no DNA s-
qunswith oth of thos proprtis' for th dvlopmntal import of a
DNA squen-how and whn it is transrid-has turnd out to vry
snsitivto th loal gnti nvironmnt. Gns an swithed n and
off nd an vn rad in diffrnt ways, dpnding on th prsn or
asnof rprssors and promotrs lswhr in th gnom. Sin ths
rgionsnd not adjant to th DNA strth thy influn' thy m
not passd on whn that DNA squnitslf is rpliatd (Griffiths and
Numann-HLd 19991 Griffiths 2002). volutionary iologists oftn a-
strataway frorn thse molulr dtails of gn opying and xprssion
andinsteadthink of gns as infomation arrs.volution requirs inhr-
itan:unlss sssful organisms ar apt to transmit th haratristis
tht mk thm sssful to thir offspring' a population annot hng
adaptivlyovr tim in rspons to sltion.Inhritan is mdiatd y th
flowof auratly opid gns from on gnration to th nt' and volu-
t i o n a r y i o l o g i s t so f t n t h i n k o f t h s g n s s i n s t u t i o n s ,a s r r y i n g i n -
formtionaout th organism of whih thy ar a part. Sin th flow of
gnsis ditd by natural sltion,with stritalons making it t th nxt
gnrationand unsuital ons bing dltd by th dth of thir arris,
Wan think of th population gn pool s arrying information dirtly
about oranisms nd indirtly aout th nvironmnts in whih thos
20 Pbilosoph of uoltior Thought

ognisms hav livd and rprodud. Th suss of som gns and th

failur of othrs is a signal out th natrrr of thos nvironmnts (.!7illiams Why Bilogy att
19921 anard Smith 2000a).
This pitur of ross-genrati<lngn flow as a flow of instutions has gn- I hav n disussing
ratd vigorous dat. Paul Griffiths, Russll Gray, Susan oyama, and othr is rlvant to thshal
dvlopmntal-Systmsthorists think tht it is wonghadd in two rspts that would sttl (for
(Griffiths and Gray 1994; ama 1985). First, thy argu that this pitur twn maro- and mirt
ovrstats th importan of gn-asd inhritan. No on dnis that volution. on rol for
th flow of gns twn gnrations is on of th mhanisms that ar spify tIy what th
rsponsil for th rsrnblansbtwn parnts and offsping. But th is rlvant to vllutior
dvlopmntal-systmsthorists argu that mny othr mhanisms r impor- dd, it is rlvant in at
tant too: symbioti miroorganisms, hmials in ll ytoplasm' food trasin plantory agnda for
matrnal milk tansmitting th mothr's food prfrns'havioral imprint- hy. (2) It provids
ing on th natal sit' and many mor. There ar mny intrating mhanisms, must fspond. () It pr.
thy argu, that ar responsile for ross-gnrationsimilarity, and a good vo- adapt for its own prolt
lutionary dvlopmntal iology annot uilt if ths othr sptsof inhr- vry diffrnt from th r
itan ar ngltd(oyama t al.2001). a s d . I s h a l l d i s u s s( 1
Mrovr' although gns ar rtainly important, dvlopmntal-systms on (4) in th following s
thorists dout whthr gns arry information in any distintiv sns. 1. Lt m gin witl
Gns undnialy rry predictiue information aout phnotyps. Holding twntith-ntury philos
nvironmntl fators onstant' if you know th gns an organism has part of th natural wold
inhritd' in prinipl you an prdit its phnotyp. But all fators ausally prolm: to What xtn
rlvant to dvlopmnt arry prditiv information. Th plnt on whih and our world mpati
th uttrfly lays its ggs prdits faturs of th uttrfly phnotyp, namly' pI volvd iohmi
th plant spis on whih insts lik this kind of uttrfly will lay its ggs. W ar slf-awar; dlit
Th tmpratur of th tuatara's nst prdits th s of its offspring. Thus tional; and awar of m
although gnsdo indd prdit phnotyps,thy ar not alon in doing so. Prhps this pitur is t
Moreovr, if information is just a ovariation twnsignal and southat us as gn-rpliting rn
nabls us to prdit fatursof th sour from th signal, information n- hoi' rationality, or r
not misrad or misusd. It would mk no snsto say of ths ..instru- o m p e t i il i s t n r u r a I i s
tions'' that thy wr ignord or misrad. In rtain dvlopmntl otxts thy prhaps rquir sor
typial human gnotyps prdit tht th rsulting phnotyp has vstigial has dfndd th id tl
lims: in those ontts thy ovary with that outom. But \X/illiams,ay- hois (Dnntt200)
nard Smith, and othrs do not think that humn gns arry instrutionsto xampl, in this ih;
grow vstigial lims if onfrontd with thalidomid or tht aorns arry argud that th natural l
gns that tll thm to rot just aus most aorns prditaly rot. Thy truth'' is an illusion. Th
nd an altnativ notion of information that maks sns of th ida that oral lif systms xr
gns nd only gns dirt dvlopmnt along prditaltrajtoriswhn down th dstrutiv pu
th dvlopmntal pross oprats as dsignd, ut also maks sns of th nothing to do with it. Hu
fat that gnti information is misrd whn th gn-radingsystm mal- anstors;lthrsdid l
funtions. It has provd surprisingly hard to dvlop suh an aount (ss- p h y y n k i n gu p r h t t
pially th ommntris to aynard Smith 2000a: Godfry-Smith 2000; slf-portrait.
Srkar 2000; Strlny 2000; and th rply y aynard Smith 2000). Thus 2. volutionary iolo1
thr is still work to don to show that th volutionary onption of pls. uh philosophy l
gns and gnti inhritan intgrats smoothly with molular onp- most always thought
tions of th gn. philosophy, just as thy

h srtssof som grrsand the
"<lfthosnvironntnts(!7illiams
w as a flow of instrutions has gn-
sll Gray, Susan oyama, and othr
it is wronghadd in two rspts
First, thy argll that this pitur
ir-rhritan. No on dnis that
; on of th mhanisms that ar
:n parnts and offspring. But th
ml]).orhmhanismsar impor-
:als in ll ytoplasm, food trs in
od prfrns,havioral imprint-
ar many intrating mhanisms,
nrationsimilarity, and a good vo-
uilt if thsothr spts of inhr-
inrportant,dvloprrrntal-systms
) r t n t i o ni n a n y d i s t i n t i v S n s .
ration aout phnotyps. Holding
(now th gns an organism has
phnotyp.But all fators ausally
inforrnation.Th plant on whih
oi th uttrflyphnotyp,namly,
s kind of uttrflyrvill lay its ggs.
dits th s of its offspring. Thus
pS,thy ar not lon in doing so.
ion twnsignal nd sour that
-.fronr th signal. infornrtion an-
] no snsto say of ths ..instru-
In rtain dvlopmntal ontxts
: rsultingphnotyp has vstigial
that outom.But Williams, ay-
h u m l rg n s a r r y i t l s t r u t i o n st o
thalidomid or tht aorns arry
nost aorns prditaly rot. Thy
. that maks snsof th ida that
rlong prditaltrajtoriswhn
lignd,but also maks sns of th
lhn th gn-radingsystm mal-
o dvlopsuh an aount (ss-
nith 2000a: Godfry-Smith 2000;
,y Maynard Smith 2000b). Thus
at th volutiona onption of
sloothly with molular onp.
P hilosophtf uolutior hought J21
Why Biology Mattrs to Philosophy
I havn disussinghallngsto th no-Darwinin synthsis.Philosophy
is rlvantto thshallngs,for th is no singl -primntr prdition
tht lvould sttl (flr mpl) th fat of Gould's viw, of th rlations -
twnmro- and nrirovolr.rtirln or lf Drvkins's viw of tlre rtll of gns in
volution.on rol for philosoplr is to ring ths viws into sharp fous: to
spifyatly what they lim aout lif and its histlry.Just as philosophy
is rlvantto volutionry thory' tlrat thory is rlvant to philosophy. In-
dd,it is rlvant iI-rrtlast four wavs: (1) volutionilry iology 1rlpsst th
planatoryagnda for tln majo trovmntof twl-ltith-nturyphiloso.
phy. (2) It provids ruial st f xampls tl wlrih philosophil thoy
nrustrspond. () It provides an important st of tools that philoslphy an
adaptfor its own prolms. (4) It is n mplar of sinin ation, ut it is
vrydiffrntfrom th usual modls on whih philosophy of sinhirs n
a s dI. s h a l l d i s u s s( 1 ) , ( 2 ) ,a n d ( 3 ) i n t h i s s t i ( ) n n l s p n d t h m o s t i m
on (4) in th following stion.
l. Lt m girr with th plrrtory agnd. on major movmnt of
twntith-ntury philosophy has n naturalism: sing humans s wholly
partof th natural world. In tun, naturalismgnratsa diffiult philosophial
prolm:t what tnt is our ordinry' ommonsnspitur of or.rrslvs
and our world ompatiIwith th ft that w ar ntrtingmor thn Om-
pl volvd iohmialmahirrs?<lmmon st-ts viws humns ils a!]ents:
W r slf-awar;dlirativ nlakrs of ral hlis;rfltiv;oftn ra-
tional;and awar of moral onsidrionsand somtimsrsponsivto thm.
rhapsthis pitr.rris undrmind v an volutionry prsptiv that ss
us ils ]n-rpliatig mhins:as vlrilsuilt y and for gns.what ould
hoi'irtionality'o l-noality ut illusions if tht is tlr truth ir911152
ompatiilistn:ltllrlistsargu tht th two pirLrrsl onsisttr,th<lr-rgh
thyprhaps rquir som visions of ommon sns.Thus Danil l)nntt
hs dfnddth ida that volvd raturs lik us nonthlssnrak ral
hois(Dnntt 2003). But ompatiilism is rtinly not unhallngd.For
atrrpi' in this MihlRus (1986, Z07) and Rihd Joy (2006) hr,
rgudthat tlr rrarural histor1' of lrumn oprzrtiot-tshws us tlrat ..tiriai
truth''is an illusion. hr ar no rol fts for lrunrnsto rsptlsivto.
Moral lif systIllsist mly aus thy indu tlopration and damp
down th dstrutir.pursuit of slf-intrst;rognition of moral truth has
nothingtl do with it' Humans with oopratin-induingmorl systmssot to
anstors;tltrsdid not (Joy 2006)' So iologv is inrportant to plriltlstl-
phy y ranking r"rpth tnsion t'nvn or folk slf-lnptand <lltrsintifi
slf-portrait.
2. volutionary iology is also important as a sourr tlf ral-world xm-
pls.uh philosophy ryls a linritd st <lfar-npls,and ths ilr l-
most lwys thougt xprimnts' Thought xpinlnts hav ol in
philosophy,just rsthy do irl tlr sins.All tlroris mirk lirirns lut
22 h iloso1ll ll, of u ol u tilr T h ougbt

ountrfatual ass' and so it is appropriat to lnsidountrfatuals- S thos wh thirrk t

nrios. B thr ar good rasons for mor srilus :rttntioll to al plr- naturl wold nd to
nomna as wll. First, thought xprimnts ar oftn undrdsrid.Thy ship twn syrl ;
strat way from th rih dtail of ral ss. Morovr' marry of ths and his llvironment
thought primnts lal to l:rsh of irrtr.ritions, partly baus intuitiv n r i n d i s s 1 ' r l i oflo r s p i
judgmnt is snsitiv to just how ln dsris th thought -primnts. to this qustion. An intr
on st of ths imrgdsnarios involvs Str rh-styl tltransportIs targts in th world. P
and snrsto sh<llvthat prsonal lntiriuit,vis not th sm as th ontinu- drs. This is a nttrls
ity of our physial odis. But whil it is asy to tll ths storis so tht grams. Howv, almos
tltransporting sms to a way of atually travling whil laving on's to ojtivly sin-ril
old ody hind' it is also asy to rnk rh.tsm lik killing nrahins aaran of ral siri
tht rplrr th dupd trvlr with an impostor. Tlris snsitivity to th atd y th way hman
mod of prsnttiOnsuggststhat th dtails mattr. Vry th stting' nd othr thor'rghtsmuh m
you vary tlr jirdgnlnt. But if Grd is in th dtils, notlring ut th rl som mntl stats
.What
will do. not imaglik. rsl
Morovr, philosophrs hav oftn not fully onsidrd th rang of a. was th foundr f logi
tul ss. Asn is oftn vidn.Dnntt's u,ork is importnt irr this r. So th rlation tlv
spt. H shows how ignorirrgrality has allowd philsophrs to rptdly ship. But prhaps it is a
slid into sustituting a diotomy for a ontirruum y ngltingth rng othrs. Think in partr
of rl ass. For xampl, in thinkirlg aout rtionI agny, philso- r l - r i s r I i l I t
phrs hv oftn ontrastd ration:rl intntional agl-lywith mly m- n's tank. That is wh
hanial sYstms. Our flxil, informationlly snsitiv rspons to otlf So an intrnal statof Pt
rvold is ontrastd rvith th rigid ar:toratiiyof artifats nd ilrsts.Dn. th ing spid in h
ntt points out that volutionary iology shows that th must b a rangof i o n s h i p l r t w nl t n i n t
intrmdiat agntsl for informational snsitir,ityis a omplx adaptation, fatur of th nvironm
nd ompl adapttions do rrot aris from rigid systrsin a singl stp will prlably not dl itl
( D n n t t| 9 9 5 ' 2 0 0 \ . pption. Lots of my
. A furthr important rol for iology within philosophy is as a sourof win aout. But th ruI
t o o l s n d i d a s .T h u s D a v i d H u l l ( 1 9 8 8 ) h a s p p l i d v o l u t i o n vm d l s t o possil for us vr ttl
t h d y n a m i s o f s i n i t s l f ,a n d R t h M i l l i k a n ( 1 9 8 9 a , 1 9 8 9 b ) ,D a v i d P a - arahnophoi, nd hrs
pinau (2003)' and thrs hav attmptd tl us th iologial notion of will oftn think ..Spid
funtion to solv oll or polm of hilosopy: th natur of mrrtalsym- snting his nvir<rnrn
ols. Blifs and prfrnsar rprsnttior-rs of th world. But what in th symol-ta gt rlatiins
physial rvorld is a rprsntation?I liv that tigs ar th most hand- ..spidr''
thoughts lv
som mmrs of tlr at fanrily. Btrt what mrksm tigr thor'rghtsabol Hll thy r nt>tsvmL
tigrs? What maks rtin nural struturs within n-ryrain a smbol of gory of spidr look-ali
tigrs? Historially this was sn as a ig prolm for any physialist thory forth). And sin Ptr
of mind iruspopl an think aout nonxistntoits and bstratob- liks,h has mad no
its. Maning nnot a physil rlationship twn nrind and th David Papinau nd I
rvorld, aus thr ar n physil rlationships to angls or numrs. th rlationship twn
or rntly pilosophrs of mind lrirvnotid that tough prolms aris funtion. Th iologial
vn fo simpl rsslik rhoughts irout spidrs' for thr is nl simpl rla- o n al t t t n . h o s . g g
tionship twn spidr thoughts and spiders ing psnt to th thinkr' t o r s t O s p r l t a g i n s t p i
Ptr an lrav spilrthouglrtswithor'ltspidrs,for h irn tk nroth for a to lrth than thos tn
spidr; and h an fail to hirvspidr thoughts vn in th prsnof a spidr maks it tru to s1ythat
y thinking that spidr is ir moth. On any partiular gg t

)rlattO onsidr ountrfatul s_
mor srious ttntion to al ph.
nts ar oftn undrdsrid. Thy
'al ass.orovr, many of ths
i n t t t i t i < l tpl s.r t l y b
d s r i tsh t h o u g h x p
lvs S/.rr r e k-st|e tltransportrs
ruitis not th sam as th ontinu-
t is sy to tll ths storiS so that
,tuallytravling whil laving on's
k t-rsm lik killing mahins
tn impostor. This snsitivity to th
dtailsmattr. Vry th stting, and
in th dtails' nothing ut th ral
rot fully onsidrd th rr-rgof a-
nntt'swork is important in this r-
Lsallowd philsophrs to rpatdly
a ontinuum y rrgltingth rang
rg aut rationl gny' philoso-
intntional agny with mrly m-
lti<lnall1.
snsitiv rspons to our
natiityof artifts and insts. Dn-
y showstht thr must a rang of
snsitivityis ompl adptation,
tror rigid systms in ir singl stp
;y within philsophy is as a soure of
i) has pplid volutionry modls to
Millikan (|989a,1989), David Pa-
ptd to us th iologial notion of
hilosrlphv:th natur of mntal sym-
nttiosof th ''vrld.But wht in th
livthat tigrs ar th most hand-
vhat maks my tigr thoughts bout
lturswithin m,v rin a smbol of
ig prolm for arry physilist thory
: nonistntljtsand abstrat o-
rlatiorlshiptwn mind and th
rlationshipsto ngls or numrs.
v notid that tugh prolms aris
'rt spidrs,for thr is no sirnpl rl-
spidrs ing prsrrttO th thinkr.
spidrs'for h n tak a moth for a
lughtsvn in th prsnof a spidr
Philosopb of uolutior hought 2
So thos who think that humans ar oth thinking ings and part of th
naturalwlrld nd to plirin th ntur of mrrtlsymbols nd th rlation-
shiptwna syrnol and its targt.Y/hat fats out Ptr'srain, his lif'
and his nvirnmnt mk it tru that a partiulr intrnal fatur of his
a u s i n t u i t i v
mindis a sylnol for spidrs?Philosophrshav xplord a rng of rsponss
i m n t s . to this qustion.An intuitivly appalig ida is that symols rsembl their
targtsin th world. Ptr'smntal stat is aout spidrs if it rsmlsspi-
drs.Tlris is a natural su8gstionif thtlughts ar lik imges, maps, or dia-
grars.Howvr, almost rainly it Will not do. ntal imags do rrot sm
to b ojtivlysimilar to nduring thr-dimnsionalphysial animals; th
appran of ral similarity twnpiturs and things is an illusiln gnr'
atlb1'thwav lrumans irrtptpiturs.Ptr'sspidr imag wili rsml
thrthughtsmuh mor than it smlsany animal. orovr, vn if
sommntal stats ar lik imgs, surly most mntal rprsntatlonsar
not imaglik.Whirt is th inragthat ospondsto th thorrght ..Aristotl
Wasth foundr of logi, arrd ulid tlr foundr Of gomtry''?
So th rlation twn symol and targt is not a rsrlanrlation-
ship.But prhaps it is a usa] rlationship.Som vnts r natural signs of
thrs.Think in partiulr of instrurnts.An instrumnr is ful gaug if
thris a rlial latiorrtwn th rading on th gug and th ful in
one'stank. That is what maks a symol on th gauge man ..quartr full.''
Soan intrnalstatt>fPtr is ir spidr syrrrlrolonly if it lialy ovaris with
thrbinga spidr in his viinity. Th symol-targtrlationship is th rl-
tionshiptwn n innr strutrrrand its nvironmntal orrlation: th
featurof th nvironmnt that aussth innr struturto light up. But this
will prolynot do ithr. [n th first instan,this pitur applis i1t st to
prption.Lots of my win thughts our prisly aus thr is no
winaout. But th runh prolm is that this piur sms to mak it im-
possilfor us v to mk r-t-tistaks,
t surly w do. If Ptr is somwhf
arahnophoi'arrd his frirrdsplagu him with lvr imittion spidrs, h
rvilloftn think ..Spidr!!''whn thr is no spidr around. H is misrpr-
sntinghis nvironmnt. Yt aording to th ovariation pitur t.,f th
symbol.tagt rlationship' lr would no misrprsntirrgat all, ftlr his
..spidr''
thoughts ovry with th atgory of spids or imitation spidrs.
Hnthy ar n()tsymols of spidrs atir ll; thir targt is th roadr at-
goryof spider look-liks (inluding ral spidrs, ur spiders, nd so
forth).And sin tr thirrks..Spidr!!'' only in th fa of spidr look-
aliks,h has mad no mistak.
David Papinau and Ruth Millikan hirv suggstdan altnativviw of
thrlationshiptwnsyrrrbolsand rhir targtstht is zrsdon iologil
funtion.Th iologial funtion of, say, th pattrn on oystrathrggs is
onaImnt. Thos ggs are mouflgel.ggs that wr diffiult for pda-
tolsto spot against typil akgoLrndsof sand nd dris w mor likly
to hath than thos that w sir to find. Tlris sltiv histoy is what
maksit tru to say that th iologial funtion of th pattrn is amouflg.
on any partiular gg th pi1ttrnhas that iofur-rtionven if it dos not
24 bilosoph1'lf uolutirlnr Tbtlught

hav tht fft. T'h pattrn is suplsd

to orrl tlr gg vn if th eggrs
sn anyway, and vn if th egg is ths gnral prini
vry sily sn ..;;;. th parnt
laid it on whit sand irgainst*i. it has t i o n s o f n t t r s i
stands out. Similarlv, thotrghtshav
ologial funtions' Thir fr-rntionis to bi- highly ompt sum
dit hvior tht adapts an orgn-
ism to a spifi fatur of its nvironmnt. ing onptions of g
Vervt monkys hv distintiv
alls and fsponss trl th siglrt of a n tradd in tlrl
|oard, and thi lopard-rognition
thoughts are bout lopards us geeralizatlons. o,
thir funtion is to ada.ptvrvt bh.rv.
tor to th prsnof leopards in tlrir pistmology of sre
nvirltrmnt.hos u..u.., with a dis.
positiorr to sraml to th top gnralizations' tog
of trs wn thy hd distintivly
lopardy visual pinwr rrror dliv quantitativ i
likly to liv than thos without it.
th loard rspons mans ..loparl'' So t h r s u | to f s p i f i
vn whn it is a fals alarm' for that
disposition irr th vrvt ists s a sult sults of natural pr
of seltionro. t.opn. voidan.
his viw of maning, of th symol-world frring its loation fl
lationship,ho, ,.,''. vIy at. vidntil wight d1
trativ faturs.[tplins why w
Wa,'t tO say of th vrvt tht it is afraid
of lopards rathr than ing afraid physis nouragd a
of lopi.rrdsnd lopard look.liks
though (I ltav rro ]out)vrvts would vn is the bse cs: th<
..,p.,nd th. ."-. Wy to dummy
lopards onstrttdy havioral domains-hmistry
ologists. }.or ltrprd dummis hav
playd no part in th volution rf n d h i r g n r l i z l r t
vrvts and thir sap bhaviors,
vrvt alarm alls ar aout lopards, so izations. orr of th l
not llpardlook-liks. his viw of
symols eplains how it is possii sttion that Kplr's
t<lmisrprsrrt.Misprsrrtatron is fai[-
ur of funtion. 1r)7hn as of NWtonian m(
hikrr's lrawk rsp<lnsis triggd
1.a duk ovr- fiation of rh rl;rt
lrad, and it fls in ti.ight,th hikn
as misrprsntdits nvironmnt.
h hikn intrnl stat is not prforming physis.
its sltdroi of hwk avoid-
n. This last point is r,ry irnportarrt. From th lt 1960
any irr his fild hav thoglrtthat
misrprsnttionis fatal stumling undr wid-rnging fir
lok for nturalizd virvsof tlrought.
Th lim is that ror' or risrprsntation, th physial sinsv
is a ormtiuntion, and nor.
nrtiv lims annot dfind in fatul sin tht for.rglrtov
trlns. A tl'.o.y of -.,ni',g th,. ( | 9 7 0 | , n d t h t r a d i
imports funtionl onpts-fronr
iology saps this prolr. Funtinl
fas ar tts aout th sltiv Laktos ..dgn
histo".of that Orgn. Thv r ntural
fats. morphology of Riar
th British rsistan
honlas Kuhrl ut his
Sinand th Mirror f Physis trining s a historin
rzozs (Kuhn 1962).T
Until th l970s (prhapsvn latr) enti Paui Fyra
hilosohy of sinrvas lomintdby tion of thirrking aor-r
anrplsdrawn tom th physiai sins,
spiallyphysis' Th stndard
amls of tho1,hng inludd history mpsizd drl
tlr oprrrianrvolution, the transition
from Nwtonin lntionsof spa In sussiv sintifi
nd .i''. to rltivist .on..p.ion,, ond
somtims th rvolution in atomi p l d . B u t t h s r i
physis that sw th stalisl-rmnt
quantllm mhanis.Ths wr rrot of t i o n w i t h p h y s i se n d
th only amples of sinin th wild,
ut thy wr stok eampls, and rnarr,, Kuhn) did thy hall
thrs w... .l.a*,, frorn tlr physi-
al sinstoo. Ths irmpls .,'.o..,"g.d a n d p i s n d q u a n i
th id tat or ltur of
sinwas th frmulation of gnral Yt this piture fail
l,-.. smll st o ptronlssgn-
rl prinipls tlrt hratriza sintifi " Whrl-organism ioll
dtlnrain. gnral prinipls that
th rpiriisttradition xmplifid
.Within y Ngl,slassi .l.hStrtltur
of Si (Nagl |961), tlrrwas oiogy nd volutio
g.,ud dal tlf daaout th natur
" of for ampl, th vuln

o Onalth gg vn if th gg is
]asilYsn us th parnt has
ls lut.Similr[v,th<lughtslrav i_
t havilrthat adapts an orgn-
t. Vrvt monkys hav distintiv
rrd.nd thir Ioprd-rgnition
fLrnti<ln is to adpt vrvt hav-
rollmnt.Thrsvvtswith dis-
ls whn thy hd a distintivly
ly to liv than thos without it. So
:n rvhnit is a fals alarm, fo that
lf sltionfor loprd avoidan.
lrld rlationship,has som vry at-
to say of th vrvt that it is afraid
pardsand loprd lok-aliks vn
rspnd th sm Wy to dur-rrr-rry
lgists. For lopard dummis hav
lts and thir sap bhaviors' so
: lopard look-aliks. This viw of
'prsnt. Misprst-ltation
sponsis triggrdy a duk ov-
t s n l i s r p r s n t idt s n v i r t l n t n n t .
ing its sltdrol of hawk avoid-
lany ilr this fild har,thought tlrt
k for naturalizlviws of thought.
n, is lrmtiL'notion' and nor-
Ll trms.A thory of maning that
/ silpsthis prolm. Funtional
i, of tht organ. Thy ar naturirl
;ophy of sinwas dominatd y
:s,spiallyphysis. Th stndard
l p n i nr v o l u t i t l nt' h t
l tii to rlativistollptions' nd
;is that sW th stalishmnt of
nly ampls lf sin in th wild'
othrs Wdrarvn from th physi-
gd tlr ida that a or ftur of
vs:a small st ot ptionlssgn-
: domin.
id y Nagl's lirssi The Strutr
Ldl of datzroutth ntur of
llilosopb rlf uolutior Tbought 2'5
thsgnral prinipls: dat aout whthr thy Wr aurat dsrip.
tions of natur as it rally is, or whthr thy wr tt undrstood as
highlyornpat sunrmarisof tlr phnomna. Notwithstanding ths diffr-
ing onptionsof gnralizations,though, it was ommon grund that si-
n tradd in thm. Th aim of sin Was to find and rfin suh
gnralizations.orovr, this fous nouagd a rtain pitur of th
pistrnologyof sine:sintili warrant divs from th fat that ths
gnralizations,t<rgthr with appopritly spifid oundary onditions,
dlivrquantitativ and oftn vry pris prditions. Somtims ths ar
th rsult of spifi xprimntl manipulations. Somtims they ar th r-
sultsof natral xprimnts,as in th lrtddisovry of Nptun v in-
frring its lozrtion from prturtions in th orbit of Uranus. ithr way,
vidntialwight dpnds on quantitativ prdition. Firrally, this fous on
physisnouragda partiular pitur of th unifiation of sin.Physis
is te bse s: t thory of vrything' Thoris of spifi sintifi
domirins-hmisty and iollgy-ar just spial ass of physil systms'
and thir gnrlizationswill turn Out to spia1ass of physial gnral-
izations.on of th arly triumphs of Nwtonian mhanis was th dmon-
stlation that Kpl's laws of plantary motion wr mor or lss a spial
is fil_ asof Nwtnian nrhanis,and this ampl was takn to an mpli-
fiation of th rlati<lnshipLrtwnspifi, domain-limitd sins nd
physis.
From th lat 1960s th mpiriist tradition in philosophy of sinam
undrwid-rangingfir (sLaktos and usgrav 1970)' But this fous on
thphysial sinsWas on Olnmon lmnt irr th ompting aotrntsof
sinthat fouglrt ovr its orps. Karl Poppr (1959,,|96),Imr6 Lktos
(1970I'and th tradition thy stalishdtradd in ampls from physis:
Lakatos's ..dgnatingrsarh programs'' did not fous on th id:irlist
morphologyof Rilrard Own ( l 848, 1849) in th mid-nintnthntr]ryor
th British rsistn to lssi gntis in th rly twntith nruy.
Thomas Kuhn ut his tth on th oprnian rvolution (1957), arrd his
training as a historian of physis prvads is Structur of Sitc Reuol-
/los(Kuhn 1962)' Th falsifiationisttrdition, Kuhn' and th irasil nd
ntriPaul Fyand(1975) haiingd muh irr th nopositivisttrdi-
tion of thinking out sin.In partiular, thir pitur of sin :rnd its
historymphasizd disontinuity rathr than smooth inrmntal inrass.
n s i t i o n In sussivsintifi rvolutions old thoris r rfutd, suprsdd'nd
rplad.Brrt rhs itis of nopositivism did rlot hallng its proup-
tion with physis and physial hmistry, rror (with th prtial ption of
Kuhn) did thy hallng th proupation with univrsal gnralizations
and prisand quantitativprdition.
Yt tlris pitur fails to fit iology and spially volutionary iology.
Whol_rganisrrr iol<lgydos not sm to trad in small sets of xptionlss
grral prinipls that dlivr pris prditions of iologial phnomna.
ologyand volutionary iology plnatorily powrful: thy plain,
for ampl,th vrrlnilityof Nw Zaland's nativ id lif to introdud
26 Philoso1lh of uolutionr Thought
.lrs
stoats. irds volvd in n nvironmnt fr of any trrstrialpdator In onlusion, philo
nd fr of any prdator at ll that finds its prey y smll. But ology and way pump. It pumps
volutionary biology rarl nabl us to mak pris quantitativ prdi- many diffitrlt polm
tions. Thr ar gnralizationsin iology, and on of thm is th fragility of aris out of th xtra
island osystmsand th vulnraility of island iota to tintion. But gn- ntury. And it umps
ralizations of this kind ar nithr ptionlssnor quantitativ:thy ar ampls, tools, and mod
littl rsrnlanto Boyl's law. Biologists sm to disovr ausal mha- losophy,s own projts
nisms: natu[al sltion' omptitiv lusion' ologial sussion,and th writings of Rihar
gn rplition arrd transription.But th fftsof thsausal mhanisms finst and most sutl
ar ontxt spifi,and thos ontxts vaty ovr sp and tim. So th dis- bl (|996) is susri
ovry of mhanisms dos not translat into ptionlssrgularitisthat autifully writtn, th
ar th onsquns of th opration of thos mhanisms. Thr ar gnr- Gould's viws, see Tht
alizations about volution on islands: larg animals tnd to gt smallr' and pallingly long ut rs
small I-rimalstnd to gt largr. But thr ar many exptionsto this gnr- ial matrial an b sl
alization. B u e $ r( l 9 9 5 ) ' i
orovr, th sirnpl piu of th unifition of sin fits iology synthsis and post.syn
poorl. hmial proprtis lik aidity indxd, or Wr thought to ind' a I7tt, Dri's Dger
singl undrlying physial proprty that was itslf rsponsil for th ph. splndid arpls of v:
nomnologial proprtis of aids. vn in hmistry it is likly that th rla- Se d Dtb: n Int
tionship btwn hmial and physial proprtis is vry oftn ompl Paul Griffiths (1999),
indd. But this omplxity fist am ovious in iology, for ll iology is in muh mor dtai
and molrrl:rriology do Ilot oIrform to th modl of rdution suggstd
v aidity (Kithr 200). lls not mirl fatoris:thir biologial a-
tivitis ar onstrained y physial prinipls. To that tnt physis rllyis |L!oGRAPHY
th thory of vrything. Bur th lationship twn' say' th onpt of a
dominant gn and th undrlying molular mhanismsrsponsilfr th Burt, A.' and R. Trivrs.2
phnotypi siln of on of a pair of htrozygous allls turns out to b Imts.Camridg
ain, A. J., and P' . Sh
vry ompl indd. Finally' th history of iology is not a srisof rvolu-
nmoralis.Hrdit4
tions and rplamnts.Th history of iology oftn smsto manifstinr- 1952.Th fft
mntl improvmnt.All sinshav thir rvisionsand fals trails, but th nmorirlis.Hrdit(
history of ll iology, plrysiology, gntis' or vn volutionary iology ]954. Nturl s
dos rrot aound with major rvrsalsof or dotrins omparal with th arroll, R. L. 1997.Ptt
m r i d g [ I n i v r s
rvolution in plat ttonis in th arth sins.In short, it is hard to oth
raraft,J., and . Dono
tak iology with full sriousnssas a sinand apt a onption of si. oford UnivrsityPr
n drivd from th physial paradigms.Philosophrs hv gon oth ways Dwkins, R. 1976.h S
on this. Al Rsnrg is th lost prominnt of thos who hv aptd 7982. Th tltt
tlr physiists'onptin of sinand as 2lonsqunhav aptdth \996. limbigN
viw that som omponnts of iology ar not quit up to snuff (Rosnrg 2.04.T sr
.!idnfld
nd Niol
7994). John Dupr6 is th most prominnt ontmporary who rjtsth im-
Dnntt,D. . 1995.Dri
ag of sin in th physiist's mirror on th grounds that it fails tl fit iol- Llfe. Nw Yk: Sim
ogy (Dupr6 2002). Most of thos who work in philosophy of iology ar 200. Frelomt
losr to Dupr6 than to Rosnrg on this issu: thy think that volution- Dpw, D., nd B. H. W
ry iology is a pardigrrr of a sussful sine, and hn th physis- tbe Gnlogl tlf Nt
-.
D o z h a n s k ' 1 9 3 7 ,G
asd onption of sin is too narrow. Thr is no onsnsus' howvr,
UnivrsityPrss.
on tly what is Wrong with th physis modl, or on how it nds to b Dup,J.2002. Hums
xtnded. ldrdg,N. 1995.Rizl

lnt fr of any trrstrial prdator
its pry y smll. But ology and
mak pris quantitativ prdi-
, and on of thm is th fragility of
island iota to tintion. But gn.
'ionlssnor quantitativ:thy ar
ts smto disovr ausl mha-
lusion' ologial sussion, nd
fftsof ths ausal mhnisms
ry ovr spa and tim. So th dis-
into xptionlss rgularitis that
hosmhanisms.Thr ar gnr-
; animals tnd to gt smallr, and
a r m a n y x p t i o n st o t h i s g n r -
nifiation of sin fits iology
dxd, or wr thought to indx, a
was itslf rsponsil for th ph-
r hmistry it is likly that th rla-
proprtis is vry oftn omplx
lvious in iology, for ll iology
l th modl of rdution suggstd
iral ftoris:thir iologial a-
lls.To that tnt physis rally is
;hip twn' say' th onpt of a
lar mhanisms rsponsil for th
)trozygousallls turns out to
of biology is not a sris of revolu-
'logy oftn sms to manifst inr-
:ir rvisions and fals trails' ut th
tis, or vn volutionary iology
:ordotrinsomparal with th
ins.In shrt, it is hard to oth
n and apt a onption of si-
Philosophrshav gon oth ways
rinnt of thos who hav aeptd
ls a onsqunhav aptd th
not quit up to snuff (Rosnrg
ontmporary who rjts th im-
th gounds that it fails to fit iol-
vork in philosophy of iology ar
is issu:thy think that evolution-
;l sin,and hn th physis-
v. Thr is no onsnsus. howvr.
:s modl, or on how it nds to
Philsophof uolutilnr hought 27
In onlusion, philosophy of volutionary iology ats as a typ of two-
Way pump. It pumps philosophy into volutionary thory, for thr ar
many diffiult prolems of intrprtation' larifiation, and integration that
aris out of th xtraordinary dynamism of th lif sins ovr th past
ntury.And it pumps volutionary iology into philosophy: a stram of -
ampls,tools, and modls that an prhaps adaptd to driv forwrd phi-
losophy's own projts. If you want to rad mor, I suggst you turn first to
th witings of Rihard Dawkins: The tedd Phnotpe (1982) is his
finstand most sutl df of gn sltin;limbig ount Improb-
ble (1996) is a sustaind and onvining dfns of grdualism. Both are
autifully writtn' though Th tded Photpe is quit thnial. For
Gould's viws, see Tbe Structure of uolutionr Tbeor Q002),It is p-
pallingly long ut rasonably radal, and a good dal of th purly histor-
ial matrial an skipped. Drtuiism uoluig' y David Dpw and
Bru \Wr(1995), is a good vrviw of th historial dvlopmnt of
synthsisand post-synthsisvolutionary iology. Two ooks y Dan Dn-
tt' Drlui's Dngros Ide (1995) and Freedom uolus (2003), ar
splndidampls of valu of volutionary insights within philosophy itslf.
S d Deth: ltroductio to tb Philosoph of Biolog' y m and
Paul Griffiths \1999), ovrs many of th issus of this ssay and othr top-
is in muh mor dtail.
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hilosoph 86 281_297.
Philosophoi Si 56: 288_302.
ms i t Llgi of Sitifi Disour.
 two wll-kno,n nd <
Spnr, Ptr Krpol
soial thinkrs who in
trs in suh rnt o<
e: Dsigig Ntu
volution and Soity familiar popularizrs
Rathr, w xplor l
soity v sking o
NInfrdD. Lbihlr nd
Jn ienshin Dfinitions

B uoltio W ma
ral volutionary id
hang of population
volution and soity ar onntd in many diffrnt ways. First and for. and lad to divrg
most' humans' as a podut of volution, liv in divs soitis.Ths ar adaptd t thir nvi
an outom of volution, alit in mor ompl and oftn mor indirt ally, with variations <
ways than th voltion of dirt morphologil nd physiologial adapta. and spis nd popu
tions. Sond' th initial dvlopmnt of volutionary thory ourrd in stant thms: l[ o
spifi soial miliu onduiv to this prriular intlltual endavor, and prnaturalism as a (
susqunt dvlopmnt of volutionary thory has also rfltd its volving nvironmnt tlrt in
historial sttings. volutionary thory, lik any othr sin, has thus n vord ar etter dp
th produt of a partiular human soity.Finally, and as proaly th most tions that mattr for (
visil onntion of volution and soity, spially in th Unitd Stats,w larning btwen gn
find surprisingly prsistnt dats aout whthr volution or som form of mission.
.W
rationism aounts for th origin of spis (s ugni . Sott disusss also shar th r
in th main ssay ..Amrian Antivolutionism: Rtospt and Prospet'' in ory of volution is t
this volum). som tims in th o
At som tims, in som plas, and in som wys' soity has nthusiilsti- othr of rhsthoi
ally mrad volution; at othr tims nd plas and in othr was, soi- ity, struggl ;rrrdom;
ty has not. In th pross euoltio has mant diffrnt things, as has all part of volution
soiet..W thrfor start with dfinitions and thn look at soity's impat onsists of irrtronn
on volution in two ways: th way in whih soity in th sns of soial' ul- this ssay.
tural, and intlltual ontt has shapd volutionary thory, and also th B soit w dsig
soial rption of volution. Thn w turn th dirtion of influn around th rlvant lmnt
and addrss how volutionary thory today and in th past has ontriutd ft many othrs. In t
to planations of soity.In onlusion w distlssurrnt rlations btwn tingnt, nd populist
volution and soity in a forwrd-looking way. b taught in th sh
.W
do not fous on muh-disussd rlzrtionshipssuh s standard irrtrpr- mands rationist
tations of soial Darwinism or rtra th lristory of ways in whih soial ial olrorts ply dif
ommntators hav usd (and misusd) volr.rtion.Thr is suh rih litra- groups as w go lon1
tur in ths gnral aras that any gnral Googl or Amazon sarh will of (human) soitis
yild dozns of offrings, of whih arl Dglr's I Serh of Hm Ntur: mans any group of
The Decli d Ruiul of Drtuiism i mrin Soil Thoght (1992| or struturd way and '
Rihard Hofstadtr's Soil Drinism i Amri hought (1959) ar just 19751Bod and Rih

0

)1ty
nd
rny diffrnt ways. First and for-
liv in divrs soitis.Ths r
ompl and oftn mor indirt
llogial and physiologial adapta-
volutionary thory ourrd in a
rtiular intlltual ndavor, and
rory has also rfltd its volving
l any othr sin' has thus bn
Finally' nd as proaly th most
, spiallyin th Unitd Stats,w
hthrvolution or som form of
is (as ugni . Sott disusss
nism: Rtrospt and Prospt'' in
om Ways' soity has nthusiasti.
n d p l a sa n d i n o t h w a y s ' s o i -
as mant diffrnt things, as has
s and thn look at soity's impat
h soityin th snsof soial, ul-
volutionarythory, and also th
n th dirtion of influn around
ay and in th past has ontriutd
. disussurrnt rlations twn
3 way.
donships suh as standard intrpr-
r history of ways in whih soial
olution. Thr is suh a rih litra-
ral Googl o Amazon sarh will
g|er,sI Serh of Hm Ntr:
mrin Soil Thought (1'992) ot
Americ Thought (1959) ar just
uolutio d Soit 1
two wll-known and diffrnt typs of ampls. Radrs an look to Hrert
Spnr,Ptr Kropotkin, Andrw arngi, and othr popular xampls of
soial thinkrs who invokd volution, or they an tak up th volution hap-
trsin suh rnt ooks as Brnard Lightman's Vitorin Poplrizrs of Si-
e:Desigi11gl\Iaturflr |'treAudiecs (2007) for an introdution to lss
familiar populaizrs. Again, it is nt on this wll-trod ground that w roam.
Rathr, w plor lss familir aspts of th intrstions of volution and
soityy asking aout th rlationships in a varity of ways.
Dfinitions
B euolutio w man th volutionry thory of iologists rathr than gn-
rl volutionary ids, tht is, naturalisti, matrialisti xplanations of th
hange of populations ovr tim suh that inhritd variations ar prsrvd
and lad to divrgn of forms and th origin of nw spis that ar
adaptdto thir nvironmnts.This pross an happn mor or lss gradu-
ally' with varitions of various sorts' inhritan working in ompl ways'
and spisand populations dfind diffrntly.Nonthlss, thr a on-
stant thms: all aounts of volution gin with naturalism rathr than su-
prnaturalismas a or assumption. Al1 assum that organisms xist in an
nvironmnt that in fft favors som variations ovr othrs sin th fa-
vord ar etter d|lted' 1rnd all vrsions assum that it is inhritd varia-
tions that mattr for volution, though inhritan may om ulturally with
larning twn gnrations' as wll as iologially through gnti trans-
mission.
! also shar th viw of rnst ayr and othrs that th Darwinian th-
ory of volution is omposd of a varity of sparat thoris, and that t
som tims in the ours of th history of volutionry iology on or th
othrof thsthoris domintd th dbats(Mayr 1982I. Variatin, hrd-
ity, struggl and omptition, and mrgn and prsrvation of th nw r
all part of volutionary thory. Indd, vlutionary thory is ompl and
onsists of intronnetd omponnts that we xplr in diffrnt ways in
this ssay.
B societ w dsignata varity of kinds of soial groups. In som ass
th rlvant lemnt of soity nters on ladrs who mak disions that af-
ftmany othrs. In othr ass th rlvant soial group is mor loal, on-
tingnt'and populist, as whn ivi ladrs may ag that volution should
taughtin th shools in any ivilizd soity' ut a voal popular group d-
mandsa rationistantivolution aount of th oigin of man. Diffrnt so-
ial ohorts play diffrnt rols, oviously, nd w will look at diffrnt
groupsas w go along. Similarly, whn w disuss volutionary planations
of (human) soitis, w follow an qually atholi approah; soiet ere
mansany group of animals or humans that intrat with h othr in a
struturd wy and whos survival dpnds on ths intrations (.lfilson
1975;Boyd and Rihrson 1985; Kupr 1994).
2 uolutkl d Societ

assumptions of unitl
volutionary Thry Shapd y Soity soity shard th as
hang ovr tim tha
volutionary thory xists in rtain soitis nd is dvlopd y mmrs 2002; Rihards 200,
of ths soitis.That muh is ovious. Darwin is th mst visil ntral th intlltual nvi
playr, and historins hav shown th signifian of th fts that soially partiular vrsion of
Darwin ws uppr middl lass and a Vitorian nglishman. For ampl, progrssiv impliat
th British gntlman would prsumaly hav found it asir than th poor an pliitly and
son of a long-poor working-lass family to om to Darwin's viw of inhri. impat, just s soi
.What
t a n a s a p o g s s i v n d p o s i t i v f o r i n p r s r v i n gv r i t i o n o t o s was sn as
hrdity as naling rathr than onstraining. Th way Darwin saw th ntury also rflts
world from .S Begle was surly shapd y his ontt and valus, as his- ory (Bowlr 1983)'s
torins hav shown (Dsmond and oor 1991; Browne 1'995,2003;odg randomly arising vz
and Radik 2003). His world Was th rihly ut gntly ntangledank of an and a fous on th i
nglish gardn rathr than n arid dsert, a divrs and pungnt rain forest, ganism in its nviror
or an Alpin tundra, h of whih might hav ld him to diffrnt qustions prtations tht app(
and mphass. rformrs who wr
Furthrmor, as Darwin sholars hav dmonstratd larly, th British ti dtrminist or h
.silliam
ontt of natural thology shapd volutionary thory. Paly saw ary thory, and th
ah organism and ah typ as adptd to its pla in th world. Of ours' If th hritabl vari
it was Paly's Anglian God th Crator who had don th adapting in that prsumably influt
.W
as, but th fit of lif form to nvironmnt arrid ovr natly to Darwin's annot ovr
viw of lif. Tru, a Thomas Rort Malthus might point to th human o. ing soity, though
nomi dilmmas intrinsi in th tndny of vry individual to inasinto Hr th point is th
a population that rapidly outgrows availal rsours.But althus did not volutionry thory,
all into qustion that humans wr adaptd to thir nvironmnt. Indd, to ask and whih as
th tndny to produ too many individuals just givs natur th opportu- rtainty whr som
nity to hoose som ovr othrs. Or, as Malthus suggstd, prhaps man as wll as intrnal t
ould do th hoosing with wis population ontrol. At any rat, soial viws study voiution: nt
shapd th volutionary thory that mrgd,and th rsultswr optimisti. trol and manipulatir
For althus, population ontrol would mak lif ttr for ah prson and whol orgnisms, pi
..halthy,vigorous, and happy''
for soity as a whol. For Darwin, only th parts lik bhaviors
.$isman
would surviv and rprodu. A happy pitur inded (Rus 1979,2003; August
Hodg and Radik 2003). 1'892 Kimplsml
Anothr part of Darwin's Vitorian miliu that was trmly important nvironmntal influ
for his vntual formulation of th thory of natural sltionas an plana- trial of volution,
tion of volutionary hang was his los ontat ovr many yars with many ptition among his
animl and plant brdrs(Brown 2003). Indd,th los omprison - long visibl hrom
twn artifiial sltion (with morphologial and havioral hangs in pop- matrial units, was
ulations that ould indud y ontinud rding)and natural sltion qustions aout hov
(hangsthat would a rsult of th ..strugglfor istn''in natur) is a th fin dtails of th<
..long
major part of Darwin's argumnt'' Was part of a sinti
Th Grmn intlltual movmnt Nturphilosophi, with its romanti mirosopi osrv
onption of natur, rought othr prsptivs that shapd volutionary ah organism, and
.!ismann's
thory as it playd out in Grmany. Johann Wolfgang von Goth's organi- volt
ism that saw an unfolding of form' Fridrih Shlling and Gorg Hgl's impat on th inh

olty
tis nd is dvlopd y mmrs
Darwin is th most visil ntral
nifian of th fats that soially
]torian nglishman. F.r xampl,
rav found it asir thzrnth poor
) om to Darwin's viw of inhri-
in prsrving variati;n or to s
ining. Th way Drwin saw th
l by his nttand vlus, as his-
1 9 9 1 ;B r o w n e 1 ' 9 9 5 , 2 0 0 ;H o d g
ly ut gntly ntirngldank of an
divrsand pungnt rain forst,
t v l d h i m r o d i f f r n tq u s t i o n s
dmnstratdlarly, th British
ionay thory. !illian-rPly saw
l its pla in th world. of ours,
,ho had don the adapting in that
rt arrid ovr natly to Darwin's
Lusmight point to th human o-
' f v r yi n d i v i d t r a l i r l r a s i n
tl rsours.But althus did not
td to thir nvionnrnt.Indd,
rals just givs natur th opportu-
althLrs suggstd,prhaps man
l ontrol. At any t, soi:rlviws
: d ' a n d t h r s u l t sw o p t i m i s t i .
rk lif ttr for ah prson and
r ..halthy,vigorous, and happy''
itur indd (Rus 1979' 20:
i u t h i r tW a s x t r m l y i m p o r t a n t
lf natttralsltionas an xplana-
)ntat ovr many yars with many
Indd,th los omparison -
l a n d h l r a v i o r l h n g si n p o p -
:d rding)and natural sltion
lgglfor xistn''in natur) is a
'urphilsophie'with irs omnti
)tivsthat shapd volutionary
n.Wolfgngvon Goth's organi-
rih Shlling and Gorg Hgl's
uolutiln d Soit 3
assumptions of unity of natur, and Karl r's sarh for an unfolding of
soityshard th zrssumptionsof mrgn or of form from nonform and of
hngovr time that lads to irlprovmnt:rnd progress (Rus 1996; ok
2002; Rihards 2002)' Ths soial and philosophial thrads hlpd shap
th intlltual nvironmnt into whih volution arrivd. rnst Hakl's
prtiular vrsion of volutionary thory in Grmany shard th positiv and
progrssivimplitions of volutionary hang and dvlopd th idas in
an xpliitly and r\triallymtrialisti wy tht hd tmndous soial
impat,just as soity had shapd th sintifiidas.
..elips of Darwinism', at th nd of the nintnth
What was sn as an
ntury also rflts impats from outsid sin itslf on volutionary th-
ory (Bowlr 198), shifting away from rlian orr what Drwin hd sn as
randomly arising variations. ndlism intodd gns, hritbl units'
and a fous on th intrnal wokings of th oganism mor than on th or-
ganism in its nvionmnt. This brought a shift from th mor organi intr-
rtations that appaled to ar and othr lat nintnth.ntury soial
rformrswho wr alling for natural hang. Instad,w s a ris of gn-
ti dtrministor lrrditarinthinking, whih in turn inflund volution-
ar thory' rrd th dtrminismof whih appaldto diffrnt soial groups.
If th hrital variations wr gntially dtrmind, thn volution ould
prsumaly inflund y modifying ths gns.
.W
annot ovrmphasize tat this is not simply a mattr of sin afft-
ing soity,though that has n rh primary fous of historial disussion.
Hr th point is that iors within soity wr affting sin' shaping
volutionarythory, and influrringth hois mad aout whih qustions
to ask and whih assumptions to mak in a ompl thortial arna of un.
rtaintywhr sm assumptions must nonthlss mad. Soial fators,
as wll as intrnal sintifi dvlopmnts, shp whih mthods to us to
s u d y v o l u i o n :n a t u r a l i s t i f i l d w r k a n d d s r i p t i o r ol r p i m n r l o n -
trol and manipulation of varials, and study of populations or individuals,
whol organisms, parts of organisms, invisil parts lik gns' or mrgnt
parts lik haviors and othr traits.
.W.ismarrn
August providd n inhritan-asdvolutionary thory. is
|892 implsm ortaindsqustrdgmplasm that was prottd from
nvironmntalinflun. This was th matrial of inhritan nd th raw ma-
trial of volution, providing sustan for volutionary lrang though om.
ptitin among his hypothtial ids, dtrminants, and idants' all arrangd
along visil hromosoms. Th rlvant population, that of ths inhritd
matril units, ws rrow insid th individual organism, and this raisd nw
qustionsabout how far th ation of natural sltion an tak us and aout
.sismann,
th fin dtails of the mhanisms of volutionary hang. For who
Was part of a sintifi tradition that fousd on ll iology' dvlopmnt, and
mirosopi osrvation' th important natural sletion took pla insid
ahorganism, and tlr nvironmnt was th intrnal nvironmnt.
'Wismantr's
volutionary thory disllowd dirt xtrnal nvironmntal
impat on th inhritd hromosoms. His drrial that Lamarkian us and
34 uolution d Socit

disus of partiular parts ould hav any ffton volution is takn in rtro- te 1970s to qustin
spt as putting that ida to rst' though it did nt atually do so. Instad, ( l d r d g a n d G o u l d l .
no-Lamarkism ontinud to find a pla in soial and politial ontxts mulation of morpholo
that mphasizd th importan of hang through ffort and will, suh as ..
with mor suddn
tht of Rihard Smon arly in th twntith ntury nd with a lin of lss othrs, as passing thro
wll-known suPportrs through susquntgnrations (Smon 1908). By trnl fators might
th sond half of th twntith ntury' politi4l figurs suh as T. D. port ompting intrpr
Lysnko gaind onsidral authority in th Stalinist Sovit Union' and not thir viws wr intr1
solly for distortd politial rasons. Lysnko's no-Lamarkism appaldto th validity of volutio
th hop that if only sintists ould ultivat sds in th right nvironmnt, A ntral point hr
thy ould produ nough grain to sav th popl from starvation. Thr t h o r y t h a t r s u l t dv
was trmndous lif in th powrs of sin, and vn though no- appard. Old-shool
Lamarkian idas hav latr n idntifid with Sovit-styl matrialism, at f o h a n g .N o t o i n i
last initially many of thsproposals mphasizdth rol of ultu and th young and fanid th
soial nvironmnt in shaping th fat of human soitis. hang. This dos nor
Som would insist that this was larly ad sin, and for thos fully fa. than thos who favor
miliar with ontmporary sintifi paradigms, it was. But for thos for hav not had trmnd
whom volution was not a ntrl part of thir duation and worldviw, Thy hav.Our point l
suh an volutionary thory that would pla th ausSof variations dirtly tists asord hav sh
in us and disus in rspons to th nvironmnt mad sns.It was not a Today w s anotl
long strth from Darwinian volution to sing tmpratur hangs as a volutionary iolog i
sorrr of nw variations through mutation and sltion and thrfor as th tionary iology. Popul
sour and aus of th origin of spis.Lysnko's vrsion of no-Lamarkism within population. Bu
promisd a spdirvolution, and sin volution Was sn as progrssiv,it isms do, and th way
thrfor offrd a fastr rout to progrss and improvmnt. Soial nds variations has importa
shapd sin' whih thn playd out in soity (Tods 7989\. hangs. wo notions'
.s
s anothr ampl in th so-alld modrn volutionary synthsis of hav widr soital ir
th lat 1940s and aftr. Partiipants at ky mtings, suh as te |947 intrations (Maynard
Printon mting' saw this as a tim of optimism for volutionary iologists t h a t n o t l l v r i a t i o n s.
(Mayr and Povin 1980). Th prwar tnsions in urop and th Unitd vlopmntal and phys
Stats may wll hav nouragd th sort of synthsis that Julian Huxly, as vritions tran
a politial ativist with th Unitd Ntions, sought in is uolutio: The r t i so f t h d v l o p i n
odern Stbsis (1942). Huly was not a major intlltual shapr of the pl interations amor
privd synthsisut ntral voi in oniving of it in that way (Smo- togthr ontriut to
ovitis 7996). Howvr, vn whil volutionry iologists suh as rnst dvo (short for volut
Mayr and Gorg Gaylord Simpson agrly mrad th ida of synthsis s i z s a n p i g n r i a n d
with volution at th or, alrady othr iologists wr pushing in ountr- This prsptiv has
dirtions.ll iologistsand gntiistsrd mor aout th intrnal work. ida of Iimitson what
ings of lls and thir ontnts than aout th biggr piturs of volution. ws first mad popul.
Rapidly inrasdfunding for mdial rsarhand mdial dmands of wars om'' and th nvir
hlpd driv wdgs twn spialtiswithin th iologial ommunitis. on intrtions , v nb
volutionary iologists oftn found thmslvsas omprativ spialistsin f i r s t m a n i f s t t i o n si n
natural history musums' whil mor rdutionist rsarh programs gaind rlvant within dvlo
priority in many univrsitisand spiallyin mdial shools. ally, th ompltion
Dats aout th ffiay of adaptation rval similar soial shaping of nation of th gnti p
volutionary thory. StphnJay Gould, Nils ldrdg, and othrs gan in a n d p i g n t i n a t u r

l.ton volution is takn in rtro-
t did not atually do so. Instad'
in soial and politil ontts
: through ffort and will, suh as
lth ntury and with a lin of lss
lt gnrations(Smon 1908). By
, politil figurs suh as T. D.
h StalinistSovit Union, and not
ko's no-Lamrkism appald t
r s d si n t h r i g h t n v i o n m n t ,
h popl from starvation. Thr
sin, and vn though no-
l with Sovit-stylmatrialism, at
rasizdth rol of ultur and th
uman soitis.
'ad sin,and for thos fully fa-
ligms, it was. But for thos for
f thir duation and worldviw,
th aussof variations dirtly
onmnt mad sns.It was not a
singtmpraturhangs as a
and sltin and thrfor as th
;nko'svrsion of no-Lamarkism
o l u t i o nW a ss na s p r o g r s s i v ,i t
s and improvmnt. Soial nds
rcity (Tods 1989).
modrn volutionary synthsis of
ky mtings, suh as t'e 1947
timism for volutionary iologists
nsions in urop and th Unitd
lf synthsisthat Julian Huxly, as
ns, sought in ts u<iltio: Tb
a major intlltualshapr of th
:onivingof it in that way (Smo.
rtionary biologists suh s rnst
ly mradth ida of synthsis
i o l o g i s t sw r p u s h i n g i n o u n t r -
rd mor aout th intrnal work.
th iggr piturs of volution.
rrh and mdial dmands of wars
' i t h i n t h i o I o g i l o m m u n i t i s .
lvsas omparativ spialists in
tionistrsarhpograms gaind
in mdialshools.
n rval similar soial shaping of
ils ldrdg,and othrs gan in
uoltion d Societ 33.'
th 1970s to qustion how far natural sltion an rally fft hang
(ldrdgand Gould 1,972).Isvolution rally allgradualism and slow au.
..puntuatd''
mulation of morphologial diffrns? Or ould hang
..plosions''?Prhaps som hangs ould nal many
with mor suddn
othrs, as passing through a ottlnk to th opning yond an do. Or
trnal fators might lad to puntuation. volutionary thory ould sup-
port ompting intrprtations gf mhanisms' ths rsarhrs flt, though
thir viws wr intrprtd y antivolutionary rationists as hallnging
th validity of volution.
A ntral point hr is that this ritiqu and th rvisions in volutionary
thory that rsultd wr, in part, shapd y th soity in whih thy
appard.old.shol adaptationism and gradualism providd a slow pross
for hang. Not oinidntally, thos who wr alling for puntuation wr
young and fanid thmslvs soial rformrs who favord rapid soial
hang. This dos not man that thy wr wrong sintifially any mor
than thos who favord gradulism. Nor dos it man that thir hallngs
hav not had trmndous positiv impt on volutionary thory gnrally.
Thy hav. our point hr is that th soity and th soial valus th sin-
tists asord hav shapd th volutionary thory signifiantly.
Today W s anothr transformation of th adaptationist paradigm in
volutionary iology in th form of a mrgr of dvlopmntal and volu.
tionary iology. Population gnti modls hav mphasized gnti hangs
within population. But gns do not intrat with th nvironmnt, organ-
isms do' and th Way gnti variations rlat to orrsponding phnotypi
variations has important impliations for our undrstanding of volutionary
hangs.Two notions, in partiular, that ar part of this nw paradigm also
hav widr soital impliations, namly, th idas of onstraints and of
intrations(aynard Smith t al. 1985). Th ida of onstraint mphasizs
that not all variations ar qually likly or vn possil. It stats that th d-
vlopmntal and physial oundaris of an rganism limit th ways gnti
variations an trnslatd into phnotypi varitions aus of th prop-
rtisof th dvloping systm. Ths ar, of ours, a onsqun of multi-
pl intrations among gnti' ellular' and nvironmntal fators that
togthrontriut to organismi dvlopmnt. Th nw synthsis of vo-
dvo (short for volutionar and dvlopmntal ilogy) thus also mpha-
sizsan pignti and intrativ viw of iology.
This prsptiv has many sintifi, ultural, and soital rsonans; th
ida of limits on what is possil within th oundaris of a ntural systm
..population
was first mad popular in th ontxt of th worry about a
om'' and th nvironmntal movmnt. Similarly, th growing mphasis
n intrations' vn twn distant parts of a larg osystm' has found its
fistmanifstations in this ontxt. But today ths onpts ar also sn as
rlvantwithin dvlopmntal and volutionary biology, whr, quit ironi-
ally' th ompltion of th many gnom proits-in mny ways th lmi-
nation of th gnti paradigm-has ld to insights into th highly intrativ
and pignti natur of oth dvlopmnt and volution (Hall 1,998)'
6 uolutio nd Soit
sltion as an pl
That volutionary thory has not n insulatd from soial influns is
207). Anothr influ
no surpris to historians of sin' ut it is oftn surprising to sintists who
natural history and
s sin as rlativly insulatd and progrssing aording to its own intr-
is of volutionary h
nal logi and rsponsS to opportunitis. It is also surprising t som soial
whil, in Vinna m
historians' who rarly study th history of sin and who tnd to tak si-
dvlopmntal mha
n as givn and to s it as shaping soity on oasion rathr than to s
th work of th iol
th intrations in both dirtions. Darwinism. as Dsmond and oor
primental basis fo
(1991) and Brown (1,995 , 2003 ) hav shown learly, is iust as muh a story
ontt in favor of a .
of Darwin in soity as of Darwinism in soity.
on volution (Przirar
All ths ampls
ity nvr Was jrrsta
Soity'sRption of volution
rlationship that in tu
ary thory in intrsti
Historians hav long dsrid th rption of Darwinism, in partiular, to
.shy
show th variations in ration. is volution takn as a hallng to s-
talishd valus in som ountris or loal soitis and not in othrs or to
volutionary Th
som rligions and not othrs? \/hy hav som groups radily ndorsd th
naturalism at th or of volutionary thory, as Germany did in th lat
Thr ar no a priori l
nintnth ntury, whil thr is hostility at som tims and in som groups
planation as long as
to th apparnt randomnss and lak of purpos that thy s in volution?
multitud of diffrnt
To what tnt is th rspons shapd y loal ontingnis of dominant in-
and ologial intra
fluntial individuals, prhaps' and to what tnt dos th ration flow from
(Rus 1996).Ovious
th logi of th values and assumptions within th ultu? Th answrs to
mnts of slim molds l
ths qustions ar ompl and involv dtaild aounts of th soial' ul-
orhstra who ar pl
tural, onomi, and politial history of ths soitisand ountris. For
th rsult of th intrp
mor dtaild aount, th radr should onsult th sustantial ody of lit-
.$7assr- th framwork of vo
ratur in this ara (fo xampl, u|| 1971Kohn and Kottlr 1.985;
mnt, kp in mind t}
sug and Ros 1984).
ioral, an onsidr
Hr w want to point to just on aspt' namly, how diffrnt soitis'
Natural sltion f;
rption of volutionary idas ontriutd to th frthr dvlopmnt f vo-
manag to inras t
lutionary thory. \Within Darwin's innr irl Thomas Llnry Huxly most
is dsrid y th t
visily mphasizd th importan of volutionary thory to promot a liral
sris th onsqun
and matrialisti agnda. To him, volution implid that thr is no intrinsi
that an rprodu 1
valu in heritag and pdigr, and tht all that should ount ar th ailitis
1,998).Th rpliator
of popl. As a onsqun, h promotd sin as a profssion rathr than
a voation and fought a onstant attl gainst rligion as th stalwart of r- its atually ontriu
variabl faturs ar r
ivd valus (Dsmond 1997)' Similarly, rnst Hakl fousd on th mat-
a (loal) optimum as
rialisti impliations of volutionary thory, whih h pandd into a whol
thrmor' th rpliat
systm of monisti philosophy. Nithr Huly nor Hakl thought highly of
ag to rprodu th
natural sltion as a mhanism of volutionary hange. Rathr, ah inor-
rprodution happns
poratd th prinipl of volution into his sintifi disiplin (mostly mor.
proprtis do not disir
phology) and his highly sussful tahing. In tahing thy oth intratd
potntial and atual l
with and shapd urrnt trnds within thir rsptiv soitis' whil thir
uls, gns' lls, l
sintifi ontriutions inflund svral gnrations of iologists.
groups' and idas, s
A similar pattrn an sn in Russia, whr ids and osrvations
1984; Sor and.Wils
aout symiosis and oopration also hallngd th dominan of natural
 uolutio d Socit 7

sltion as an xplanation of volutionary hang (Tods 1989; Akrt

nsulatd from soial influns is
2007|. Anothr influen in Russia was th prsn of a strong tradition of
' oftn surprising to sintists who
.ssingaording to its own int- natural history and ology, whih laid th groundwork for susqunt stud-
is of volutionary hangs within loal populations (Adams 1994). an-
t is also surprising to som soial
whil, in Vinna mphasis on primntal work within physiology and
sinand who tnd to tak si-
dvlopmntal mhanis paird with fous on an organism's lif history in
:ty on oasion rathr than to s
th work of th iologistg who workd in th Vivarium. This providd an
zinism, as Dsmond and lV1oor
xprimntal asis for proposals that qustiond th thn-mrging gnti
wn larly,is just as muh a story
ontt in favor of a dvlopmntal (inluding no-Lamarkian) prsptiv
ity.
on volution (Prziram 1904).
All ths ampls show that th rption of volutionary thory in a so-
ity nvr was just a on-way strt. In all ass this was a highly intrativ
rlationship that in turn ontriutd to th furthr dvlopmnt of volution-
ary thory in intrsting ways.
ln of Darwinism, in partiular' to
rlution takn as a hallng to s-
I soitisand not in othrs or to
volutionry Thory plaining Soity
iom groups radily ndorsd th
]ory' as Grmany did in th lat
Thr ar no a priori limits on what an th subjt of an volutionary x-
lt som tims and in som groups
planation as long as rtain asi onditions apply. From monad to man' a
rrpos that thy s in volution?
multitud of diffrnt lif forms and thir haviors, as wll as thir soial
l a l o n t i n g n i osf d o m i n a n t i n -
and ologial intrations, hav all bn shapd by volutionary fors
]tnt dos th ration flow from
(Rus1996). Oviously thr ar diffrnsbtwn th oordinatd mov-
ithin th ultur? Th answrs to
mnts of slim molds that ar forming a fruiting ody and th mmrs of an
:taildaountsof th soial, ul-
orhstra who ar playing a Bthovn symphony, ut oth ar ultimatly
tssoitisand ountris. For a
th rsult of th intrplay of omplx haviors that ist in aordan with
onsult th sustantial ody of lit-
th framwork of volr.rtionarythory. In ordr to undrstand this last stat-
; Kohn and Kottlr 1985; !assr-
mnt' kp in mind th ways in whih any fatur, morphologial or hav-
ioral, an onsidrdth produt of volution.
t' namly, how diffrnt soieties'
Natural sltion favors variants with highr fitnss, namly, thos that
to th furthr dvlopmnt of vo-
manag to inras thir rprsntation in th nxt gnration. This pross
r] Thomas Hnry uly most
is dsrid y th univrsal rpliator quation that mathmatially d-
ionary thory to promot a liral
sris th onsquns of natural sltion and that applis to all objts
l i m p l i dt h a t t h r i s n o i n t i n s i
that an rprodu thmslvs (Dawkins 1'976; Hofar and Sigmund
that should ount ar th ailitis
1998).Th rpliator quation dos not speify what proprtis of ths ob-
;ins a profssion rathr than
jts atually ontriut to fitnss diffrns; it only prdits that whatvr
rinstrligion as th stalwart of r-
varial faturs ar rsponsil for ths ffts, ths will vntually rah
rnst Hakl fousd on th mat-
a (loal) optimum as long as nvironmntal onditions do not hang. Fur-
, whih h pandd into a whol
thrmor, th rpliator quation dos not spify how thos variants man-
cy nor akl thought highly of
ag to rprodu thmslvs. All that is rquird for natural sltion is that
onary hang. Rathr' ah inor-
rprodution happns with a suffiint dgr of auray that th favorabl
sintifi disiplin (mostly mor-
proprtisdo not disintgrat too fast. any diffrnt struturs an thus
;.In tahing thy oth intratd
potntial and atal rpliators, or units of sltion. Ths inlude mol-
ir rsptivsoitis, whil thir
uls,gns, lls, lons (in th otanial sns),grops' inluding soial
)nrationsof iologists.
groups' and idas, so-alld mms (Dawkins 1982; Brandon and Burian
a, whr ids and osvtions
1984;Sr and.!7ilson 1998).
llngdth dominan of natural
38 uolution d Socit

Th rpliator quation spifis th road limits of what kind of propr- oordinatd. ow t

tis an volv-thos that ithr inras th fitnss of thir arrirs or thos fasinating prol
that ar nutral with rgard to fitnss (thos' howvr, ar th sujt of a Ths rif thn
diffrnt dynamis; s Kimura 1983). If w now ask in what way a spifi ary thory also hl
nw phnotyp suh as a morphologial fatur or a form of havior an nurtur in a diffr
th produt of volution, w hav to addrss an dditional qustion. Bfor tintion is on onl1
suh novel phnotyps an ontriut to fitnss,thy must first mrg within tion btwn natu
thir arrirs; that is, thy must thmslvs produts of volution. In othr A multitud of fat
words, th molular' llular, and dvlopmntal ontt of organisms is th opmnt and formal
first part of th planation of any nw fatur. Only thos phnotypi vari- gram rquirs all tt
ants that ar possil at all an om th raw matrial for natural sltion, As w hav lar
us phnotyps dtrmin th fitnss of thir arrirs. Th fat that d- rlativs, w ar
vlopmnt plays a ruial ro1 in this gnration of variation has rntly this fat is that in h
n th sujt of inrasdattntion (Raff 1'996;Hall 1998). titud of nvironm.
For our disussion of volutionary xplanations of soitis' a diffrnt in our volutinary
aspt of th volutionary proSs is also of intrst. As w hav sn, all that tion of languag'w
is rquird for natural sltion to at is th availaility of auratly trans- ns (Diamnd 19
mittd variation in fitnss. This rlationship is ommonly masurd y hri- also nal larnin
taility of a haratr. Hritaility is atually dfind as th orrlation twn sion' all vidn t
parnts and offspring' and vn though in many ass th rason for this diffrnt fators (
orrlation will gnti' it is not limitd to th ffts of gns (Falon in xplanations of
1989). This aspt of hritability is espially important in th ontt of any on of ths fa
disussions aout ultural volution. Not only an idas or mms intr- us avoid th natur
prtd as rpliators, ut in th ontt of human, as wll as animal, soi. aspets of ou bL
etis larnd haviors an transmittd to th nt gnration. From this judgmnt about t
astrat prsptiv thr is thus no ontradition twn th intriat ths haviors dor
pattrns of human history rfltd in th hangs and transformations of inrasingly realiz
human soitis and th asi prinipls of evolutionary iology. Thr ar, and managmnt
of ours, many additional fators sids gns that ontribut to human our rif analys
history. Nothing lss would xptd from th viwpoint of volution- ity notwithstand
ary thory aus mor ompl systms add many diffrnt lvls and ary iology th
dgrs of frdom' ut all this happns within th onfins of volutionary planations of hl
prinipls. 1959). Sn thu
It is important to not that this prsptiv dos not imply any fom of mutual rlationshi
iologial redutionism; rathr, th volutionary analysis of omplx sys- at rtain tims s[
tms, suh as human soitis,rquirs that w simultanously onsidr ffts mnts of th thn-
at all diffrnt lvls of omplxity. Indd, in volutionary iology th most dation of wid-rar
intrsting qustions always aris in situations whr any simpl planation th ys of an voh
rahs its limit and whr w find a onflit twn diffrnt ntitis. Ths On th on hand,
..major
ar th transitions in volution'' dsrid y th late John Maynard thn-inomplt st
Smith and irs Szathm6r tat ring aout somthing fundamntally nw, our prsnt undr
suh as th first multillular organisms or th first human soitis (May- aus it implis r
nard Smith and Szathm6r 1995).In ordr to surviv,thsnw ntitisnd srvation an voll
..ontrol'' th havior and slfish intrst of th parts; in highr organ-
to ass of volution
isms, lls hav to giv up thir own rprodutiv intrsts, and in a omplx to mphasiz only
soity with a high dgr of division of laor' ths ativitis also nd to identifid as th m
 uolutio nd Soiet 9

ad limits of what kind of pfopr- oordinatd.How thsrgulationsom about and volv is on of th most
th fitnssf thir arrirs or thos fasinatingprolms of volutionary iology (Margulis 1981).
os, howvr, ar th sujt of a Ths rif thnial ommnts aout the asi assumptions of volution-
ir'now ask in what way a spifi ary thory also hlp us s th long-standing ontrovrsy ovr natur vrsus
:aturor a form of havior an nurtur in a diffrnt light. In th ontt of volutionary iology this dis-
:ssan additional qustion. Bfor tintion is on only of dgr and not of kind. Thr is no lar.ut spara-
nss'thy must first mrg within tion twn natur (or iology) and nurtur (or ultur and soity).
i produts of volution. In othr A multitud of fators, oth iologil nd ultural, ontriut to th dvl-
lmntal ontxt of organisms is th opmnt and formation of any human ing; indd, our dvlopmntalpro-
tur.only thos phnotypi vari- gram rquirs all ths diffrnt stimuli as ruial inpts.
: raw matrial for natural sltion' As w hav larnd from volutionary iology, ompard with our losst
of thir arrirs. Th fat that d- rlativs,w ar orn prmaturly (Portmann 1951). On onsqun of
nration of variation has rntly this fat is that in human infants th dvloping rain is stimulatd y a mul-
fif 1996 Hall 1998). titud of nvironmntal nd soial fators. In addition, w hav larnd that
lanationsof soitis' a diffrnt in our volutionary history th mrgn of soial skills, inluding th volu-
'f intrst.As w hav sn, all that tion of languag, was muh mor important than aumlatd gnti diffr.
'h availailityof auratly trans- ns(Diamond 19921Kupr 1994). Of ours, prisly thos haratristis
Lipis ommonly masurd by hri- also nal larning and th transmission of aquird knowldg. In onlu-
y dfindas th orrlation twn sion, all vidn thus far points to highly intrativ rlationships among
in many ass th rason for this diffrnt fators (iologial and ultural) in human volution. onsquntly,
.d to th ffts of gns (Falonr i n p l a n t i o n so f h u m n v o l u t i o n n d s o i t y W n n o t g i v p i o r i t y t o
ially importnt in th ontxt of any on of ths fators ovr all th othrs. This ralization shold also hlp
only an idas or mms intr- us avoid th naturalisti fallay; knowing th volutionary history of som
rf human' as wll as animal, soi- aspts of ur havior and soial organization dos not imply any valu
l to th nt gnration. From this judgmntaout ths proprtis. But ralizing th volutionary rasons for
ntradition twn th intriat thshaviors dos hlp us if w want to nfor soital ols, a fat that is
hangsand transformations of i n r s i n g l yr l i z d y a g o w i n g o m m u n i t y o f l g a l s h o l a r s . o n o m i s t s .
lf volutionry iology. Thr ar, and managmntonsultants.
]s gns that ontribut to human our rif analysis of th rlationship twn volutionary thory and so-
from th viwpoint of volution- ity notwithstanding, it is also tru that throughout th history of volution-
ns add many diffrnt lvls and ary iology th majoity of ttmpts to apply volutionary prinipls to
ithin th onfins of volutionarv xplanations of human soitis hav n grandios failurs (Hofstadtr
1959). Sn through historians' ys' ths ass ar prim xampls of th
]tiv dos not imply any form of mutual rlationship twn sin and soity disussd in th first stion;
utionary analysis of omple sys- at rtain tims speifi soital lifs and assumptions rsonat with l-
t w simultanouslyonsidr ffts mnts of the thn-urrnt stat of volutionary thory and om th foun-
J, in volutionary iology th most ..planations,'
dation of wid-ranging of natur and soity. Sn through
ions whr any simpl xplanation th ys of an voltionary biologist, ths ass tak on a diffrnt maning.
it twndiffrnt ntitis. Ths on th on hand, on might rgu that ths ass ar a rfltion of th
sribdy th lat John N4aynard thn-inomplt status of volutionary thory, at last whn ompard with
)ut somthing fundamntally nw, our prsnt undrstanding.Takn at fa valu, this viw is prolmati -
or th first human soitis (}ylay- aus it implis a rathr niv onption of sintifi progrss. Anothr o-
to surviv, ths nw ntitis nd srvationan volutionry iologist would mak, looking ak at prvious
:rst of th parts; in highr organ- assof volutionary planations of soity, is that all ths proposals tnd
,dutivintrsts' and in a omplx to mphasiz only on or a fw lmnts of wht rnst ayr nd othrs hav
'or, ths ativitis also nd to idntifid as th multipl elmnts of volutionary thory (Mayr 1982).

40 uolution nd Soiet

This obsrvation allows uS to omin historial ontxtualiztion of vo. tion' ruially dpn
lutionary planations of soity (what was th sintifi, soial, politial, hrditary matrial.
and onomi ontt of ths proposals?) with valuation in trms of thir Galton saw possiil
sintifi mrit (what wr th prolms with and wrong assumptions of volution and th fu
ths proposals? what was lft out' and what was aggratd?).This ap- th Vitorian progr
proah allows us to analyz ths various proposals oth in rlation to thir aus and ontrol it
historial ontxt nd with rgard to thir sintifi ondnt. Unfotunatly' rding with th n
W do not hav th spa to disuss ths idas in dtail, ut w want rifly In nd Super
to prsnt a fw eampls of how volutionary idas wr applid to human progrssiv intrpr
soitis in th past for onluding with a skth of how prsnt-day vo- that som ar natur
lutionary biology approahsth issu of human soitis. soity is urrntly
It will om as no surpris that as soon as thr was volutionary thory, it no informd rdin
was applid to plain aspts of human soity. rrt Spnr' a popular ftions now and als
philosophr and advoat of idas similar to thos of Darwin, oind th prmn.
trm sLlruiul of tbe fittest to dsri th onsquns of natural sltion. Indd, ugnis l
Dtahd from its stritly iologial maning' this trm ould (and was) Amrian visionari
applid to a varity of soial ontts. It ould takn to support a onsr- joind th all for go
vativ status quo whn it was omind with idas and osrvations that gradually did sint
volution favors onsrvation of xisting traits and hviors. In this way drgirdd th ugn
volutionary thory providd a naturalisti and matrialisti xplanation of ugnis was not as
soial diffrns y dmonstrating how some popl ar mor sussful . tors in rtrospt.S
aus thy ar fittr in Hrrt Spnr's sns. Thir variations ar ttr ugni programs a
adaptd to th omptitiv soial nvironmnt in whih thy liv, and thy ugni idas' tog
thrfor naturally nough ris to th top. raial hygin' ontr
Not surprisingly, it was th grat industrialist Andrw arngi who most nat ..unworthy lif,'
nthusiastially mrad this intrprtation of volutionary thory as - oalition formd to <
plaining soity. But th ida of natural sltionwas lso usd in support of logial dfinition of
radial transformation of soity, as th initial positiv rption of Darwin mnt on ra. This
y l\4ar, ngls, and othr lading soial dmorats attsts.Thy, of ous' antidisrimination pr
mphasizd th transformativ powrs of natural sltion rathr than th rathr than iologi
onsrvativ ons. onsistnt with stud
\/hatall ths initial appliations of volutionary thory to soity hav in nS among humans
ommon is that thy fous mostly on omptition' struggl' and a progrssiv nti diffrns in m
intrprtation of (volutionary) history, as wll as a hirarhial notion f populations hav ls
diffrnt soial lasss and ras. Furthrmor' thir notion of omptition is nalogial rlationsh
mor or lss dirt or hands-on. Struggl was oftn sen as a onsqun of our spis in ast A
diffrnt strngths of individuals, groups, stats' or ras. Indd, th arly sujt of gnti dif
yars of th twntith ntury rought svl attmpts to adapt Darwinian find a growing ons
logi to work as a guid to intrnational affairs and politis (Zieg|er 1918). of mankind an i
\/hat all ths appliations of volutionary thory to soity missd was n dvlopmnts in th 1
undrstanding of th intrnal onditions of organisms and th mods of ial tratmnt on th
transmission of hrditary matrial' whih wr' aftr all, ruial lmntsof Th two most imp
Darwin's thor. and human) havio
Th ntrality of hrdity to th thory of volution Was not missd y tion of smingly altr
Darwin's osin Franis Galton. Inhis l-Irditr Gnjus (l869) h ralizd tions, and hr sp
that th umulativ fft of natural sltion. *...,i.u for sradual volu- and othrs (Hamilton

istorial ontxtualization of vo-
as th sintifi,soial, politial,
with valuation in terms of thir
with and wrong assumptions of
,,hatwas xaggratd?).This ap_
lroposalsoth in rlation to thir
sintifiontnt. Unfortunatly,
las in dtail, ut w want rifly
rary idas wr applid to human
a skth of how prsnt-day vo-
uman soitis.
s thrwas volutionary thory, it
lity.Hrrt Spnr,a popular
.to ths of Darwin, oind th
:onsquns of natural sltion.
ing, this trm ould (and was)
luld takn to support a onsr-
with idas and osrvations that
traits and haviors. In this way
: and matrialisti xplanation of
m poplar more sussful-
sns.Thir variations ar ttr
nnt in whih thy liv, and they
'ialist Andw arngi who most
on of volutionary thory as x-
tionwas also usd in support of
itial positiv rption of Darwin
lmortsattsts.Thy, of ours'
natural sltion rathr than th
iutionary thory to soity hav in
t i t i o n s, t r u g g | . n d a p r o g r s s i v
; wll as hirarhial notin of
o r ,t h i n o t i o n o f o m p t i t i o n i s
,as oftn sn as a onsqun of
stats' or ras. Indd, th arly
ral attmptsto dapt Darwinian
'ffairsand politis (Zieg|er 1918).
y thory to soity missd Was an
of organisms and th mods of
wr. aftr all. ruil lmnts of
, of volution Was not missd y
,ditr Geius (1869) h ralizd
tion, nssaryfor gradual volu-
uolution nd Soiet 41
tion, ruially dpnddon oth th vailaility and th staility of adquat
hrditarymatrial. Fousing on inhritnand stability aross genrations'
Galton saw possiilitisfor ontrolling populations and thry ontrolling
volutionand th futur of human populations. This was an ntiing ida to
th Vitorian progrssivist. Not only ould w hav progrss, ut w ould
aus and ontrol it through iology, Galton suggstdwith his all for good
brding with th ngativ.and positiv sltions of ugnis.
In nd Suprm (190), Gorg Brnard Shaw also rfltda mor
progrssivintrprtation, though still vry muh groundd in assumptions
tht som ar naturally bttr than othrs. !(/hy not aknowldg that our
soity is urrntly wakr than it ould and should aus w hav
.!7
no informd rding program' Shaw suggstd. an plain our imp-
ftionsnow and also sk to improv, gnrating a soity of mn and su-
prmn.
Indd, ugnis rought hop for progrss spially to uropan and
Amrian visionaris in th arly twntith ntury. Lading gentiists
joind th all for good rding and for ttr soity through ilogy. only
gradually did sintists apt how littl solid sintifi knowldg rally un-
drgirddth ugni assumptions' so that in th 1910s and into th 1920s
ugnisWas not as ridiulously unsintifi as it has smd to ommnta-
tors in rtrospt. Soial sltiv prssurs rinford thos who adoptd
ugniprograms asd on biologial laims (Kvls 1985; Paul 1995).
ugni idas, togthr with a iologial onpt of raial supriority and
raial hygin,ontriutd to th Nazi gnoids' th ampaign to trmi-
nat ..unworthy lif,'' and finally th Holoaust. As a onsqun' a growing
oalition formd to ountrat what was privd as an inappropriat io-
logial dfinition of human ras, ulminating in th 1950 UNSO stat-
mnt on ra. This statment thn am th foundation for furthr
antidisriminationpoliis and laws. Ths dlarations mphasizd thni
rathrthan iologial diffrnstwnhuman populations, whih smd
onsistnt with studis that showd that th vast maiority of gnti diffr-
nsamong humans our within populations. Howvr' in rntyars g-
nti diffrns in mitohondrial and Y-hromosomal DNA twn human
populations hav also n usd to illumint migrarion pattrns and g-
nalogial rlationships twn hman populations' inluding th origin of
our spisin ast Afria aout 130'000 yars ago. Today, vn though th
sujtof gnti diffrnstwn human populations is still touhy, w
find a growing onsnsus that nithr th iologial nor th ultural history
of mankind an ignord. This is spially tru as svral thnologial
dvlopmntsin th postgnomiag inrasour ailitis to ustomiz md-
ial tratmnt on th asis of th gnti onstitution of patints.
Th two most important thortial insights into th volution of (animal
and human) havior wr Hamilton's notion of kin sltionas an plna-
tion of smingly altruisti havior and th ppliation of gam-thorti no-
tions, and hr spillr th onpt of stratgis, y John aynard Smith
and othrs (Hamilton 1|64;anard Smith |9s2). Ths idas rprsntda
42 uolutil d Soit

major hang in volutionary plnations of havior and st th stag for mpirial data and th
th dvlopmnt of rigorous mathmatial modls. mal and human ehav
Th thory of kin sltion nd of inlusiv fitnss fousd th dat on Th soioiolgy dt
th appropriat units of sltion in planations of havior. Rathr than lutionary xplnation
just looking at on individual's fat as th sol dtrminant of fitnss, th th- of th many dimnsio
ory of kin sltion hangs th wys in whih vo/utionary aounting is ism, supportd y his
.sithin ulation gntis at thil
don. th framwork of population gntis hangs, volution is rp-
rsntd as hangs in gn frqunis. Wht Hamilton ralizd Was that intrativ prsptiv
thr is mor than on way in whih a spifi gn in a spifi individual rs' and his ritis w
an mak it into th nt gnration. It an passd on to its offspring sintifi, nvironmn
dirtly or through th produtiv suss of Ios rlativs, who will shar molular iology). T
th sam gn with a rtain proaility (.5 for silings, .125 for ousins).In ns of that tim, wI
alulating th fitnss of a spifi gn' on thus has to fous on th inlusiv dats.
fitnss, whih is th sum of all th diffrnt mans y whih a spifi gn Today, mor than
an om to rprsntd in th nt gnration. tionary planations (
Prvious vagu notions, suh as th ida that a havior an volv - popular within iolo;
aus it is good for th spisor th group' wr rplad y onpts that asis for this rsarh
aordd with th prditions of population gntisand th thory of natu- ral dbat and or
ral sltion. Howevr, it is also important to not that ths notions Wr ogy. Yt som of todi
highly astrat, and whn thy wr first proposd, sintistsdid not rally soitis bttr rfl
know anything aout how ths proposd gneti rpliators atually ould any diffrnt filds
aus th rsptiv haviors. At this tim th argumnts wr lrgly tho- study of th volution
rtial: if a gn/rpliatorauss altruisti havior, it an favord by as for intrdisiplin
natural sltion as long as th ost to th individual ating slflsslyis kind of rsarh;as
smallr than th nfit to its ripints multiplid y th offiint of rlat- havior' it is in dan
dnss twn thos individuals. A similar approah guidd th many bat, vn if sintifi
gam-thortionpts, suh s th prisonr's dilmma and idas aout r. aout volution in g
patd gams and riproal altruism that wr usd to plain many asPts
of animalhavior.
Th appal of ths nw idas Was normous, and iology smdapal onlusions
of xplaining muh of soial havior. Th idas providd an important
oost to many disiplins, suh as havioral ology, ut also invitd far- volutionary thoy i
rahing gnralizations and in turn gnratd many' ftn polmial' ontro. th sins, iology
vrsis.Th most visil of th sandalsruptd with th puliation of . o. ogy and iomdiin
.lfilson's hildrn's, lot will im
Sobbiolog tn 7975.In this nylopdi tom./ilson' an prt
on ants and soial insts' olltd an normous amount of mpirial data aout th ngativ
on what Was thn known aout animal havior and organizd it in th futur of human soit
ontXt f th nW thortial framwork. In th final haptr h applid to at stak. Biolog;
ths idas to human havior and th volution of human soitis.Fur- srvativ iongings for
thrmor, h proposd that this nw synthsis would lad th soial sins us on th history of
to intgratd within th framwork of th iologial sins.Th ra- stag.
tion was vivid. Howvr' nsuing dat fousd mostly on th feard politi- It sems almost inl
al impliations-Wilson's most voal ritis long t th Lft-nd on th assoiatd transformz
onsquns of th rdutionism impliit in l/ilson's mphasis of gnti . triggr ration y ful
planations and in his modl of th rlations amog t\sins. Surprisingly ontxt of volution
littl attntion ws paid to th ulk of th ook. th normous amount of mnts highlights imp

of haviornd st th stag for
odls.
siv fitnss fousd th dat on
nations of havior. Rathr than
ol dtrminantof fitnss,th th-
w h i v o I u t i o n a r y o u n t i n g i s
g n t i s h n g s ' v r l l u t i o ni s r p -
Vhat Hamilton ralizd was tht
ifi gn in a spifi individual
:an passd on to its offspring
s of los rlativs, r,r.howill shar
i f o r s i l i n g s ., 1 2 5 f o r o u s i n s ) .I n
l ths has to fous on th iniusiv
tt mans y whih :r spifi gn
sration.
la that a havior irn volv -
p' Wrerplaed y onpts that
n gntisand th thory of natu-
t t() not tht ths notions Wr
rroposd,sintistsdid not rally
gntirpliators atually ould
l th argumntsWr largly tho-
i havior,it n favord y
th individual ating slflssly is
rltiplid th offiintof rlat-
rilar approah guidd th many
n'sdilmma and ids out r-
rusd to plain many aspts
nlus,nd iology smdapal
.h
idsprovidd n important
lral ology, ut aiso invitd far-
:dmny, oftn polrial, ontro-
Lptdwith th puliation of . o.
ylopditor $ilson' n xrt
)rmous amount of mpirial data
havior and orgar-rizdit in th
r. In th final haptr h applid
. o l u t i o no f h u m a n s o i t i s .F u r _
siswould ld t soial sins
tlr iologial sins.Th ra-
lr'rsd mostly on th fard politi.
s long to th Lft-and on th
n W i l s o n ' s m p h a s i so f g n t i x -
s aog th sins.Surprisingly
: ook, th nornrous amount of
uolution d Soiet 4
enrpirildat and th rigorous appliation of volutionary prinipls to ani-
lnlnd humn hvio.
Th soioiology dat rsnrld th many othr attmpts to pply vo-
lutionary planations to human soitis in that it ovrmphasizd just on
'Wilson's
of th many dimnsions of volutionary thory. gnti rdution-
isr-l-l,
supportd y his prin with soil ants' as wll as th stat of pop-
ultion gntisat this tim, lft Iittl oom for othr fators or for th mor
intrativ prsptiv that has mrgd mor rntly. \Vilson' his support-
rs' and his ritis Wr all produts of thir soial and politial, as wll as
sintifi, nvironmnts (at that tim dominatd y th rnt susss of
molular iology). This ld to a rathr polarizd limat within th lif si-
ns of that tim, whih in prt plains th onfrontational ton of ths
dats.
Today, mor than 30 yars ftr th puliation of Sociobiology, volu-
tionary xplanations of soity and disussions of ltural volution ar still
popular within iology. Howvr, the onptual, as wll as th mpirial,
sisfor this rsarh has om muh mor pluralisti. Thr is still onsid-
raldt and ontrovrsy,spially surrounding volutionary psyhol-
ogy. Yt som of today's approahs to undrstanding human (nd animal)
sitis ttr rflt th insights of volutionary thory skthd arlir.
any diffrnt filds of volutionary iology ontriut' and indd th
study of th volution of ompl soial systmshas om a paradigmati
asfor intrdisiplinary rsarh.As suh, it fas all th hallngsof this
kind of rsarh,; as ah spilty tks on on spt of soity and human
bhavior, it is in dangr of ovrmphasizing that part. And th nsuing d-
at, vn if sintifially produtiv, is rivd y a soity that is unasy
aout volution in gnral.
Conlusions
volutionary thory is unqustionaly th foundation of iology. And of all
th sins, iology has th largst impat on today,s soitis. Biothnol-
ogy and biomdiin arry th hops of illions that thir, or at last thir
hildrn's,lot will impov, whil nvironmntal sinswarn almost daily
aout th ngativ onsqns of our own ations. Nothing lss than th
ftur of human soitis-som might all it thir volutionary fat-sms
to at stak. Biology offrs oth gloom and glory, utopian drams and on.
srvativlongings for an idalizd past, and volutionary iology with its fo-
us on th history of lif, s wll as th manisms of hang' is at ntr
stag.
It sms almost invital tht today's rapid sintifi dvlopmnts and
assoiatdtransformations of soity and human (slf-)undrstanding might
triggrration y fundamntalists' with thir dsirs for sur answrs. In th
ontt of volution an| soity, th popularity of fundamntalist mov-
mnts highlights inrportant filurs in ommuniating what is promising in
44 uoltio nd Soit

th fundamntal prinipls of volutionary thory and in ploring implia- without ing undul
tions and limitations of ths sintifi insights for our urrnt soitis and ial organization and
for human slf-undrstanding (sth main ssays ..volution and Rligion'' prinipls of th rpli
y David N. Livingston and ..Amrian Antivolutinism: Rtrospt and produt of volution
Prospt'' y ugni C. Sott in this volum). l/ thus of{er a fw rmarks nomna (suh as th
on futur dirtions. studied within just or
volutionary analysis of human havior and soial organizations will r- solvd whn thy art
main an important rsarh ara within volutionary ilogy and will hav a urrnt xtnsion of
high potntial for insights, as wll as for onflit. volutionary mediin is al- ontrvrsil ontinu
rady transforming th ways w undrstand and trat rtain disass (s of ours, all this
th main ssay ..volutionary Biology of Disas and Darwinian diin'' tionary thory might
y ihal F. Antolin in this volum). volutionary psyhology offrs in- vid a justifiation. I
sights into th history of human havior and our potntial to modify and for thial thory, fl
ontrol havior, inluding possil transformations in our lgal systms and ..oug
annot lad to
pratis (sth main ssay ..SoialBhavior and Soioiology'' y Danil I. hav,xpt insofr
Runstin in this volum). Th mrging of onomi thory with volution- ory as good sini]
ary thory nfits oth filds and has givn volutionary thory W OnP-
tual and mathmatial tools, suh as gam thory and th notion of stratgis.
It has also introdud nw qustions y rplaing traditional idas of an ..in- B|LIoGRAPl-
visil hand'' with th havior of individual ators shapd y thir volu-
tionary history and onstraints. Appliations of volutionary thory vn Akrt, L. 2007. Th ,.
,l'885-1940.
transform rtain filds of nginring and omputr sin as th prinipls Jlur
A d a m s , . B . 1 9 9 4 .T |
of volutionary dsign and of gnti algorithms hav opnd nw vnus for -hought
d i Rt
solving omplx optimization prolms, suh as th dsign of airplan wings Bowlr,P, l, 1983.h
(Rhnbrg197i Ho|Iand 1995 ). i the Decdsr
Th rlationship twn volution and soity also provids an important Boyd, R., and P. J. Ril
as study for intrations twn sin and soity mor gnrally. Th Univrsityof hi
B r n d o n 'R . N . , n dR .
history of volutionary thory is among th st-studid aras in th history
otrouersislu
of iology and has rvald intrsting ways in whih sintifi rsults mrg Brown,J. 1995.hr
in a ompl intrplay twn sintifi invstigations and what an 203' hrls-
alld thir soial and ultural ontts. Although ths studis hav ld to Univrsityrss.
insights aout how sin happns and thus hav ontriutd to popular un. Dwkins, R. 1976,h
"1982.The t
drstandings of sin, thy also hallng philosophrs of sin to rthink .W.
. Frman.
som fundamntal assumptions aout th natur of sintifi vidn and Dglr,. 1992.I S
how historially ontingnt xpimntal and thortial pratis an lad Driism i Amt
to inrasingly ..aurat'' rprsntations of natural phnomna. Som vn Dsmond'^. J' 1997.I
go so far as to dsrib this ..soial'' pross of finding th ..truth'' in sin Rading'A: Ad
in volutionary trms' thus oming full irl. Dsmond,A. J., andJ.
Diamond, I. . 1992-
But th most important aspt of th ongoing intrations twn volu-
HumnAimI,N
tion and soity lis in th ways in whih volutionary thory transforms our ldrdg,., and S.J.
slf-undrstanding as oth iologial and soial/ultural ings. As propo- phyltigradualis
nnts of ultural volution suggst' thr is no inhrnt differn twn San F'raniso:Fr
th ways our odis and haviorsand our soitisand ulturs volvd.In- F a l o n rD' . S . 1 9 8 9 .1
G l t o n 'F . 1 8 6 9 . H r
dd, th rigorous appliation of volutionary thory to suh aras s th
London: Mamilla
volution of languag and of ultural transmissin.hasallowd ths propo-
Hall, B. K. 1998.uolt
nnts to formulat a thory of human soity and ultur that is matrialisti all.

'thr and in ploring implia.
ightsfor our urrnt soitis and
..volution
1 ssays and Rligion''
Antivolutionism:Rtrospt and
m). W thus offr a fw rmarks
r and soial organizations will r-
lutionarybiology and will hav a
nflit.volutiorrarymdiin is al-
nd nd trat rtain disass (se
)isasand Darwinian Mdiin''
volutionary psyhology offrs in.
and our potntial to modify and
)rmations in our lgal systms and
ior nd Soioiology'' y Danil I.
[ ononrithory with volution-
r volutionary thory nw onp-
hory and th notion of stratgis.
llaing traditional idas of an ..in-
.ual ators shapd y thir volu-
ions of volutionry thory vn
omputrsinas th priniples
ithmshav opnd nw vnus for
h as th dsign of airplan wings
;oityalso provids an impotant
: and soity mor gnrally. Th
st-studidaras in th history
s in whih sintifi rslts mrg Brown,J. 1995. hrltlsDri: Volgittg,N'*'York: Knopf.
invstigationsand what an
'lthough ths studis hv ld to
ls havontriutdto popular un-
philosophrs of sin to rthink
natur of sintifi vidn and Dglr,. 1992. [ Srhlf Hum Nture: Th Dclie d Ruiulof
lnd thortial pratis an lad
of naturl phnomna. Som vn
;s of finding th ..truth'' in sin
:l.
going interations twn volu-
olutionarythory transforms our
soial/ulturalings. As propo-
is no inhrnt diffrn twn
soitiesand ulturs volvd. In-
nary thory to suh aras as th
;mission.hasallowd ths propo-
ty and ultur that is matrialisti
uolutio nd Soiet 4''
without ing r"rrrdul gn ntrd and rdutionisti.Th insight that so-
il organization and ultual tanstnissiona govnd y the sam gnral
priniplsof th rpliator quirtion implis (1) that not vrythingthat is th
produt of volution has to grrtiallydtrmind and (2) that many ph-
nomna (suh as th volution of oopration) that rmain puzzling whn
studidwithin iust on l/l of analysis (gntior soial/ultural)an r-
solvd whn thy ar approahd from oth dirtions. \/ithoutdout this
urrntxtnsion of volutionary theory to soity will an important and
ontrovrsilontinuation of th long history of volution nd soity.
.!
of ours, all this an takn too fa. must kp in mind that volu-
tionary thor might provid an planation for soity' ut it nnot pr-
vid a justifiation. volution also annot provid an pistmi justifiation
for thial thory, for ampl, dspit th numrous attmptsto do so. ..Is''
annot lad to ..otrght.''vlution annot tll us how soitis ought to -
hav,pt insofar s it follows that soitismst apt voltrtionaryth-
ory as good sinif thy gr to apt sinirt all.
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of rligi
(quotd in Brown 2
natur of Darwin's o
volution and Rligion at what point? Did
to do with it? Did
ntivolutiorristsali k
Duid N. Liuingston h ida that Drv
ontstd y thos l
to th nd'' (Moor
spiritual momnts lr;
ooks dating frorl tl
..Sopy Sm'' \trilrfor,th ishop William Paly's o
vr sin th mythi nountr twn
..Darwin,s sign; and th prson
of oford, and Thomas Hnry Huly, ulldog,'' at th 1860 mt.
trin'' of trnal rtri
ing of th Bitish Assoiation for th Advanmnt of Sin' thr has na
dath of his dlightf
widspad snsof an ongoirrg and invital lash twn volution and r-
thos who insist tlrtrt
ligion. Th lristorial narrtiv of tlris nOulltr' howvr, provs to th
mind, and motion (
lss lar-ut that suh irnprssionism miglrt suggSt.In th pags that follorv
athism or mtapy
w xamin th rligious viws of Darwin himslf and somthing of th histo-
ims\i tutnd arar
riography of th date for introduing atholi, Protstnt' Jwish, and
rusading sulrist
Islami rsponssto his thory. A numr of rurring thms 1rgfrom this
Sin irrslrtti
historial survy' and a rviw of thsissusorrltldsthis ss:ly.
iudgmnt of ths
ing to diagnos D:
hsitany ar mor
Darwin's Rligious volution wavring to whih -
fr xampl, h r,vr
Syms Covirrgton Was }lS Bgl's fiddl, ship's oy, and odd'jo man. For this is alwavs pair"
just ovr si yars h was also harls Darwin's srvnt.Bush was th athistially. But I r
evr-prslltassory to Darwin's gogrphi arrd sintifi travls, it is wish to do, vidn
hardly surprising that this virtually invisil Darwinian adjunt would othr hnd I anno
soonr or latr ath a novlist's imagintion. Th final fw sntnsof spially th natur
Rogr Donald's fitionirl aount of <lvirrgtontn r. Drtuin's Shootr brut for'' (Burkh
(1998' 410) rystalliz an irrrliguityat th hart of any aoDnt t>fth l- of vaillation sttrf
..H saw Darwirr on his kns,ad ..trukl''
tionship twnvolution and rligion: to puli
thr was no diffrntwn prayr and pulling a Wor from th grass. th origi oi S1li
As for r ovington, h prayd in th old-fashiond way.'' Th lision - ] 00 tims. Aor'rn
twn th transndntalrrd th asthti tlrat is apturd hr rsonats Josph Dalton Ho]
with Mihal Rus's onlusion to his rnt ilount in Drtuitl d Dsig. 1879 note tl J' For<
Fully irwirof Darwin's rpatd rsort tO th languag of uty and worrd tuats'' (quotl in
..of (|986, 27) onlus
in th ntural ordr, Rus (2003, 335) rn-lindshis raders th gnuin
lov and joy''-an ..ovrwhlming xprin'' touhing on th spiritual- to and flow..
that volutionists snsin thir nountrsr'vithth organi rvorld. Hr w mti athist; at hig
..Hosannah'' in intoirtddlisht asially agnosti-
ar alrtd tl th Drwin who brathd
at th sulinrity of th primval Bzilin rin frlrst. mit hirnslf o suh

48

ligion
Soapy Sam'' Wilrfor' th ishop
)arrvin'sulldtlg,'' at th 1860 mt-
nmntof Sin,thr has n a
al lash twnvolution and r-
)untr' howvr, provs to rathr
;ht suggst.In th pags that follow
himslf and somthing of th histo-
rg Catholi, Protstant, Jwish, and
cf rurringthmsmrg from this
u s o n | u d tsh i s s s a y .
:r,ship'soy, and odd-jo man. For
arwin's srvant.Baus h was th
3raphi and sintifi travls, it is
rvisil Darwinian adiunt would
nation. Th final fw sntns of
Covington in r. Dri's Sholter
h hart of any aount of th rela-
.. saw Darwin on his kns, and
rnd pulling a Worm from th gass.
ld-fashiondway.'' Th lision -
rti that is apturd hr rsonats
]ntaount in Dri d Dsig.
l th languagof auty and wondr
'mindshis raders ..of th gnuin
in''touhing on th spiritual-
rs with th organi world. F{r w
..Hosannah''in intoiatd dlight
rain forst.
uolutkln d Religio .t49
Givn th omplitions that lsvationsof this lass introdu, not to
mntionThomas Hnry Huly's rmark to harls Kingsly in 1860 that a
..dp
sns of rligion is ompatil with th ntir asn of thology''
(quotdin Brown 2002, 10), it is not surprising that opinions diffr on th
naturof Darwin's owi rligious onvitions. Did h los faith ntirly? If so'
at what point? Did his thory of volution y natural sltionhav anything
to do with it? Did h undrgo a dathd onvrsion? volutionists and
antivolutionistsalik hav a stak in th ft of Darwin,s soul.
Th ida that Drwin movd inxoaly from lif to agnostiismhs n
ontstd by thos who ar onvind that h rmaind ..muddld thist
to th nd'' (oor 1985' 438). Similarly, a vrity of supposdly diagnosti
spiritual momnts hav n idntifid: th matrialism disrnil in not-
ooks dating from th lat 180s; his growing douts aout th adquy of
!illiam Paly's aount of ration with its roust onfidn in divin d-
sign; and th prsonal moral dilmmas h suffrd ovr th ..damnal do-
trin'' of trnal rtriution with th loss of his father and th snslss,rul
.re
dath of his dlightful daughtr Anni (oor 1989, 797). ar' too'
thoswho insist that an intrst in ultivating a naturlisti aount of spis,
mind, and motion annot onstrud s vidn of ithr thoroughgoing
athismor mtaphysial mtrialism (Gillspie |979) vn though Darwin
himslf turnd away from hristianity. Rgardlss, th imag of Darwin as
rusadingsularistfails to do justito th omplitis of th as.
Sin irrsolution on Darwin's spiritual stat sms to b th olltiv
1udgmntof thsportrayals, th languagof rtainty nd prision in sk.
ing to diagnos Darwin's spiritual ondition is misplad. Amiguity and
hsitanyar mor appropriat' all th mor so sin thy onvy th snsof
wavringto whih Darwin himslf gav voi. In an 1860 lttr to Asa Gray,
for ampl,h wrot: ..\ithrsptto th thologial viw of th qustion;
this is always painful to m.-I am wildrd.-I had no intntion to writ
athistially.But I own that I annot s' as plainly s oths do, & as I shd.
wish to do, vidn of dsign & nfinon all sids of us . . . on th
othrhand I annot anyhow ontntdto viW this wondrful univrs &
spiallyth natur of man, & to onlud that vrything is th rsult of
brutfor'' (Burkhardt and Smith 1985_2005, 8:224)' Thn agin his sns
of vaillation surfadin his shifting judgmntsaout whthr h was wis to
..trukl'' puli
to opiniln nd rfr to th Pntatuhal term rltion io
the rigi of Specis(1859), thoghh atually usd it nd its ognatsovr
100 tims. Aount too must takn of his ommnts in an 1870 lttr to
..my thology is
JosphDalton Hokr that a simpl muddl'' and thn in an
1879 ote to J. Fordy that on thologial mattrs ..my judgmnt oftn flu-
tuats'' (quotd in Brlwn 1986,25, 1). Suh rmarks onfirm Brown's
(1986,27) onlusion that Darwin's rligious lifs ..nvrntirly asd
to and flow . . . At low tid, so to spak, h was ssntially n undog-
matiathist;at high tid h Was tntativthist;th rst of th tim h was
basiallyagnosti-in Sympathywith thism ut unal o unwilling to om-
mit himslf tln suh impondralqttstions.''
.'0 uolutio d Religion

Th impat of rligion on Darwin, howvr, annot rstritdto flutu- prsidnt of ornll

ations in his spiritual trnpratur.Th influn of Paly's Nturl Tholrlgl' Theolog i bristr
(1802), for ampl' lirrgrdlong. Th onnrions Paly livd h oLrld muh to onfirm th i
disrn twn divirr rtivity nd human agll),providd Dawin with fors of nlightnd l
an asoring ar-ralogy:natural sltion' Nature, Drwin judgd, was in- But th risp larit
volvd in th sltion of organi forms rathr lik th mannr in whih pi- Stration for histori
gon fanirs pikd out th most promising variations. As h lrimslf rd. For a start' at ii
famously put it: ..Natural sltionis daily and hourly srutinizing' through- of th Gnsis rord
out the world, vry variation' vn th slightst;rjtingthat whih is ad, stratgis to maintain
prsrving and adding up all that is god'' (Darwin' |959, 168-169).In laoratd, and th id
som wys Darwin's Natur took on th attriutsof Paly's God. T this w minds of arly Vitol
right add tha suh onpts as organi adaptation and th harmony of n- took a varity of fo
tur wr as tral to Paly's osmogony as thy Wr to Darwi's. Yt Dr- hallng, and of ou
win's ognitiv indbtdnssto thology i1lltlot limitd to arhittural Anglianism and stri
hos of Paly. Sor of his most profound onvitions mrgdin dialogu th dsign argumnt \
with-or in dfian of-nvntional hristian dotrin. His ritiqu of and William 'Whw
natural thology, it has n suggstd,was umilially onntd to his ration y natural l
growing onvitions about th links twnhunran and animal-how oLrld opnd th door to tl
th produts of an adv:rnd monky-mind trustd? His allrgy to th ways in whih volut
miraulous was inflamd y a rading of Paly's L,ideesof hristiltit takn, nd ontinu tr
( \ 7 9 4 ) ; h i s s n s i t i v i t , tvo s r . r f f e r i n ga,s D n a l d F l m i r r g( 1 9 6 1 . 2 3 1 ) s p u - What furthr omp
latd, may hav owd its intnsity to a ..yaningaftr a ttr God than ists and rligious l
God''; and his douts aout tlology w nurturd y an inaility to r- th sid of sular z
onil th dotrin of divin providn witlr lif's daily dtails (Brook Thoms Hnry Huxl
1985). Takn ovrall' sinand rligion wr, in on Wy or anothr' thor. lub, who ddiatdt]
oughly intrtwind in Darwin's lif and thought. alik. No soonr did
of volution than h f
ology to insist that
laring th Grund nism to that vigoro
moral' and soial lif
On th fa of it, Dawin's thory posd lrallngsof pi propotios to 1991' 308).Thn, in
hristian lif at vry turn. It mythologizd tlr osai narrtiv of spial through gologial sg
ration; it smashd through th prati of using ilial gnalogisto dat rap, murdr & ars
arth history; it rmoved th ida of divin dsign from natur y dmon- (quotd in Dsmond
..sintifi
strating how spisam aout through th rdinary, humdrum prosss out th 1
of natral sltion; it rvald that humans wr! in som fundamntal to ration''' dnomi
..t
sns)no diffntfro-raninrzrls;it rootd morl snsibilitynot in th hun-ran daining frinds to
sujt dignifid as God's imag r ut in th pimitiv irnpuls of a tlolog'''
struggl fo survival; it ws nt-door niglror to th Frnh m:rtrialism At th sam tim a
tht had n infiltr:rtinginto Britain through Lamarkian-nthusdmdial ial agndas in dlivi
radials who moilizd it to attak th Anglian-Tory stalishmnt.Not as of Drapr, who
surprisingly, th idea of a protratd attl trvn sin and rligion r- mation' th ral obj
ivd its gratst imptus from nintnth-ntLlrytratiss that hartd, as dotrin of papal infa
in th as of hmist-tLrrnd-historian John Dpr, te Histor of tb o- prophsy that it woul
flit bttu Rligio d Sice (1875 ) or, in th rvords of Andrw \ffhit, frindly govrnmnts
 uolutil d Religio . 1 - \I

.vr'annot rstritd to fltu- prsidntof ornll Univrsity, A Histor of th Wrfr of Siee luith
flunof Paley's Ntrl Tholog holog i hristedom (\896). Th vry titls of works lik ths did
:onntionsPaly livd h ould muh to onfirm th imprssion of a monumntal dath struggl twn th
man agnyprovidd Drwin with forsof nlightnd sin and nightd rligion.
. Natur, Darwin judgd, was in- But th risp larity f this rivd imag sustituts monohrom ab-
athr lik th mannr in whih pi- stration for historial thniolor. Thr is a rihr narrativ to unov-
,,days',
mising variations. As h himslf rd.For a start' at lst sin ugustine, many ad read th ration
[y and hourlv srutinizing' thrugh. of th Gnsis rord symolially rathr than litrally. Various harmonizing
lightst;rjtingtht whih is ad, stratgisto maintain onord twn Gnsis and gology had long n
,od" (Darwin, 1959, 168-169). In laoratd,and th ida of a lngthy arth history was wll stalishd in th
lttriutsof Paly God. To this w minds of arly Vitorian hristian gntlmn.gologists. Natural thology
adaptationand th harmony of na- took a varity f forms' som lss susptil to Darwin's antitlologial
.as thy wr to Darwin's. Yt Dar- hallng' and of ours thr wr traditions, lik rtain strands of high
nnot limitd to arhittural Anglianism and strit unsntimntal alvinism, for whih th signifian of
nd onvitionsmrgd in dialogu th dsign argumnt was ngligil. And thr wr thos lik John Hrshl
hristian dotrin. His ritiqu of and William !hwll whos rligious outlook prdisposd thm to strss
, was umilially onntd to his ration y natural law rathr than y divin intrvntion' a mov that
nhuman and animl-how ould opndth door to th ida of volution as God's mthod of ration. Th
rind trustd? His allrgy to th ways in whih volution and religion ould mad to fit on anothr hav
>fPaly's uides of hristiit takn,and ontinu to tak, myriad forms.
lonaldFlming (1961, 231) spu- lfhat furthr ompliats mattrs is that whn onflit twn volution-
..yarning aftr a ttr God than ists arld rligious livrs did our, th ampaign was oftn wgd from
r nurturd y an inaility to r- th sid of sular zalots as muh as from thologial rationaris. Tak
: with lif's daily dtails (Brook Thomas nry uly, a ky mmr of that Darwinian gingr group, th
l Wr'ln on Way or anothr' thor- lu,who ddiatd thmslvs to driving out of powr Palyits and parsons
hought. alik' No soonr did h har of th atholi St. Gorg Mivart's advoay
of volution than h frrtd out a opy of Franiso Suarz's Sholasti th-
..omplt
ology to insist that volution was in and irronilal antago-
nism to that vigorous and onsistnt nmy of th highst intlltual,
moral,and soial lif of mankind-th atholi hurh'' (quotd in Brook
c hallngsof pi proportions to 1991, 308). Thn, in irritation at th stratgisof thos wh rad Gnsis
..prov that
1izdth osai narrativ of spial throughgologial sptals,h dlard his dtrmination to
..positivly
of using ilial gnalogisto dat rap,murdr 6( arson'' wr njoind'' in th old Tstmnt
zin dsign from natur y dmon- (quotdin Dsmond and Moor 1991, 472). Bsids,h was forvr talking
..hymns
l th ordinary, humdrum prosss aoutth ..sintifipristhood,'' prahing ..lay srmons,'' singing
lmns Wr' in som fundamntal to ration''' dnominating himslf a ..Bishop'' of th nw lsiology, or-
..th ..molular
l moral snsiilitynot in th human dainingfrinds to hurh sintifi,'' and pounding his
ut in th primitiv irpuls of a tlolog.''
righor to th Frnh matrialism At th sam tim advoats of onflit historis hav also had thir own so-
ough Lamarkian-nthusd mdial ial agndasin dlivring pugilisti ounts of sin and rligion. In th
Anglian-Tory stalishmnt. Not asof Drapr' who rmarkd that sin was th twin sistr of th Rfor-
'tl twn sin and rligion r- mation' th real ojt of opprorium Was th Romn atholi hurh; th
th-ntuytratissthat hartd' as dotrinof papal infalliility nuniated in 1 8 70 appalld him and ld him to
lhn Drapr, te Histor of tb on- prphsythat it would ring th atholi hurh into onflit with hithrto-
.!hit' .W.hit,
) or' in th words of Andrw frindly govrnmnts. As for th opposition that his nonstarian
52 uolutio d Rligil

ornll Univrsity had attratd from lrgymn strngthnd his solvto Frnh Dominian pri
kp sin at th forefront of th urriulum; ornll would an asylum o r g i q s( 1 8 8 7 ) ,a n d
fo sin liratd from th striturs of rligius dogma. It was manuvrs rol in th dvlopmnt
of this kind that promptd som historians to ronptualizth so-alld |ished uolution nd Dt
warfar twn sin and rligin in Vitorian soity as a soial struggl Both ths writrs su1
twn two ontnding intlltual lits-namly, th old-fashiond par- it within rtinlimi
son and th nwly professionalizdsintist (Turnr 1978|.In this snario foundations, was dirt
dbats aout sintifi knowidg in gnral and volution in partiular sim. thologial nmis of t]
ply am a furthr arna in whih tussls for ultural suprmay wre might th produt
played out. Sard ongrgation ol
. . v o l u t i o nt h o r yi
Howvr alluring grand narrativs may , storis of th nountr - tt
twn volution and rligion that trad in systmi onflit or, for that mat- human ody'' (quotd
tr' oopration srifi historial intriay t linar simpliity. In fat, ook was nsrdwas
rligious liversrspndd in vastly diffrnt Ways to vlution, and thir spdily rtratd in tht
stans do not follow any straightforward taxonomy of thologial orinta. find ways of rvisiting t
tion' dnominational affiliation, or dotrinl proupation. Bsids,volu. of th ook, ut it too :
tionary thory Was nvr nountrd in a ultural vauum' and dats It was muh th sam
aout it routinly am a mans of giving voi to a rang of othr ani- rognition for his S
tis. Somthing of th divrsity of ths ngagmnts my gland from tb oli Sietists (789
th following thumnail skth. st of 1 as th aU
nd Dogm was to d
..to admir'
muh in it
Th Viw from Rom xx). Likwis ground
ti gsisto justify a
atholi opinion on volution has long n dividd. From th arly days th intrnational suss,a
opposition of a group of Jsuit thinks rought togthr y Pop Pius I to of a fuming iuilt t
omat th fors of modrnity Was prssd through th pags of La lnt with rklss ass
iuiltd ttolic' an outlt that rtd vry onsidral influn among sumptions. Th ongr
ItalianJsuits. Thus in th arly 1860s th nssityof maintaining th fiity vry largly on th ma
of spisagainst Lamarkian idas of transformism was vigorously promul- lutionism was part of z
gatd by G. B. Piniani in its pags (Brundll2001). By ontrast' th fforts tion of ttul ritii
of th nglish omparativ anatomist St. Gorge Jakson ivart' a atholi lsiastial affairs. L
onvrt' to histiniz volution through a Lamarkian rading of transmu. voral rviw h r
tation in his 1871 volum the Gesis of Species was praisd in th pains, prSntlyam l
Catholi prss. Huxly might snip that Mivart ould not a sturdy sldir Various fators play
of sinand a loyal son of Rom, ut Pop Pius I wardd him th dgr i51''-1hg tndny or
of dotor of philosoph ln 1876, and the Blgian ishops prssdhim to tak indpndn and to :
up a hair t th Univrsity of Louvain in 1884, although h latr fll into suh for. This indr
disfavor on aount of rtain thologial writings. ntury atholis wr
Howvr voifrous th opponnts of volution undoubtdly wr, th of- with whih immigran
fiial ustodians of Catholi dogma did not tak any ation against Catholi rns rotatd round t
volutionists until th 1890s whn attituds gan to hardn as th aging arly trnds, notably t
Pop Lo III, hithrto a for for modration, lost ground to th inreasing iology wr thus ofl
influn of Roman Jsuits and th mahinations of th iuilt ttoli o- to stablishd ways o
tri. So whil ivart's fforts sapd at last for a tim th nsur of th authority and who h
Vatian' othr atholi volutionists did not far so favoraly' notaly th Thos who found v

gymn strngthnd his solv to
lum; ornll would an asylum
.ligiousdogma. It was
manuvrs
Lsto ronptualizth so-alld
torin soityas a soial struggl
_namly, th old-fashiond par-
st (urnr 1978). In this snario
al nd volution in pzrrtiularsim-
sls for ultural suprray wr
' , storis of th nountr -
s,vstmionflit or, for that mat-
ily to linar simpliity. In fat,
]ntWys to volution' and thir
taonomy of thologil orinta-
tl proupation.Bsids' volu-
a ultural vauum' and dats
rg voi to a rang of othr anxi-
ngagmntsmay glarrd from
n dividd.From th arly days th
lught togthr y Pop Pius I to
'rssd through th pags of L
:ry onsidralinflun among
nssityof maintaining th fiity
sformismwas vigorously promul-
tll 2001). By ontrast' th fforts
orgJakson Mivart, a atholi
r Lamarkian rading of transmu_
is of Spis was praisd in th
vat ould not a study soldir
Pius I awardd him th dgr
:lgiarrishopsprssd him to tak
1884, although 1rltr fll into
rritings.
llution undoutdly wr' th of-
tak any ation against atholi
s egan to hardn as th ging
i o n . I o s tg r o u n d t o t h i n
rtiorrsof te iuilt ttolicct o-
astfor a tim th nsu of th
ot far so favoraly, notaly th
t,ollttio d Rligio .].'.]
Frnh Dominian prist M.-D. Lroy, author of L'uolutio des esps
orgiques(1887), and th Amrian Jhn Zahm, who playd ir signifiant
rol in th dvlopmnt of th Univrsity lf Notr Dam in Indiana and pub-
|ised,uolutiod Dogm in 1896.
Both thswritrs suppotd th th;logialaptaility of volution' al-
it within rtain limits' Lroy's apologia, rtd foursquar on Mivart's
foundations,was dirtd against both thisti rnditiorrsof volution and
thologialnmisof th thory. But aush hintd that th human ody
might th produt of volutionary fors, th ardinal prft of th
SardCongrgation of th Indx of Prohiitd Books issud th judgmnt
..volution thory
that is tmrarious and anti-hristian whn pplid to th
human ody'' (quotd in Brundll 2001' 88). Although th finding that his
ook was nsurdWas onvyd to him privatly' Lroy was dvasttdand
spdilyrtratdin th pags of Le ond in arh 1895. Latr h trid to
find ways of rvisitingth whol issu y prduing a nw' orrtd vrsion
of th ook, ut it too fll foul of th uthoritis.
It was muh th sam with Zahm, who had alrady ahivd widsprad
rognition for his Sottd d si (1892) and tholi Scine d
tholi Scietists(1893). Although h had arlir ondmnd Darwin's De-
sntof as th aus of soial inquity, his purpos in writing uolutio
d Dogm was to dlonstratits thologial aptailiry.Indd h found
..to admir' muh
muh in it that is nnoling and inspiring,' (Zah-l,|896,
x). Likwis groundd in ivart's sstm' Zahm engagd in dtild patris-
ti xgsisto justify a tlologial rndring of volution. Th ook was an
intrnationalsuss'and an Italian translation rapidly appard,ut radrs
of a fuming iuilt ttoli wr assurd that it was all a tissu of lis' rdo-
lnt with rklss assrtions' and hoplssly ompromisd y duious as-
sumptions.Th ongrgation of the Ind dnouned tt in 7897, fousing
vry largly on t mttr of human origins. To traditionalists' Zalrm's vo-
lutionismwas part of a pakg of radil proposals tlrat inludd th adop-
tion of txtual ritiism and th ir-rrasingsgrgation of dmi and
lsiastialaffairs. Lik Lroy, Zam withdrw his fforts dspit th fa-
voral rviw h rivd from th nglish ishop John Hdly, who' for his
pains, prsntly am undr th whiplas of iuilt ttoli tongus.
.,Amrian-
Various fators playd thir parts in ths intrigus. A far of
ism''-th tndny on th part of Amrian Catholis to display intlltual
indpndnnd to adapt atholiism to arr Amrian OntYt-was on
suh fr. This indits that Darwinian ontrovrsisanlong turn-of-th-
nturyatholis wr oftn lss aout sinthan aout issus of idntity
with whih immigrant ommunitis wr grappling. Anothr suit of on-
rnsrotatd around th routin onflation of Drwinism and modrn shol-
arly trnds, notaly th nw ilial itiism. Rations to volutionary
s i n g iology Wr thus oftn all of a pi with rsponss to widr hallngs
to stablishdways of thinking. ritil too Was th mattr of intlltual
uthorityand who hd th right to intptth tradition rivd anon.
Thos who found vlution in th writings of Aquinas nd Augustin wr
54 t,oltio d Religio

arrogantly failing to pay du dfrn to th history of nosholasti lassil writings of phr

ommntary. Latr, frs of Darwinian-inspird ugni poliis playd their n d t h l i k o n t h o r i g i nt
rol in shaping atholi valuations of volution (Apply 1999\' sourd, as Was th lvork
Dspit ths prsistnt anitis, Huxly's fforts to prosri ivart's Rodrigo d Arriaga. Ths
atholi Darwinism' and th diffiultis that Lroy andZam prind at r o f s p i f i o n l u s i o n s
th hands of th ongrgation of th Ind, a sussionof tholi sholars ..hristitrrr
tur nd that
hav ontinud to dfnd th thologial proprity of Darwinian biology. th hurh fthrs on th
During th final dds of tlr nintnthntury, th atholi astronomr g rmindd his rdst}
Gorg Searl dlard that th thory of volutiorrnjoyd a strong fatual ondmnd volution as tl
sis and that human volution would not jopardiz atholi thology. At h rmained onvind tha
th sam tim th Harvard anatomist Thoms Dwight insistd that thr was ody-though not th sor
no ontradition btwn volutiorr and tlology. Th thologin John fully orrrpatilwith ath
Gminr addd his sllpport in 1884 whn, in oder Sitific Vi,s d It would mistakn tr
hristi Doctrilts ompred, h wnt so fa s to laim that Augustin v o i s . k , f o x n r 1 r l g
had atually prolotd a vrsion of volutionary thory and suggstdthat i i n i n B i r m i n g h a m .p r s i
Darwin ould argualy onsidrd his disipl. of anthropology in St. Nti
Prhaps th most onsidrd atholi statmnt n th sujt in tlr arly and orrspondnt of Tlr
dads of th twntith ntury appard in Frnh tn 1927 (and in nglish tion s ir tholi polog
transiation y rnst ssng th following ya) y th dirto of th Go- tholi of Sci (1
logial Institut at th Univrsity of Louvain, anon Hnri d Dorlodot, who taild srutiny in his work
dfndd not just th lgitimay ut th intrinsi dotrinal plausiility of Dar- .Writing
1917. at a tim WL
winism. Not only Wr thr no sriptural argumnts against th thory' h in- ovry of Mndl and th
sistd in Drtuiism d tholi Thought, ut th ..tahingof th Ftlrrsof proposals' Windl ws a
th hurh is vry fvoural to th thory of Asolut volution'' (Dorlodot atholi vrsion of o-La
|922,4)' ntral to Dorlodot's diagnosiswas his dvlopmntof th onpt Dspit ths fforts ant
of what h alld hristian naturalism. onivd as an antidot to athisti gtis from th Frnh Jst
and matrialisti volution' as wll as to th intrvntionist stan f figurs th mil-twntith ntr-rr
lik Gorgs uvir and Alid d'oigny, this onpt mphasized th imma- atholis sympathti to
nnt workings of th rator through ntural prosss.So mphati was Dor- ttd :rdvrsretioll
lodot in this judglrrntthat h pronound it lgitimat for Catholis to go vn C a t h o l i t h o l o g i a n sa s J
furthr than Darwin had don whn h attriutd th initial origirr of living larly' Tilhard d hardin
things to n at of spial divin intrvntion. atholi thology, h main- proval and was prlhiitd
taind, prdisposd its adhrnts to an advand systm of transformism and lss did lris work liit th
ven oligd thm to pt th ida tht all living bings wr drivd from t r M d a w a r ( 1 9 6 7 ) ,t h }
a fw lmntary organisrns' All of this was rpotdl in kping with tlr fully dismissd th Phlul
tahing of th hurh fathrs. St. Grgory of Nyssa's aount of origins, for A m o r r g t h o | o g i al r i t i t . st
..asolut
ampl, was paradd as a thory of volution'' that lft nO room s i s o f r h h u m n s p i s.
..vn
for spial intrvntion at th origin of lif'' (Dorlodot 1922,79)' onrns' whthr ovr pol
Givn ssng'srol as transltor of Dorlodot's volum' it is nor sur- to tholiism than ithr
prising that h himslf turnd in 1931 to a dtaildanalysis of th thologial sltion pr s bar'rstlr
proprity of volution that uilt on Dordolot's ..rilliant pi of wok'' tair-rdat all thologilo
(Mssngr 193 1, xxiv). Stagd in th introdutory ssay y harls Souvay papal ronounmnts hal
..noisy
as poisd twn th dvoats of Protstant odrnism'' nd th of th human soul y dir
..stuornranks'' of ProtstntFundamntalism,Mssngr'svolum undr-
onsidr Pius II's n
t<lok n tnsiv arhaologial twl through hurh history to vindiat thinkrs wr warnd of th
n volutionary aount of humn origins (ssngr 1931, xviii, ix)' Th .,in
tis' So whil rsah

to th history of nosholasti
pirdugnipoliis playd thir
[ution(Appl t999).
y's fforts to prosri Mivart's
rt Lroy andZam xprind at
, a sussionof Catholi sholars
proprity of Darwinian iology.
ntury' th atholi astronomr
:volution njoyd a strong fatual
. jopardizatholi thology. At
ras Dwight insistd that thr was
tlology. Th thologian John
'. in odr Sietifi Vittls d
so far as to laim that Augustin
'ionary thry and suggstdthat
isipl.
ltmnton th sujt in th arly
in Frnh in 1921(and in nglish
rg yar) y th dirtor of th Go-
n, anon Hnri d Dorlodot, who
insidotrinal plausiility of Dar-
rgumntsagainst th thory, h in-
..tahingof th Fathrs of
ut th
of Asolut volution'' (Dorlodot
las his dvlopmnt of th onpt
nivd as an antidot to athisti
h intrvntionist stan of figurs
hisonpt mphasizd th imma-
al prosss.So mphati was Dor-
: lgitimatfor atholis to go vn
triutd th initial origin of living
tion. athli thology, h main-
and systm of tansformism and
rll living ings wr drivd from
,as rportdly in keping with th
' of Nyssa's aount of origins, for
olut volution'' that lft no room
of lif''(Dorlodot7922,79).
D o r l o d o t ' sv o l u m , i t i s n o t s u r -
dtaildanalysisof th thologial
'dolot's ..brilliant pi of Work''
odutory ssy y Charls Suvay
f rotstant Modrnism'' and th
talism,Mssngr'svolum undr-
rough hurh history to vindiat
(ssngr1931, xviii, xix). Th
uoltiln d Rligio 55
lssialwritings of phrm, Basil, Grgory' hrysostom, Ambros, Aquinas,
and th lik on th origin of living ings,partiularly th human ra' Wr
sourd,as was th work of ltr Sholastislik Suarz,Ans Sylvius, and
Rodrigo d Arriag. Ths xavatlonsWr marshald in support of num-
r of spifionlusions-that spontanousgnrationWas taught in Srip-
tur and that ..hristian naturalism'' undrlay th prolamations of many of
th hurh fathrs on th passiv mod of divin ation. rtainly ssn-
gr rmindd his rdrs that standard manuls of atholi dogma routinly
ondmndvolution as thologially fals and philosophially asurd, ut
h rmaind onvind that th volution of plants, animals, and th human
body-though not th soul, whih rquird a dirt at of ratiln-was
fully ompatil with atholi orthodoy.
It would mistakn to think that Dorlodot and ssngr wr lon
vois.Tak, for xampl, th as of Sir Brtam Windl, FRS, dan of md-
iin in Birmingham, prsidntof Qun's ollg ork, and, latr, prfssor
of anthropology in St. Mihal's ollg' Toronto. A onvrt to Catholiism
and a orrspondntof Thomas Hardy, Windl surd a signifiant rputa-
tion as a Catholi apologist through his publiation of works |lke Tlue
tholi e of Si (|912). Not surprisingly, volution am undr d-
taild srutiny in his work T rh d Sic, whih first appard in
1917. \/ritingat a tim whn Darwinism was in lips busof th rdis-
ovryof Mndl and th availaility of various non-Darwinian volutionary
proposals,Windl was al to ploit to th full his inlinations toward a
atholi vrsion of no-Lamarkisr.
Dspit ths fforts and th sustaind sussionof volutionary apolo-
gtisfrom th Frnh Jsuit and palontologistTilhard d hardin during
th mid-twntith ntury that srvd as somthing lf a rallying point for
atholis sympathti to Darwin, atholi support for volution has at.
tratd advrs rations. Mssngr rportd th antagonism of suh
atholi thologians as Janssns,Pignataro, Hugon, and Van Noort. Simi.
larly, Tilhard d Chardin Was on th riving nd of pr*Vatian II disap-
proval and was prohiitd from spaking pulily on rtain sujts.No
lss did his work liit th iting nsur of a pofoundly unsympathti P-
tr Mdwar \7967), th Nol Priz*winning immunologist, who disdain-
fully dismissd the Phlmo|1of as a work of philosophial fition.
Among thologialritis dat has routinly ongrgatdaround th gn-
sis of th human spis,a proupation that onfirms that anthropologial
onrns'whthr rvrpolygnism or human anstry' Wr mor unnrving
to atholiism than ithr uniformitarian gology or th prinipl of natural
sltionpr s austh onsanguirrityof th human ra had to r-
taind t all thologil osts (Astor 1996). Thus various twntith-ntury
papal pronounmntshav n onrnd to prsrvth spial ration
of th human soul y dirt divin intrvntion.
onsidr Pius II's nylil Humi cris (1950). Hr atholi
thinkrswr warnd of th dangr of too agrlymraing sintifinovl-
..intl
tis.So whil rsarh th origin of th human ody as oming fom
56 uoltion d Religio

pr-istntand living mattr'' was allowd' anv suggstionthat th humn th imag of a divin .
ra Was f plural origins was asolutly ond-rndausof its implia- ontraSt' Samul Wil
tions for th dotrin of original sin. A similar stipulation prtaind to th ally and philosophial
origin f th lruman sor.rl,whilr Pius insistdwas yond tlr rh of vlr'r- ].860 mting f th
..Th Franis Opn ois,
tionary transformisnr: atholi faith oligs us to hold that souls ar
immdiatly ratd y God'' (Pop Pius II 1950' [36). shir, who authord
or rntly Johrr Pul II addrssda plnry irssmlyof th Porrtifial Drinism (1869) and
Aadmy of Sinsin Otor 1,996r>nth sujt of volution and its sig. thought tht Darwinisr
nifian for undrstandingth human agnt. Ralling Pius II's nylial th Atlanti th lrgy
and noting that evolution had now gon yond th status of a mr hvpoth- gllags in Boston Tho
sis, h ondd th liklihood that th humn ody had originatd fronr hoi napsulatdin
prxistnt living mttr, ut h still rstatd th tditilnl rationist- ntion too should bt
as opposd to trduinist-dotrin of rlr rigin of th human soul irr ordr doyn of Amrian Prs
..th ontologial lap'' twn humanity and its Drtuiism?(l874) wit
to sustain what h alld
forars.By this mov mtaphysialmtrrs rotilting around morality, on. a judgmnt on Darwin'
sin,frdom' asthtippriation, and spiritual prinwr dl- ltion itslf.
gtd to phiiosopial anlysis rathr than to volutionary iologv aus Th swp of opinior
..momnt Th nglish Nononft
t of transition to th spiritual'' was simply not opn to mpirial
..osrvation''(Pop Thomas R. Birks, tlok
Joanns Pulus II 1996, II 6).
Not surprisingly, this statmnt has attrt]th ttntion of atholi Spnian form, in bo
ommntators. on th mor rationary sid, John Marthy osrvdthat uolutio pulishd in
th addrss had gnratd dismay and onfusion among th faithful. H ity's ultimat hoi wa
thrfr workd hrd to minimiz any opnnssto volution that th po- Seltio' His fllorv ot
nounmnt might foster y th introdution of a numr of srplrtinin- ously sought to Drwtn
trprtations and y rminding his radrs that mmrs of th Pontifial ( l88). Lss spulat
Aadmy of Sinswr hosn rgardlssof ligious rd. By ontrast' who tok up th ps
in 1998 th Vatian astronomr Gorg V. oyn Warnlly mrird th Univrsity) in 1868, t<
pope's announmnt of evolution's fatual status and wnt on to urg that Allian that instadof
..ontinuous ophrs would ttr
sin volution ws tlr mthod of divirr rtion' th ida of
ration'' Was th st way to oniv of th mrg of th human fllow Sotsmn, th th
spis (oyn 1998, |60). ah, and Hnry ldr
of thisti vlution. I
insistd in th first d:
Th Protstantosai lution was simply anot
This pathwork of l
Protstarrtrsponssto volution Wf no lss divrs.From th arlistdays around 1900 is math
thr wr thos who snsd in volution th mthod of God's modus Wer onntd with vl
oprandi in natur. Anglian pi1rson'novlist' and hristian soilist harls own mntor, th Angli
Kingsly, for arrrpl'Was nthusistiabout the rigi irnd wrot to Da- p r s s d a r l y o p p o s i t i
win tlling him that although th thory mant that h must giv up many of wik nd th Sotti
..just mathmatial physiis
th things h livd,it was as notal a onption of Dity, to -
liv tht H ratd priml forms apal of slf-dvlopmnt''(Brrrkhardt physial omputations
and Smith 1985_2005, 7: 380). Simllarly, Frdrik Tmpl, latr to om tim ndd for th o
arhishop of Cntrury, pssdhis support as arly as 1860, ontnd- And infamously th Pl1
ing that a world md to mak itsl{attrllhirndth rator's noilit. rspondnt of Darwiir
To suh figurs, a univrs govnd y lratural law was nror dignifying tha volution and rlid o

ld' any suggstionthat th human
:ondmndausof its implia-
milr stipulation prtaind to th
d was yond th rah of volu-
L oligsus to hold that souls ar
: I I1 9 5 0 ' I3 6 ) .
plnry assmlyof th Pontifial
h sujtlf volutin and its sig.
nt.Ralling Pius II's nylial
:yond th status of a mr hypoth-
human ody had originatd from
atd th traditionlrationist-
: origin of th human soul in ordr
l iap'' twnhumanity and its
rs rotating around morality, on-
nd spiitual prinwr dl-
n to volutionary iology aus
' Th nglish Nononformist and ofoundr of th vanglial Allian,
was simply not opn to mpiril
)6,\ 6).
:tratdth attntion of tholi
id, John arthy osrvd that
:onfusion among th faithful. H
pnnssto volution that th pro-
. i o n o f a n u m o f s r p n t i n i n -
rs that mmrs of th Pontifial
ss of rligious rd. By ontrast,
V. oyn warmly mrd th
al status and wnt on to urg that
..ontinuous
ration,th ida of
of th mrgn of th human
lssdivrs.From th arlist days
ion th mthod of God's modus
list' and hristian soialist harls
out th rigi and wrot to Dar-
lant that h must giv up many of
lal a onption of Dity, to b.
of slf-dvlopmnt''(Burkhardt
Frdrik Tmpl, latr to om
supportas arly as 1860, ontnd-
lly nhandth Crator's noility.
tural lw Was mor dignifying than
uolutio d Religion 57
th imag of divin onjurr who mgikd nw spis into xistn. By
.!7ilrfor
ontrast, Samul thought that Darwin's thory was oth mpiri-
ally and philosophially unsound and famously pourd sorn on it at th
1860 mting of th British Assoiation for th Advanmnt of Sin.
Franis Orpn orris' Vitorian ornithologist and Anglian rtor in York-
shir' who authord a squn of pamphlts with titls lik Diffiulties of
Dariism (1'869) and The Demnds of Dr,iism o redulit Q890),
thought that Darwinism dsrvd only uttr ontmpt and drision. Aross
th Atlanti th lrgyman Luthr Tray Townsnd, who taught ilial lan-
guagsin Boston Thologial Sminary, plad for his radrs th stark
hoi napsulatd in th titl of his trat F,uolutio or retio (1896).
ntion too should mad of th Printon thologian Charls Hodg,
doyn of Amrian Prsytrins, who answrd his own qustion in.Wht ls
Drtuiism? (1874| with th trs quip ..It is athism''' though this was mor
a judgmnt on Darwin's antitlologial stan than on th prinipl of vo-
lution itslf.
Th swp of opinion rprsntd hr ould rpatd many tims ovr.
Thomas R. Birks, took up th udgls against volution, partiularly in its
Spnrianform, in ooks lik odr Phsil Ftlism d tbe Doctrin of
uolutio pulishd in \876. Th Sotsman.lfilliam Millr flt that human-
ity'sultimat hoi was' to us th titl of his 1897 volum' God, or Nturl
Seltio.His fllow ountryman Hnry Drummond' on th othr hand, vigor-
ously sought to Darwiniz thology inhis Nturl L, i the Spiritul World
(1883). Lss spulativ and mor traditionally alvinist, Jams osh,
who took up th prsidny of th ollg of Nw Jrsy (latr Printon
Univrsity) in 1868, told th 1873 Nw York mting of th vanglial
Allian that instad of dnouning the thory of volution, hristian philos-
ophrswould ttrmployd in pounding its rligious dimnsions. His
fllow Sotsmn' th thologians Rort Rainy, Gorg athson, Jams lvr-
ah, and Hnry aldrwood, all addd thir vois in support of som form
of thisti volution. Latr th Amrian Baptist thologian A. . Strong
insistdin th first dad of th twntith entury that th prinipl of vo-
lution was simply anothr nam for hrist.
This pathwork of Protstant opinion from th thologians in th dads
around 1900 is mathd y th pattrn of rspons mong sintists who
Wronntd with various Protstant dnominations. In Britain Darwin's
own mntor, th Anglian lrgyman.naturalistJohn Stvns nslow, x-
prssdarly opposition' as did th amridg gologist Rv. Adam Sdg-
wik and th Sottish natural philosophr David Brwstr. Th Irish
mathematialphysiist at Glasgow, \/illiamThomson (Lord Klvin), whos
physial omputations of th ag of th arth smd to dny Darwin th
tim ndd for the oprations of natural sltion, rmaind antagonisti.
And infamously th Plymouth Brthrn naturalist Philip Hnry Goss, a o-
rspondntof Darwin and admirr of l/alla'was implaaly opposd to
volutionand rlid on th ida h put forward tn mphIos (1857) (th
3.'8 uolutil d Rligil

Grk word f<>r ul) tht God had ratd a matur univrs with th sm- infiltration. To a on
lan of ag-prohronism, as h lld it. Aross th Atl:rnti th Swiss g- sin an trad
ologist and physial gographr at rsytriarrrintorr, Arnold Guyot, tist Gorg Mrd
and his anadian ollntrpart Jolrn Villiam Dawson, prsidnt of Gill Wekst Poit i the
Univrsity, also st thir fas against Darwinism. In th mid 1880s Guyot His flood gology
advoatd a kind of pr-Darwinian lvlopmntalismthat, in th fashion of apologist, lln \/hit
th Sottish stonmason Hugh illr, rad th gologial pohs into th litralist intrprtatio
dys of Gnsis, whil Dawson alkd at Darwinis]'s antitlologial alon, and h riv
stan. By ontrast th Llarvard otanist, Congrgationalist, and onfidant slf-styld rsarh s
of Darwin Asa Gray foLrnd oth odg's and Dawson's ojtions odd tor.William Bll Ril
and workd hard to fit volution into tlologial garmnts y arguing tht was also forthomin1
it was rnistakn t rstrit th id of dsign to th suddn flashing of a rati andidat for
Palyit wth into istn.Anothr ongrgationlist' th Orlin golo- paifist, whos fars
gist Gorg Frdrik Wright, also gav volution his ndorsmnt in th Grmani militarism
arly 1880s' vn if h wavrd ltr in lif, i.rndwnt so far as to draw a st th lrtd Sops
of rlling analogis twnCalvinism and Darwinism, onluding that Dar- ron traordinair or
winism was ..the alvinisti intrprtation of natur'' (\/right1882, 255). Darrow. Of ourse, tt
othr supplrtiv ommntaris. though with varying dgrs of nthusi- absnt from ths th
asl' wr fothoming from suh sintifi pratitionrs as th alifornia phlt titld Poisoig
gologist Josph Lont, th Yal minralogist Jams Dana' th Mihigan volutionism togth
.sinhll,
goIogistAlandr and th Printon lrgyman.iologistGorg volutionary transfor
aloski' Indd, figurs lik ths did a good dal to kp lins of om- and politial.In Brita
muniation opn btwn spialist sin and radrs of dnominational ship of Amros Flr
magzins. vrsity Collg Lond
A a t a l o g o f s u h p a r t i u l a r i t i s o u l d l a o r a t da d l i i t u m , u t a s a n allowd for th poss
ovrall grrrlization' most Protstant thinkrs livd that volution Indd, in Flming's t
ould mrad without aandoning dotrinal ssntials.Th majority ration of th ili
found ways of aommodating thir thology to mor or lss rvisd vr- al findings2 a mov t
siorrs of volutionary thory. As wll as hallnging widsprad assump- wrong way.
tions of systllihostilitis twrtvolutiol-land rligion, this trjtory Whatvr th sour
srvs to all attntion to a numr of sming inongruitis. Thus num- motion dlind fair
rd among th writrs of Th Fudmetls, a squneof 72 pamphlts though it promptd
i s s u d t w n 1 9 1 0 n d 1 9 1 5 t h a t g a v v o i t o r l . vF u r - r d a m n t a l i s m , Histor of No-eu
Wr svral writrs, inluding G. F. Wright and th Sottish thologian tnt and up.to-dt lil
Jams Orr, who advoatd volution in on form or anothr. There is also ayan until th 19.
.Warfild,
th s of th Printon thologian B. B. ustodian of th onfs- iblial thlogian Jo
sioral standards of trditional lvinism and rnodrn ahitt of th do- am th foundtio
trin of bilial inrany, a mainstay of th Amrian Fundmntalist Geesis Flood. |t |im
.!arfild
movmnt. Brought up on a Kntuky stok farm' knw firsthnd nd rassrtdrati
aout inhitd variation irr animal rding arrd latr musd that h was its various dtion
lrady a ..Darwinian of th purst watr'' bfor th oming of Jams lswhr in this volr
osh to Printon (\Wrfild \976, 652). In 1915 h limd that polmis and intrn
alvin's dotrin of rtion was in fat a ,.vy pur volutionary shm'' day rationism has d
( W a r f i l d1 9 1 5 , 2 0 9 \ . ally' In fat, th th
Dspit this plurlism aross th dnominational sptrum' rtain s- ontinud to dfn
tarian strands in Protstantism rraind staunhly rsistant to Darwinian dnominational rang

l a matur univrs with th sm-
, Aross th Atlanti th Swiss g_
trin Prineton, Arnold Guyot,
Lm Dawson, prsidrrtof MGill
winism. In th nrid 1880s Guyot
lmntalismthat, in th fashion of
d th gologial pohs into th
at Darwinism's antitlologial
ongrgationalist,and onfidant
's and Dawson's oitions odd
llogial garmnts y arguing that
sign to th srrddn flaslring of a
:lgrgationlist, th ()rlin golo-
:volutionhis ndorsmnt in th
, and wnt so far s to dfaw a st
Darwinism, onluding that Dar-
t o f n a t u r ' '( w r i g h t 1 8 8 2 ' 2 5 5 ) .
with varying dgs of nthusi-
i pratitionrsas th alifornia
LlogistJams Dna' th ihigan
ton lrgyrnan-iolgistGorg
' good dal to kp lins of om-
: and radrs of dnminational
laoratdad liitum' but as an
hinkrs livd that volution
l o t r i n l s s n t i a l sT. h m j o r i t y
logy to mor or lss rvisd vr-
hallnging widsprd assum-
r t i o n a n d l i g i o n ,t h i s t r j t o y
minginongruitis. Thus num.
tls,a squnof 12 pamphlts
: voi to arly Fundamntalism,
ight and th Sottish thologian
n form or anothr. Thr is also
Warfild,ustodian of th onfs-
and modrn arhitt of th do-
lf th Arriirr-r}-undamntalist
k farm, Warfild knw firsthand
ling and latr rusd that h was
r'' for th oming of Jams
652). In 1915 h laimd that
..vrypur volutionary shm''
rlinationaI sptrllm' rtain s-
staunh[,vrsistant to Darwinian
uolutio d Rligio 59
infiltration. T a onsidral dgr th mrgn of modfn ration
sinan trd to th nrgti ativitis of th Svnth-Dy Advn-
tist Gorg r:rdy Pri, author in ] 906 of Illogil Geolog: Tb
Wkst Point in the uoltio Theor and irr 192 of Th N Golog'
His flood glogy prssd th -lr4osaisin of th movmnt's hif
apologist, lln Vhit, whos dotrin of th Saath was groundd in a
litralist intrprtatin of th ration narrativ. Pri, of ours' ws not
alon, and h rivd noriragmntfrom th Prsyterian ministr and
slf-styldrsarh sintist Harry Rimmr and th ]V1innsotaBaptist ps-
.William
tor Bell Rily. Support, though from a diffrnt idologial stal,
was also forthoming from William Jnnings Bryn' thr tims Dmo-
rati andidat for th prsidny of th Unitd Stats and ommittd
paifist' whos frs about th sming allian twn Darwinism and
Grmni militarism nouragd him to tirk up th proseution's s in
th lratdSops trial of |925.|t arnd for him th rputation of mo-
ron traordinair ourtsy of th polmis of his fisty opponnt' larn
Darrow. of ours, this is not to say that mattrs of politial idology wre
asntfrom ths othr ontrovrsialists.Pri, for instan, issud a pam-
phlt titld Poisoig Demor 1921) in whih h nttd soialism and
volutionism togth as th mainspring of modrn-day ills. His loathing of
volutionary trnsfortnisnr was thtls mr-rlridirrrrrsional-bilial,sintifi,
and politial. In Britain th volution Protst ovmnt' undr th ladr-
ship of Amros Flming, FRS, profssor of ltrial thnology at Uni-
vrsity ollg Lndon, lakd th vitriol of its Amrian ountrpart and
allowd for th possiility of som divinly guidd volutionary hang.
Indd,in Flming's own as h was vn prpard to rd aspts of th
rtion of th ilial Adam in th light of various paloanthropologi-
al findings, a mov that ertainly rued th anatomist Arthur Kith th
wong Way.
!hatvr th sours of ths sntimnts' this flurry of rationist om-
motion dlind fairly dramatially in th yars aftr th Sops trial,
though it prornptd th puliation of . T. Brwster's 1,927 rtio:
Histor of No-uoltior Thoris to alan th ..ampl and omp.
tnt and up-to-dat litratur of volution.'' rationist agitation Wnt into
abyanuntil th 1950s whn th hydrauli ngineernry Morris and th
bilial thologin John Whitom joind fors to produ in 196l what
amth foundational txt of th modrn ration sinmovmnt' l2e
Gensis Flood. It limitd goiogial histoy to no mor thn 10,000 ys
and rassrtdration in six litral dvs. Th taitory of this movmnt'
its various duational mahinations, nd its lgal Stratagmsar hartd
lswhrin this volum. It suffis hr to add that dspit its ggrssiv
polmisand intrnational suss' it would mistakn to think that si-
day rationismhas dominatd onsrvativProtstant thought mor gnr-
ally. In fat' th thory of volution, in on vrsion or nothr, has
ontinudto dfndd y Protstant thologins and sintistsaross th
dnominationlrarrg.
60 uoltio nd Reliil

for rligious institutions a

In th Light of norah and Qur'an
ats ws th ntur of Jud
Somthing of th sm s
Rligious rsponss to volutionary thory wr oftn ound up with roadr
though in th Islami as
issus that onfrontd thologial ulturs and faith ommunitis, not last
th Qur'n has mant that
qstions of how to ngag with a thratning wider soity or how to rtin
lss, Islami support for v
or rfashion idntity in the fa of xtramural hallngs.Jwish datsaout
for xampl, has laimd t
Dawinism in lat nintnth-ntury Amria ar illustrativ, for ths wr
Islam, whil th Laness
part and parl of a suit of dlirations on th futur shp that Amrian
tirly ompatil with Qur'
Judaism should tak (Swtlitz 1,999).Traditionalist, odrat Rform, and
fzaa| (7996)' who likwis
Radial Rform rabis ah had a stak in how th Jwish ommunity should
from th Holy Qur'an itslf
rspond. on th radial wing volution was mrad ausit ould mo-
winism and volution. The
ilizd to undrwrit progrssivism and tansformation. Thus th radrs of
limitd appliaility of nat
Kaufmann Kohlr's various pronounmnts during th 1870s and 1880s ..fits prftly in th ovrall
larnd that volution applid to th spiritual ralm and that Jwish ritual
Book.,' Indd, dploying tl
and rmonial ndd to rfashiond to kp in stp with modrn sin.
an volutionary aount of
To him and othrs lik him, volutionary dvlopmnt onfirmd th prinipl
mnt to natural prosss.
of progrssiv rvltion; adaptation to th prvailing nvironmnt applid
ration. In this mov lnr(
to organi and rligious ommunitis alik. From th sam thologial stal
rtus' and so on-dvlo:
Josph Krauskopf, in a squn of lturs ttt|eduoltio nd Judism and
ltd a singl pair-a mal
pulishd in 1887, radily applid volution to th growth of rligion, th
itual souls." Not surprising
txtual history of sriptur' primitiv Soity' th ida of God, and th dvl-
aounts of figurs lik Til]
opmnt of Jwish worship.
Dspit ths irni ff
Traditionalists, y ontrast' ithr rjtd Darwinism or ln som ass
from writrs lik T. . Jan:
moilizd it to support traditional Jwish thology. on th on hand, Samul
warrant and is promotd
. Isaas usd his position as ditor of th letuish sseger to launh at.
hauddin Nadvi issud a rl
taks on volution, whil Araham d Sola' a anadian immigrant, mr-
ration of Adam, whil A
shaled th ritiisms of th Montral gologist John William Dawson to
i n o r p o r a t dm t t ro r i g i n
attak th thory. Thomas Mithll, an advoat of th viw that God had
haps th most influntialIs
ratd th world in si 24-hour days, told readrs of his 1887 artil..vo.
dly an minnt Turkish r
lution and Judaism'' for eorb that thr was not a shrd of vidnfor
antivolutionary trats (lik
human antiquity or volutionary hang. On th othr hand, the traditional-
that his opponnts laim tl
ist ditors of te Amri Hebr' a good numr of whom wr traind in
of writrs. In any as,in t}
sin or mdiin, workd to ultivat a thisti vrsion of volution along ..dishonstphilosoph
at th
th lins of Asa Gray. Traditionalists too appald to th gradualism of vo-
si valus on whih th sta
lution to oppos rapid hang and any swping ovrthrow of inhritd insti-
th trat urgs, providd a l
tutions. Th tnaious nduran ovr ountlss gnrations of traditinal
ntifi status' thry xpl
rmonial tstifid to its valu in th Jwish struggl for survival against
in natural history. In anot
i n n u m r l a d v r s i t i s .
rought to humnity,'' YaI
Intrnal rivalris thus did muh to govrn raini nountrs with volu-
sar to th malign influn
tion vn though pro- and anti-Darwinian sntimnts nvr dirtly mappd
winism on figurs lik Stal
onto th ommunity's vrious fations. This mans that the sintifi dimn-
(Yahya 2001). Translatd
sions of volutionary thory wr rarly addrssd;rsponss wr mdiatd
Yahya mind-st' partiular
through onrns aout th supposd influn that matrialism had on low-
r a t i o n i s t o o k s o m i n
ring synagogu attndan' on volution's impliations for th natur of
tain plas (Konig 2001).
mind and morality, and on what advoats livd its aptan ntaild
numb of TukishDewi
 uolution nd Religio 61

for rligius institutions and ustoms. \/hat was oftn at stak in thes d-
n
batswas th natur of Judaism itslf.
Somthing of th sam sptrum of rations is also disrnil in Islam,
r oftn ound up with roadr
though in th Islami as a rathr mor prvailing litralisti hrmnuti of
rnd faith ommunitis, not last
th Qur'n has mant tht advoats of volution ar lss ommon. Nvrth-
r g w i d r s o i t yo r h o w t o r t a i n
lss'Islami support for volution is not ntirly asnt. uhammad Iqal,
hallngs.Jwish dbats aout
for ampl, has laimd that th ida of volution originatd in mdival
a ar illustrativ, for ths wr
Islam, whil th Lans sholar ossin al-Jisr insists that th thory is n-
r th futur shap that Amrian
tirely ompatil with Qur'ani thology (Majid 2002).In th as of Ahmd
'ionalist,Modrat Rform, and
tfzaa| (7996), who likwis ontnds that ..th thory of volution sprouts
lw th Jwish ommunity should
from th Holy Qur'an itslf,'' a ruial distintion is prosutd twn Dar-
:mrdbausit ould mo-
winism and volution. Th formr h finds unaptal on aount of th
lsformation.Thus th radrs of
limitd appliability of ntural sltion, whras th prinipl of volution
ts during th 1870s and 1880s ..fitsprfetly in
th ovrall shm of God's ration as dsid in th Holy
'lal ralm and that Jwish ritual
Book.'' Indd' dploying th standard soul-ody dualism, fzaa| argus for
<pin stp with modrn sin.
an volutionary aount of th human spis that alloats physial dvlop-
lopmntonfirmd th prinipl
mnt to naturl prosss and th mrgn of slf.onsiousnss to divin
prvilingnvironmnt applid
ration. In this mov inrasingly humanoid forms-Homo hbilis, Homo
From th sam thologial stl
rtus,and so on-dvlopd to a point whr ..Almighty Allah (S!) s-
rit|eduolution d Judism and
ltda singl pair-a mal and a fmal-and ndowd thm with thir spir-
)n to th growth of rligion, th
itual souls.'' Not surprisingl, fzaa| found inspiration in th antimhanisti
l, th ida of God, and th dvl-
aountsof figurs lik Tilhard d hardin and Hnri Brgson.
Dspit ths irni fforts, vigorous opposition has n forthoming
d Darwinism or in som ass
from writrs lik T. H. Janai who laim that volution is without mpirial
ology.on th on hand, Samul
warrant and is promotd for soial and politial rasons. Similarly, Sha-
: Jetuish ssngr to launh at-
habuddin Nadvi issud rfutation of volution in his 1987 aount of th
La,a anadian immigrant' mar.
ration of Adm, whil Adm Tatli's uolutio: Bkrupt Theor 0990)
logist John !illiam Dawson to
inorporatd mattr originally prsntd to th Turkish govrnmnt. But pr-
,,oatof th viw that God had
..vo- haps th most influential Islami riti of Darwinism is Harun Yahya, rport-
radrsof his 1887 artil
dly an minnt Turkish sholar, who has n so prolifi in produing
3 was not a shrd f vidn for
antivolutionarytrats (\tkeWb Drluiism ls Inomptible ith th Qur')
n th othr hand' th traditional-
that his opponnts laim that th works ar atually th produt of a group
numrof whom wr traind in
of writrs. In any as, in te \999 ook Th uolutio Deit Yahya hit out
histi vrsion of volution along
at th ..dishonstphilosophy'' of ..matrialism''that ..sksto aolish th a.
>pald to th gradualism of vo-
si vlus on whih th stat and soity rst'' (Yahya 1999, 7). Dawinism,
ping ovrthrow of inhritd insti-
th trat urgs, providd a mythi foundation for matrialism's laims to si-
rntlssgnrations of traditional
ntifi status, thry plaining why arx rootd dialtil matrialism
.ish strugglfor survival against
in natural history. In anothr 2001 rharsal of ..th disastrs Darwinism
rought to humnity,'' Yahya has attriutd war, povrty' pain, and mas-
r rabini nountrs with volu-
sarto th malign influn of ..th selfish and pitilss world viw'' of Dar-
;ntimntsnvr dirtly mappd
winism on figurs lik Stalin' Trotsky, ao, Pol Pot, itlr' and Mussolini
s mans that th sintifi dimn-
(Yahya 2001). Translatd into mny languags' th output of th Harun
Jrssd;rsponsswr mdiatd
,nthat matrialism had on low- Yahya mind-st, partiularly in Turky, has n rportd as immns, with
's rationistooks oming mor influntial than standd ttooks in r-
impliatins for th natur of
tain plas (Konig 2001). To sur, ths movs hav n rsistd y a
s livd its aDtan ntaild
numbr of Turkish Darwinians, notaly th iologist Aykut Kn and th
62 uolution nd Rligion

mdil gntiist Isik Bksoy. But marshaling support for Darwinian vo- divin wisdom and powr
lution has provd diffiult in a ontt whr politial frs hav fostrd wrot inquiry int th ni
th tahing of rationism in shools and whr dirt support has n pro- vrs. In rspons' num
vidd y Amrian Protstant rationists(dis 1994). tlologially. In gnral,
sion had a mor diffiult
plans and th oprations
Ruring Thms thusiast for th way Argyl
divin wisdom mor in h.
Various historial modls hav n anvassd in th attmpt to gt a handl Th Duth alvinist, fun
on th rlations twn sin and rligion. Our travls thus far onfirm that o f - t h - n t u r yp i m m i n
any simpl portrayal of th rlations twnvolutionary thory and rligious zoologist Jan Lvr (195
lif is doomd to fall foul of th mssinssof history, ut a numr of prsis- whih onivd of natur
tnt thms hav rurrd, alit in diffrnt wys in diffrnt sttings. ours' this dos not ma
Qustions of sriptural hermeeutis hav routinly surfad in disussions tirly disappard. Th nt
of volution. Som adhr to a litralisti xegsis of ration narrativs, surfad in rnt tims st
whil thos who ar intrstd in onordism hav strssd th rol of mta- has n alld irrduil
phor and ultural ontxt in th intrprtation of partiular statmnts. an all-or-nothing varity l
Thos who ar willing to rlgat th osi narrativ to th ralm of prim- ations on whih natural s
itiv mythology hav n ngagd no lss in a hrmnuti undrtaking. i n i t s s t t i s t i a lw i z a r d r y '
ithr Way' qustions of gsis hav prsistntly otudd. Philip Hnry pliations of Darwin fr .
Goss's whol systm of prohronism was dsignd to prsrv th litral a- thr ontinus to b phil
uray of th Bil's timsal.By ontrast, th Orintal and ilial sholar th diffrnt positions ad<
\. H. Grn rlivd rtain strands of nsrvativ rotstantism of th tinga (199 l),who dfnds
urdn of Usshr-typ hronology (whih famously datd th world's origin rnan MMullin (1991I,
to 4004 B...) whn h dmonstratdin 1890 that old Tstamnt gnalo- ural law. Th fat that
gis ould not rlid on to dat primval tim on aount of thir stratgi s o u n d i n g v o a b u l a r yu r g i
omissions from th rords. That gav som thisti volutionists th y . t r i l a n d o . ' h ti t
.loo1
hrmnuti room thy ndd. Again, th atholi writr Dorlodot dvotd y snaking through
onsidral spa to luidating atholi thoris of inspiration in ordr to t i n s i n o r d t o i m p r o v i
allow for his onvition that th thought-forms of siptur wr aommo- whol issu (Grne 1989
datd to th ultural onditions of th tim. And th Islami volutionist Aross th rligious ran
Ahmd fzaa| urgd th lgitimay of xplaining ..th rlvant Qur'ani th sptr of mterilisn
vrss in trms of a mtaphor or paral'' and spok of ..th Qur'ani lgend that volution is rtd
of th .Fall''' (t{zaa| 1996). In on Way or anothr' volution thory has moral natr and has u
prsistntly ralld attntion to hrmnuti mattrs in th intrprtation of various strnds hav also l
anint sard tts. for instan, th Catholi
From its arlist days Darwin's thory onfrontd had-on th issu of tl' A Stud of Vitlism d
olog. As Darwin himslf put it at on point, ..Th old rgumnt from dsign outlook of th primn
in ntur, as givn by Paly whih frmrly smdto m so onlusiv' fails, th antirdutionism pr
now that th law of natural seltionhas n disovrd'' (Darwin 1888, 1: and P. G. Tait, whih first
309). Th onsqunsWr potntially monumntal, for natural thology John yndall's infamous -
had undrsord th fundamntally moral haratr of natur' nthusing - linations, Windl Was at
livrs, uring starian tnsions' end inspiring mn and womn to study mattr' and h rmaind t
.!0.hig, hanial univrs. By on
God's handiwork. !hthr from th pn of \/illiam Paly, a modrat
whos Nturl Thology (1802) was dvourd y Darwin, or in th multi- sondary uss-that G
volum Bridgeter Trtises (1833_1836), whih st out to dmonstrat f o r m o f m a t r i l i s m o n s i

ling support for Darwinran vo.
r politial fors hav fostrd
rrdirtsupport has n pro-
dts 1994).
:d in th attmpt to gt a handl
our travlsthus far onfirm that
volutionarythory and rligious
lf history' ut a numr of prsis-
ways in diffrnt sttings.
r o u r i n l ys u r f a di n d i s u s s i o n s
gsisof ration narativS'
n hav strssdth rol f mta-
tation of partiular statmnts.
narrativ to th ralm of prim.
; in a hrmnuti undrtaking.
sistntly otrudd. Philip nry
signdto prsrv th litral a-
th orintal and ilial sholar
lnsrvativrotstantism of th
rmously datd th world's origin
]90 that old Tstamnt gnalo-
r i m o n a o u n to f t h i r s t r a t g i
som thisti volutionists th
]atholi writr Dorlodot dvotd
horisof inspiration in ordr to
lrms of sriptur Wr aommo-
n. And th Islami volutionist
..th rlvant
'plaining
..th
nd spok of Qur'ani lgnd
)r anothr' volution thory has
i mttrs in th intrprtation of
rfrontd had-on th issu of tel-
..Th old argumnt from dsign
t,
smdto m so onlusiv' fails,
: nd i s o v r d '(' D a r w i n 1 8 8 8 , 1 :
lonumntal, for natural thology
:haatrof natur, nthusing b-
piring mn and womn to study
!illiam Ply, a modrat.Whig,
urd y Darwin, or in th multi.
i). whih st out to dmonstrat
uolutio nd Religil 6
divin wisdom and powr in ration, th natural thology ntrprisundr-
wrot inquiry into th natural ordr. Darwin dstailizd this whol uni-
vrse. In rspons' numrous writrs workd hard to intrprt volution
tlologially.In gnral,thos who wr still nthusiastiaout Paly's vr-
sion had a mor diffiult task than thos who found dsign in arhtypal
plans and th oprations of natural law. Thus Jams osh, alrdy an n-
thusiastfor th way Argyll's b Reig of I-tu rloatd divin dsign, found
divin wisdom mor in homologial struturs than in spifi adpttions.
Th Duth alvinist' foundr of th Fr Univrsity of Amstrdam, and turn-
of-th-ntury prim ministr Arham Kuypr nd latr dvots lik th
zologist Jan Lvr (1958, 229) spoke of ..divin volutionisti ration'''
whih onivd of natur unfolding aording to a prdtrmindplan. of
ours'this dos not man that advoats of Paly's mhanial vrsion n-
tirly disappard. Th nortionist Intllignt Dsign movmnt that has
surfadin rnt tims sks to idntify trnally drivd purpos in what
has n alld irduilomplity, namly, rtain iologial systmsof
an all-or-nothingvarity whose funtions annot rdud to simplr opr-
ations on whih natural sltion an work. In its nptual strutur, if not
in its statistialrwizardr,this rads lik Paly rdivivus. In ny as th im-
pliationsof Darwin for dsign hav maind a ky arna of date. That
throntinusto philosophial milag in this whol mattr is lar from
th diffrntpositions adoptd y suh minnt philosophrs as Alvin Plan.
tinga(1991),whodfndsdivine intrvntionismin th history of natur, and
rnan Mullin (1991)' who inlins mo toward God ating through nat-
ural law. Th fat that iologists hav frquntly tradd in tlologial.
soundingvoaulary urging ..that volution is .opportunisti', that it prods
y .trialand rror', that it rahs.dad nds' or aomplishs.rakthroughs'
by snakingthrough .loophols', or that natural sltion .an rmodl pro-
tinsin ordr to improv intrations' '' adds yt furthr ompliations to th
w h o l i s s u ( G r n 1 9 8 9 ' 4 0 8 ) .
Aross th rligious rang opponnts of Darwinism hav n hauntd y
th sptr of mterilisz. Islami opposition springs in part from a sns
Qur'ani
that volution is rtdon a matrialist rd that donstrutshumanity's
.Wstrn
moral natur and has ultivatd a dadnt ultur. hristians of
variousstrandshav also n trould y similar onrns.Brtram.!indl,
for instan' th Catholi advot of volution and author of Wbt ls Life?
Stud of Vitlism d Neo-uitlism (\908), prssd sympthy for th
outlook of th primntal physiist and spiritualist Olivr Lodg and for
th antirdutionism prssd tn The Usee Uiuerse of Blfou Stwart
and P. G. Tait, whih first appard anonymously in 1875 immdiatly aftr
John Tyndall's infamous Blfst addrss. An volutionist of Lamarkian in-
linations,Windl was attratd to th ida of a Vital Powr that anrmats
matter'and h rmaind hary of sintifi inlinations toward a purly m-
hanial univrs. By ontrast, numrous writrs hav usd th dotrin of
sondaryauss-that God works through natural agnis-to argu for a
.lfarfild,
form of matrialism onsistnt with th Judo-hristian hritag. for
64 uoltio nd Rligion

xilmpl, insisd that,,a omplt systm of naturl ausation'' was ntirly told tlr radrs of his 1
..tlologialsystm implis a om- ..uslss
onsistnt with tlology and tlrat vry to pt ptlpl
plt .auso-mhanial' explanation as its instrrrmnt'' (quotd in Livingston trs of th Bil, if thy d
and Noll 2000,301). Diffrnt though all ths rsponssundoutdly ar, its first hapters'' (Pi1
thy ar witnss to matrialism as a rurring lous of dat. Th ralization that th
losly onntd with ths onrns is a st lf qustionstht ongrgat tIu"tl L'olutiontlI srip
'l
aror.rndth implizrtions of volutiorr for th ltLlrnd morlit of the a fr'rrthrpoint of issu.
hm get. Vaious stans hav n adoptd on th issu of whthr puliation of Darwin's (
human ings bu sods or ar souls. Som hav optd for th viw that th th Bil had to intr
soul is always th rsult of an at of divin ration. Thus th atholi volu- was widly doptd v t.
tionist rnst ssngr osrvd in his 1931 ount uolutio d Theol. prinipl of volutiona
..no hypotlrsis manating fr
og t,at Ctholi thologian or philosophr ould possibly allow that
th spiritual soul of man has volvd from a brrrtesoul, muh lss from inor- tion atd no stir wht
gani mattr. Th soul is immatrial, and an only om into istny a no among thosnthus
dirt at of ration'' (ssngr 19\, 14)' Othrs hav thought that th By th sam tokn, as wl
id of th soul mrging in som way from arlir forrsis ntilyom- applid to oth th instit
ptil with thir thology. As for th thial rprussionsof Darwinism, to th way in whih D:rr
som hav oild at th suggstionthat moral snsiilit has n shapd Jws. Suh uss of volut
y th imprativs of a struggl for survival. In suh a snario, as \William ing to many. Gorg Frd
..Grat in good part from his
Jnnings Bryn, th ommonr,'' mad lar, thr is no rason to
xpt humans dsndd from animals to hav any diffrntly from ani- mntay history. In S
rnals. Othrs' nthusd with th possiilitis of lsmi volution, hv sught, Williar Rortson Smith
lik Hnry Drummond, to us th languag of volution to mak sns f spookd onsrvativl
thial and spiitual dvlopmnt. Again, although diffrnt rligious om- hang smd thr tr
mntators hav takn diffrnt positions on th qustion, th issu of this attituds to Darwinism .
and volution has prsistntly manifstd itslf. thory's rlvan for ro
Howvr amiguous in Darwin's own thinking, th idas of progress and,
in rtain strands of rligious thor-rght,its thologial nxt-door nighor
shtolog hav frquntly ntrd th disssion. Som of ths assoiations Putting volution ar
wr idntifid y rnst Bnz in L-,uolutiod bristi Hop (1966), whih
assssdth rlationship twn Drwinism and a varity of futur-orintd Rlations btwn volu
idologis-arist and matrialist historis of salvation, Nitzsh's futur- Attituds, morovr, dl n
isti dotrin of th Urmnsh, zrnd spulations on volution and th fu- fssionirl traditions, or lo
tur of humanity in th writings of th Hindu Sri Auroindo. Within th light of this raliztionw
.Wstrn
Christian traditin h flusdon th volutionizd shatologyof t,ll'tion and rligio tr
Teilhard d hardin' who onivd of vlution as impelling all lif toward soial, or sintifi world
an ..Omga Point.'' Th writings of Jams Mosh, Hnry Drummond, and l l s i n d i f r n I o e t i
Gorg Frdrik Wright also atn within th ar of this analysis. Indd, politil, onomi, mora
thr is muh to said for olzrtingattituds to volution with shto. th rltionship twnt
logial stans. Postmillnrians, witlr thi oust onfidn in tlr trans- u n d l s o f i d a s .A l s st r
forming powr of hristian iviliztion, hav routinly n favoraly episds might thrfore1
disposd toward volutionary prlgrssivism,whil prmillnnialists,who - attak on th lrgy in hi
liv that th world will ontinu to dgnratuntil th millnnium is ush. Prsbytrians to rad v<
rd in, typially find volution rpugnant.Thus ir is not surprising that th with tlr thrats of ilil
modrn foundr of ration sin,Gorg rady Pri, flt that blifs Prsytriilns thr wr
aout th gir-rning and n]of tim wr umililly onntd,and that h Za|and rsponssw s

lf natural ausation'' Was ntirly
: | o l o g i asly s t mi m p | i s O m -
lstrumnt'.(qutdin Livingson
thsrsponss undoutdly ar,
'inglous f dat.
st of qustions that ongrgat
te ttur nd morlit of the
rdoptd on th issu of whthr
hav optd for th viw that th
:ration.Thus th atholi volu-
31 aount uolutio nd bol-
sophr ould possibly allow that
r brut soul, muh lss from inor-
:an only om into eistn y a
43). othrs hav thought that th
r arlir forars is ntirly om-
.ial rprussionsof Darwinism,
moral snsiilityhas n shapd
.William
al. In suh a snario, as
mad lar, thr is no rason to
' hav any diffrntly from ani-
; of osmi volution, hav sought,
g of volution to mak sns of
l t h o u g hd i f f r n tr l i g i o u s o m -
n th qustion' th issu of this
tslf.
rinking, th idas of progress and,
's thologial nt-door nighor
: u s s i o nS. o m o f r h s s s o i a r i o n s
d hristin Hop (1966), whih
;m and a varity of futur-orintd
is of salvation, Nitzsh's futur-
:ulations on volution and th fu-
.!7ithin
{indu Sri Auroindo.
r th volutionizd shatology of
lution as impelling all lif toward
osh, Hnry Drummond, and
n th ar f this analysis. Indd,
:titudsto volution with shato.
:ir roust onfidn in th trans-
L' hav routinly n favoraly
m, whil prmillnnialists'who -
nratuntil th millnnium is ush-
:. Thus it is not surprising that th
1rady Pri, flt that lifs
umiliallyonntd, and that h
uolutio d Rligio 65
told th radrs of his 1923 Sie d Religil i Nlltshll tat it was
..uslssto xpt popl
to liv in th prditions givn in th last hap-
trsof th Bil, if thy do not livin th rord of th vnts dsridin
its first hapts'' (Pri 1923, 13).
Th ralization that th prinipls of Darwinism might applid to th
ttul uolutio lf sriptur and to the merg of religion itslf has n
a furthr point of issu.Th ida of soial volution, of ours' prdatd th
puliation of Darwin's origi, and th suggstion that th txtual history of
th Bile had to intrprtdin th light of soity'sstdial dvlopmnts
was widly adoptd y ttual ritis in th nintnthntury. Indd, th
priniplof volutionary hang was so dply ingraind in th doumntary
hypothsismanating from Grmn highr ritiism that Darwin's invn-
tion ratd no stir whatsovr in that Old Tstamnt sholarly ommunity
nor among thos nthusiastiaout the puliation of sss nd Ruies.
By th sam tokn, as w hv sn' th aliztion that volution ould
pplid to th th institutional and th ritual history of Isral was rlvant
to th way in whih Darwin's hallng was rad among nintnth-ntury
Jws.Suh uss of volutionary thought forms, of ours, hav bn distur-
.Wright's
ing to many. Gorg Frdrik ltr wvring on Darwinism sprang
in good part from his onrns aout its impliations for th Bil's dou-
mntary history. In Sotland th promulgatin of ilial ritiism y
william Rortson Smith with its dploymnt of volutionary anthropology
spookdonsrvativalvinists so muh that Dwin's own thoy of spis
hangsmd rathr tam in omparison (Livingston 2004). On again,
attitudsto Darwinism wr mdiatd through what was takn to th
thory'srlvanfor roadr ultural affairs.
Puttingvolution and Rligion in Thir Pla
Rlations twn volution and Iigion hav nvr n straightforward.
Attituds, morovr' do not map natly onto dnominational ontours' on-
fessionaltraditions, or loations on th onsrvativ.liralsptrum.In th
light of this ralization w might justifialy paus to wondr whthr th trms
th
ulltioand religio trak any transndntal ssnin th thologial,
soial, or sintifi worlds. At th vry last th wid-rnging uss of ths
llsin diffrnt loations and thir dploymnt in a host of disourss-
politial,onomi' moral' and soial-alrt us to th fat that in spaking of
thrlationshipbtwnvolution and rligion w a handling rathr triky
bundlsof idas.A lss transndntaland mor loal approah to partiular
pisodsmight thrfor prov nlightning. For ampl' whras Tyndall's
attak on th lrgy in his Blfast addrss of 1874 mad it diffiult for loal
Prsytriansto read volution thory sympathtially, th proupation
with th thrats of ilial ritiism at th sam tim in Sotland mant that
Prsytriansthr wr lss risd aout iologial mattrs. In Nw
Zea|andsponsswr shapd y th politis of Maori.sttlr rlations, and
66 uoltion d Rligion

in th Amrian South raial qustions prdominatd,though with diffrnt Brundll'B. 2001. tholi
outoms in ah as. h way in whih Amrian atholis rspondd to lourl for th Ilistor
soial Darwinism in th yars round 1900 ws shapd y th soial nih Burkhardt,F. ., and S' Sm
D r t u i . 1 5v o l s . a m
thy oupid: Amrian Catholiism Was vry largly a woking-lass rli.
yn' G. V. 1998.voluti
gion of nwomrs struggling to maintain thir idntity in nw nvironmnt.
T. Ptrs,d.,Scic
In all ths instans th partiulars of gographi loation and soial spa .Wstviw
Prss.
had hug rol tl play in what ould said out volution and, just as D r w i n , . 1 8 5 9 .o t h
important, what ould hard aout it. 1888.e Life d
From th momnt Darwin put forward his thory of volution y natural urray.
|959. h rigio
sltion, th dat was ngagd on its signifian for a wid rang of
N4.Pkham,d. Phil
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Drummond,H. 188.Na,
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, {Jiurs,or, Phsil Spultions

; to Darwin and volutionary thory'

hous,ds., Dissmintig Duiism:
edr. amridg: amridg
 That irnirosity is found
ptane <lfv<liutilrr(or .
hildrrr to aandon thi
God' thn that hild is lst
tians. Furthrmor' hildr
Amrian Antivolutionism: to guid thm and thus wil
itizns. Soity would sul
Rtrosptand Prospt ommon rationist lai.rI
thy ar animals, thy will
th img of God. In ddi
ugi . Sott tion, thr a tholgial
for hristiirn lifs:if Ad;
als, spially ratd y G
ast ollt from th Gardn
W ar not inhrntly sinfu
urrtion' th pivotal vrr
Antivolutionism in th Unitd Stats is ntring its sond ntury. Bgin-
o f o n s v t i v h r i s r i a n s
ning in th arly twntith ntury and ontinuing today, with no sign of
atly as writtn, and volu
rlnting, antivolutionistsh:rv protstd th thing of volution to hil-
forms of iblial litralism
drn in puli shools. Th ontrovrsy has wad and wand irr strngth
atholis and mainstrm
in dirt rsponsto th mount of volution tht is taught in th lassroom
hurh of Christ, most
and found in ttooks. It is a onrplx ontrovrsy with a long history and
glial Luthran hurh irl
dp roots in distilltivlyAmriarr ttitudstoward rligion' duatior,and
liv tlrat God ratd th
st11.
h hyday of tlris first
attmpts wr mad in sv
Th st-known rpuss
Th History of Antivolutionism Tnnss tahr John S
itd the tahing of voluti
ANNING VoLUloN
lld 'ausof th hith
To undrstand Amrian antivoIutionism,on must undrstand th history dia publiity that it ngnd
of this ontrovrsy' whih an dividd into th tim priods (Sott on a thniality y th Tr
7997). Th first onsists of th fforts to an volution from th pu[i shool tion of th trial as a dfatf
.!7ll
]assroom. for th sond dad of th twntith ntury, volution f1|m Ihrit the Wid), in
had found its w1, into high_shoolttooks and tlrryinto the utrlu- volution in ttooks and
lum. Aording to Larson, a dnrographitrnd in uran popultion growth Inded, aording to nt
triggrd th first round of rrivolutionismin th Unitd Stats (Larson fronr tlr urriulum l'thr
2003). High shols at that tim wr largly an urn phnomnon, and th (Lrsolt 200). voIution
populations of itis-fuld otlr y hanging onomi onditions that and arly 1960s, whn p
pomptd th movmnt of popl from farms into itis and y forign fundd Biologial Sin
imnrigration-Wr irrrasing' As a rsult, mor studnts Wr ttnding mattr_of-fatlyinludd .
high shool y th sond dd of th twntith ntury than vr for. u l lt h o n t n r o i r h 1
Although volution had n part of high.shool iology for ovr dad,it ear| |'97s volution was r
did not om an issu until mor hildrn Wr xposd to it. on that txtooks ing sold in th
happnd, onsrvativ hristiarr animosity toward volution promptd riulum stimulatd th st
fforts to rmov volution from th urriulum. th priod >frtion sil
.70

lutionism:
spt
ring its sond ntury. Bgin-
rtinuing today, with no sign of
r tahingof volution to hil-
s waxd and wand in strngth
n tht is taught in th lssroom
trovrsywitlr long histry and
s toward rligiorr'duation, and
on must undrstnd th history
l into thr tim prids (Sott
' volution from th puli shool
I th twntith ntury, volution
lks and thry into th urrlu-
rnd in uran population growth
;m in th Unitd Stats (Larson
y an uran phnomnon' and th
'nging onorrri lnditions that
farms into itis and y forign
[' mor studnts wr attnding
.ntithntury thn vr for. ulatdth ontnt <lfth popular BSS ooks, and y th lat 1960s and
;hool iology for ovr a dad' it
n Wrxposd to it. on that
;ity toward volution promptd
lum.
Amrin tieuolutitlism 71
That animosity is foundd on a strongly hld ligious onvition that a-
ptanof volution (or ..lif'' in volution, s it is ommonly put) ar'rss
lrildrnto aandon thi faith in God. In this viw' if a hild loss faith in
God, thn that hild is lost to slvation, a srious issu to onsrvativhis-
tians.Furthrnror,hildn who lak lif irr God will hv no moral rddr
to guid thm and tlrLrswill hv no rason to hav proprly to tlrir fllow
itizns.Soity wuld suffer gratly, all usof ..lif''in volution. A
ommon rationist laim rlatd to havior is that if hildrn ar taught that
thy ar animals, thy will hav lik animals rathr than raturs mad in
th imag of God. ln ddition to ths pratial rasons for rjting volu-
tion, thr ar thologial rsons as wll. Gnsis is sn as th foundation
fr hristian lifs:if Adar and v Wr not ral flsh-nd-loodindividu-
als' spiaily rtd y God, whos sin of disodinausd thrn to
astout frm th Gzrrdenof dn, thn lrurnrrkindis not inhrntly sinful. If
War not inhrntly sinful aus of th Fall, thn Christ's sarifi rrd rs.
urrtion' th pivotal vnts for hristinity, wr unnssary.In th opinion
of onsrvativ hristians, th ook of Gnsis thus must e intrprtd -
atly as writtn, nd volution is rjtd baltsit is inompatil with mst
formsof iblial litrlisrn.Ndlssto say' this is not th only Christian viw;
atholisand mainstramProtstants(pisopalians,mmrs of th Unitd
hulrof hrist, most Mthodists, most Prsytrians,mmrsof th van.
glialLuthrn hr"rrhin Amria, arrdothrs)ar thistivolutionistswho
livthat Gd rtdthrough volutiorr.
Th hyday of this fist priod was approximatly 1919 to 1927, when
attmptswr mad in svralstat lgislatursto an th tahing of volution.
Th bst-known prussionof thsfforts WaSth famous 1925 Sops trial.
Tnnsstahr John Sops was trid for raking a stat law that prohi-
itdth tahing of volr"rtionin th puli shools. Th trial of th ntury, so
alldausof th hithrto-unxpindoast-to-ollstradio and print nl-
dia puliity thzrtit ngndrd,rsultd in th onvitin of Sops (ovrtund
on a thniality y th TnnssSuprm ut). Dspit th puli prp-
tion of th trial as a dfat for Fundamntalisnr (as nouragd y th ply and
ft|mInhrit th Wid), in its aftrmath am th gradual diminishmnt of
volutionin ttooks and thforin th urriulum of puli shools.
Indd,aording to on ommntator, volrttion was fftivly removd
from th urriulum y th 190s, and it did not rturn for svral dads
(Lason2003). volution rturnd to th lassoonr only in th lat 1950s
and arly l960s, whn pofssiorr:rIsintists wrking with th fdrally
funddBiologil Sins urriulum Study (BSS) wrot trooks that
mattr.of-fatly irrluddvolution. ommrial ttooks susquntlym-
ear||970s evolution was usually, though not amply, inludd in most of th
ttooksing sold in th Unitd Stats.Th rturn of volution to th ur-
riulumstimulatdtlr sond wav of antivolutionism in th Unitd Stats:
thpiod of rctilt sitt..
72 mric ntieuolutkism

ollg. |n 1972 h form

RAIoN s|EN
n t i f i a t i o n i s mu n i t \
Whn volution rturnd to ttooks in th mid- to lt 1960s, onsrvativ apparntly amialy' on
hristian antivolutionists again flt th nd t prott thir hildrn from Sant' alifornia' as
larning what thy blivd tl an antiligiousida. Not only had volu- 1992). Aftr Morris's rt
tion n rturnd to th urriulum, ut also as a rsult of ourt disions dirtorship of his son, J<
(partiularly pprso u. Arkss, 1987), volution uld no longr Morris's 1961 ook Tl
annd. Prhaps, thn, th Bil ould taught alongsid volution; Y/hitom Jr., proposd
ntivolutionistsblivdthat if qual tim wr givn to Gnsis in th lass- p I i n d y a r a s r r o p h i
room, fwr studntswould apt volution. But rathr than mrly p
But th stalishmntand fr-xrislauss of th First Amndmnt to om and Morris lim
th U.S. onstitution rquir that puli institutions,suh as shools, rli. idas. ration sinw
giously nutral: thy may nithr promot nor inhiit rligion. To tah th Cration sin was
ontnts of th Bil as fatully orrt (as opposd to tahing ompara- C h r i s t i a n o m n l u n i t y .t t
tivly aout rligion) would promot a starian rligious viw, prosrid sists mostly of ngativl
y th stalishmntlaus. Th ffort to rquir that th Bil taught as i n g ( i n i t s p u r s rf o r m .a t
a alan to volution in sinlasssthus had only a rif history: it was brupt parn in its
forordaind for rapid rjtion y th Suprm ourt. \n 197 th stat of sin that wr to
Tnnsspassd law tht rquird ttooks that inludd volution to ationism, to tught i
..thory'' ..not r o o t s o f r t i o ns i n
arr a dislaimr stating tht vllutionwas a that was rpr-
sntd to sintifift.'' (Th Bil was spifiallydfind as a rfrn a n d t h r t h n d i t s l i v i
ook rathr than a ttook and thus did not hav to display a dislaimr.) as a rsult of rativ at
In 1975 an appals ourt struk down this law in Dil u. Wters on th from oth sin and th
grounds that it violatd th stalishmntlaus. Th distast of hristian p r o p o s d t t u g h ri n
onsrvativsfor volution Was not a valid rason ithr to an voiution or only sintifi vidn
..Thr stripping rfrnsto
to prsnta starianviw suh as ilil litralism in th lassroom.
is and an no dout that th First Amndmnt dos not prmit th Statto ism, ration sinwtlt
rquir that tahing and larning must tailord to th prinipls or prohi- shools.
itions of any rligious st or dogma,', Wrot th judg in Diel. v and Harrold (199
If volution ould not annd, and th Bil ould not taught along- p l o d u a t d h r i s r i
sid it t amliorat its prsumd ngativffts,what roursdid antivo- modrn ..knowldg o
..
lutionists hav? Rligion ould not advoatd, but altrnativ sintifi valus that mphasiz
viws ould: if th ration story of Gnsis ould supprtd y sin- through th Bil,' (199
tifi vidn' rationistsrasond,thn it would onstitutilnalto prs- oth worlds: thy an m
nt that vidn alongsid volution. In fat, vr sin ightnth-ntury rligious and soial valu
disovris in gology rfutd th litral Flood story and dmnstratd Inspird y orris,s tt
..sriptural gologists'' modl lgisltion that w
an anint g of th arth' onsrvativ hristian hav
fought a rrguard movmnt' arguing fr a young arth and spial r- lution wr taught.ln ri
..sin'' h irulatd it to frind
ation (Lynh 2002.\'ration ws uilt upon th shouldrs of ths
viws. to onsrvativ stat lg
Th tru giant of rtion sin,howvr' ws hydauli nginrHnry stat lgislatrrrsof at l
M. orris' who vn for h ntrdgraduat shool gan writing ooks l 9 8 0 s ( S 2 0 0 4 ) ;s i
tht prsntdwhat h viwd as sintifividnfor th spial ration of qual-tim ills passd a
th univs' ath' and living things. Aftr a arr in usinss and aad- rvas hallngd y a oal
mia, in 797 he lft th hairmanship of th ivil nginringDpartmnt t a P s y t r i a nm i n i s t r .l
Virginia Th to lrlp vanglist Tim [,aay found hristin Hritag sulting highIy puliiz

mid- to lat 1960s, onsrvativ
:d to prott thir hildrn from
igious ida. Not only had volu-
lso as a rsult of ourt disions
, volution ould no longr
taught alongsid volution;
vrgivtlto Gnsis in th lass-
n.
ussof th First Amndmnt to
titutions,suh as shools, rli-
nor inhiit rligion. To tah th
ts opposd to tahing omPra-
:tarianrligious viw, prosrid
quirthat th Bil taught as
r s h e d o n l y a r i fh i s t o r y :i t w a s
'rmourt. |n 197 th stat of
ooks that inludd volution to
..thory''that was ..not rpr-
sa
spifillydfind as a rfrn
not hav to display a dislaimr.)
.Wtrs
s lw in Dil u'
hristian
lus.Th distast of
rsonithr to an volution or
itralismin th lassroom.
lmnt dos not prmit th Stat to
tailordto th prinipls or prohi-
'otth judg in Dnil.
Bil ould not taught along.
ffts,what rours did antivo-
lvoatd,ut altrnativ sintifi
sis ould supportd y sin-
it would onstitutional to prs-
fat, vr sin ightnth-ntury
l Flood story and dmonstratd
..sriptural gologists'' hav
rristian
Or a young arth and spial r-
rs uilt upon th shouldrs of ths
)vr'was hydrauli nginr Hnry
aduatshool gan writing books
vidnfo th spial ration of
:r a arr in usinss and ad-
h ivil nginringDpartmnt at
-aHay found hristian Hritag
meri Atiuolutioism 7
llg.|n1972 h formd th Institutfor Cration Rsarh (IR) as a si-
ntifirationism unit within th ol1g.In 1980 h and LaHay agrd,
apparntlyamialy' on a sparation, and Morris took th IR to nary
Sant, alifornia, as an indpndnt nonprofit orgnization (Numrs
\992|' tfter Morris's rtirmntin 1996, th IR ontinus tody undr th
dirtorshipof his son, John Morris.
Morris's 1961 ook Th Geesis Flood' oauthord with thologian John
.Whitom
Jr.' poposd that th arth's gologial faturs ould x-
plaindy atastrophivntsdsridin th Bil' spiallyNoah's flood.
But rthr than mrly positing thologial rasons for living this, lfhit-
omb and Morris limd that sintifithory and vidnsupportd ths
ids.Cration sinwas orn.
ration sin Was nthusiastially mrd within th onsrvatrv
hristin ommunity, though sintists undrstandaly ignord it. It on-
sistsmstly of ngativ statmntspurporting to disprov volution, offr-
ing (in its purst form, at last)no xplnation for th univrs othr thn its
aruptapparanin its prsnt form. orris proposd varitis of ration
sinthat wr to dployd in diffrnt nvironmnts. Biblicl cre'
ationism,to taught in hurhs, was up-front about th spial-ratlon
rootsof ration sin,frly adritting a lif tht th stars, th galis,
and th arth and its living things appard at on tim in thir prsnt form
as a rsult of rativ ats y God. In biblil sitifirationism vidn
from oth sinand th Bil was to usd. |n te scitif rationism
on th
proposdto taught in th puli shools, God was not to b mntiond;
only sintifividnWas to usd (Morris 1980). Th hop Was that y
..Thr strippingrfrnsto God and Gnsis from unashd ilial ration-
ism,ationsinwould md lgally suital for instrution in puli
shools.
v and arrold (199\) ontnd that ration sin has a partiular ap-
palto duatd hristin onsrvtivswho ar living and working in th
modrn ..knowldg onomy''' ut who still mra prmodrnist soial
valusthat mphasiz ..authrity, tradition, and rvlation of Gtld's truth
through th Bil'' (1991,109). ration sin allows thm th st of
bothworlds: thy an mra sinyt rmain faithful to th onsrvatlv
religiousand soial valus undrgirdd y lif in a litrally rad Bil.
Inspirdy Mlrris's tahings,North rolinian Paul llwangr draftd
modllgislationthat would mandat th tahing of ration sinif vo-
lution wr taught. In tim-honord Amrian grassroots politial fashion,
h iulatd it to frinds in a varity of stats, who thmslvs passd it on
to onsrvativstat lgislators.Suh ..qual.tim''ills wr introdud in
statlgislaturslf at last 23 statsduring th mid- to lat 1970s and arly
1980s(Sott 204); sintists and tahrs vigorously opposd thm. Two
qual-timills passd nd wr immdiatly hallenged.Arkanss Bill 590
washallngdy a oalition of rligious orgnizations and duators,with
a Prsbytrianministr, th Rvrnd Bill Lan, s lad plaintiff. Th r.
sultinghighly puliizd 1981 trial, L u. Arkss, was invitaly
74 merin tiullutirlism
..SopsII''
illd as y th prss' although during the 1'925 trial no laims vanglism ministris of
wr md for th sintifivalidity of spial ration. ral hargs of failing to
Le Was a rsounding dfat for ration sin.Th plaintiffs argud yars'' 2007D,.!7alt Br<
a straight stalishmntlaus position: ration sinWas not sin,and Spakrs fom thsmir
thrfor thr was no sular purpos for thing it in puli shool sin ountry' spaking at hu
lasss.Its tahing would thrforrsult in th stat promoting rligion,in thy an find an audin
violation of th First Amndmnt. Th stat'S witnsss (th ration sid) individual wsits pro
had littl rdil sinto prsnt' and th dfnsWas marrassdwhn invok lassi ration s
th history of th ill,s passag providd lar vidn that th Arkansas musum that laims tou
qual.tim law was proposd and passd for th purpos of singling out on AIG opnd a 90,000-sq
topi' volution, from all othr urriular sujts aus it offndd th r- northrn Kntuky. With
ligious snsiilitisof som itizns. Th law Was struk down. S firm was rady xdd its p
th dision that th stat dlind to appal to a highr ourt. tors in its first yar (..(
Th nt qual-tim law to ntd Was a vry similar ill passed in rationist Musum of i
nry Louisiana. Th distrit-urt judg didd on summary judgmnt plannd to opn nar th
(that is, on th asis of doumnts fild y oth sids,without a trial) that th Thr is, thrfor, plnt
Louisiana qual-tim law violatd th stalishmntlaus. Th stat also posd to ration sin
lost on appal and thn rqustda haring from th Suprm ourt, whih ativitis' puliations, l
aptd th as. |n a 7987 dision th Suprm ourt agrd 7-2 (Justis duation wryly rmark,
.!illiam
Antonin Salia and Rhnquist dissnting) with th lowr urts that formal duational oppo
rtion sin was an inhrntly rligious position' and thus its tahing Nlson 2002).
unonstitutionlly would promot ligion. Th Suprm ourt hos not to
ommnt on whthr or not ration sin was sintifi, living that its
NEoRAloNIst\
Ssntialrligiosity was suffiintto ar it from th lassroom. Th dision
in drds u. Aguillrd statd: Nortionism rfrs to
Th At imprmissily ndorssrligion by advaning th rligious - largly to avoid its lgal p
..altrnativ
lif that a suprnatural ing ratd humankind. Th lgislativhistory sintifi
(ID), and/or propsing t
dmonstratsthat th trm ..ration sin,''as ontmpltd y th
stat lgislatur,mrasthis rligious tahing.Th At's primry pur- ut y allged vidn :
pos Was to hang th puli shool sinurriulum to provid pr- agd y th wording of t
suasiv advantag to a partiular rligious dotrin that rjts th ration sin.
fatual asis of volution in its ntirty.Thus, th At is dsigndithr ,,Altttiue sitifi t|
to promot th thory of ration sinthat modis a partiular re-
In h dlllrds dision
ligious tnt or to prohiit th tahing of a sintifithory disfavord
y rtain rligious sts.In ithr as,th At violats th First Amnd- ould not advoatdar
mel:t. (drds u. gillrd, 578) ontnt' but that ..tahi
humankind to shoolhil
ration sin'slgislativstratgythus am to a halt at th foot of th tnt of nhaning th ff
Suprm Court's stps. But ration sin itslf did not withr; instad,it teahrs ould tah ..all l
xpandd during th 1990s and shows no sign of diminishing during th (dturds u. guillrd, 5|
twnty.firstntury. In Janury 2005 sB 2286 was introdud into (although thoris to volution' ut
..puli
it faild to pass) th Mississippi stat lgislatur, proposing that thir istn and provid
shools within this stat shall giv land tratmnt to th thory of sin- puli shools. ration s
tifi rationism and th thory of volution.'' lassi ration sinan lution; with its failur to
found in th xtnsiv w prsnsof th IR and its sistr organization, rquird rpakaging. Th
Answrs in Gnsis (AIG). as wll s in th indpndnt ration sin i n o n . . a I t r n a i vs i n

during th 1925 trial no laims
al ration.
ion sin.Th plaintiffs argud
tion sinwas not sin,and
,ahingit in puli shool sin
1 th stat promoting rligion, in
t'switnsss(th ration sid)
dfnswas marrassdwhn
lar vidnthat th Arkansas
r the purpos of singling out on
jtsausit offndd th r-
iv was struk down. So firm was
l to a highr out.
/asa vry similar ill passd in
didd on summary judgmnt
oth sids,without a trial) that th
lishmntlaus. Th stat also
;from th Suprm ourt' whih
prm ourt agrd 7-2 (Justis
:nting)with th lowr ourts that
s position, and thus its tahing
Th Suprmourt hos not to
:was sintifi, living that its
:omth lassroom. Th dision
y advaningth rligious -
rankind.Th lgislativhistory
n,'' as ontmplatd y th
:ahing.Th At's primary pur-
:nurriulum to provid pr-
ious dotrin that rjts th
hus,th At is dsignd ithr
: that modisa partiular r-
lf a sintifi thory disfavord
r At violatsth First Amnd-
s am to a halt at th foot of th
itslf did nt withr; instad' it
sign of diminishing during th
,86ws introdud into (although
..puli
;islatur,proposing that
l tratmntto th thory f sin-
:r.''lassi ration sin an
I C R a n d i t s s i s t ro g n i z t i o n ,
t h i n d p n d n t r t i O n s i n
Amri ntiuolutioism 375
vanglism ministris of Knt Hovind (until his rnt inarration on fd-
ral hargs of failing to pay payroll taxs [..Knt ovind sntndto 10
ears',207|, lfaltr Brown, John aKay, and arl Baugh, among othrs.
Spakrs from ths ministris and othrs ar onstantly rissrossing th
ountry' spking at hurhs' ollgs, and vn publi shools-anywhr
thy an find an audin to listn to th antivolution gospl. Innumral
individual wsits promot ration sin, and tlvanglists frquntly
invok lassi ration sin argumnts to attak volution. Th IR has a
musumthat laims thousands of visitors annually, and in th spring of 207
AIG pnd a 90,000-squar-footpuli musum nar its headquartrs in
.!7ithin
northrn Kntuky. th first svn months of its opning, AIG had al-
rady dd its ptd annual attndan' drawing ovr 300,000 visi-
tors in its first yar (..ration usum Unxptd Suss'' 2008). A
rationist usum of arth History, whih will fous on Flood gology, is
plannd to opn nr th popular Branson, Missouri, tourist dstination.
Thr is, thfor' plnty of opportunity for th Amrian puli to x.
posd to ation sin through th movmnt's various puli outrah
ativitis' puliations, lasss' and rvivals. As two spialists in volution
duation wryly rmark' ..any studnts hav hd ampl formal and in-
formal dutional oppotunitis to misundrstand volution'' (Altrs and
Nlson 2002\.
NoREAloNIsM
Norationism rfrs to th post- drds rpakaging of ration sin
larglyto avoid its lgal prolms (Sott |997). It inluds proposing allgd
..altrnativ
sintifi planations'' to volution, suh as Intllignt Dsign
(ID), and/or proposing to ountr volution not with an altrnativ sin
but y allgd evidn against volution. Ths approahs wr nour-
agd y th wording of te drds dision itslf, ut thy pristd in
ationsin.
,,Itrntiue
sintifi tbories,': IntIligntDsign
\n the urds dision Justi \/illiam Brnnan Wrot that rationism
ould not advoatd as an ltrnativ to volution aus of its rligious
ontnt)ut that ..tahing a varity of sintifi thoris about th origins of
humankind to shoolhildrn might b validly don with th lar sular in-
tnt of nhaning th fftivnssof sin instrution.'' H also Wrot that
tahrsould tah ..all sintifi thoris aout th origins of humankind''
(drds u. gillrd, 59_594). In fat' thr ar no sitc altrnativ
thoristo volution, ut Brnnan's ommnt invitd rationists to propos
thir xistne and providd a lgal foundation for arguing to prsnt thm in
publi shols. rtion sinWas th original sintifiltrnativto vo-
lution; with its failr to onvin th sintifi ommunity or th ourts, it
rquird rpakaging. Th most important rpakaging of ration sin
into an ..ltrnativ sintifi thory'' is Intllignt Dsign.
376 meric Antieuolutionism

Te drds dision took a long tim to work its way through th ourts, ster was publishd,
ut vn tr th Suprm Court's dision ws issud in th summr of indst: origin of Lif l
7987, man antivolutionists had sn that th antivolution movemnt would ontnt in DNA'' in 198
hav to go yond ation sin and spially yond th young-arth nt sponsorship, follow
rationism (Y) of Hnry . Morris and th IR if it wr to xpand and Philosophial frn j
sussful.Thy sought form of antivolutionismtht oth would hav of Culrur''1 1996,.,er
mor sholarly rsptaility and would avoid th First Amndmnt Pro- Sientifi ntrprise.''
.!7ht
lms of ration sin. was Soon to e lld IntlligntDsign was Som of th fas at tl
nouragd y Jon Bull' ministr who dirtd (nd still dirts) a Txas who follows tlr antiv
nonprofit organization alld th Foundtion for T.houghtand this (FT). ationists' suh as Paul N
Bull nouragdfllow antivolutioniststo downplay ilial litralism and B i l - S i n A s s o i a t i o
Gnsis imagry and to invok a vagur form of spial ration to plain latr orgniztion g
highly omplx natural phnomna. In this apprlah God th Dsignr ationist studnts at th
would not rfrdto, and th inaility of volution to plain omplity morphd into an ID ad'
would strssd.1Bull notlragdth puliation of The ster lf Lif's sarh Ntwork. ost
origi (Thaton et a|' 1984), whih was writtr thr sintists assoi- a t d w i t h n o l d - a r t hv i
atd with FT. Although th ook onntrats only on allging wak- and dirts th llasons
nsss in thn-urnt thoris of th origin of lif, srupulr"rslyrlgating suh as mathmatiian/1
disussion of dsign to an pilogu' it is now rgrdd s a founding doumnt C. lvlyr, and thologra
of ID. fw of whih inludd
str ontndd that th origin of lif was an ampl of a lass of ph- movmnt to dvlop its
nomna that wr too omplx fo plantion hrough nurl auss.his strtgy for th otsid r
was atgorially so and was not th rsult of inor-npltknowldg. Th ori- vn in th ginning,
gin of lif was not only a diffiult sintifi prolm, it was an unsolval on lassroom (Sott and M.
if on rstritd onslf only to natural alrss. Narr-rralauss Wr qui1td ouragd th writing an<
with han, and suh omplity as that hiitd in th strutur of th first t o h i g h - s h o o l i o I o g yt
ll ould not xplaind y th random joining of llular struturs. Suh tion was that Bull soug
diffiultis ould ovrom' Thaton nd his oauthors laimd, on[y y tr would mak th blt
inluding th tions of n intlligntagnt in sintifiplanation. 1 9 8 9 ) .U n d t h a u s p i
str prsntdth possiility tht th agnt was matrial (suh as in. oauthord of Pds r
tllignt lins), ut it optd for a transndntagnt..yond th osmos'' was long on ritiisms o
(Thaxton et a|. 1984,200)-God. In th pilogu of th ook th athors pla it. FT attmptdt
spok frly of spial ration and th nd to xpand sin to inlud ook was widly rjt
suh ausation as a mans of otaining omplt viw of truth. |n 1997 a law profs
str had littl impat on th sholarly world. An Institut for Sintifi Phillip Johnson, publish
Information (ISI)itation ount notS only a fw rviws, largly in philos- Although h ster <
phy and rligion journals, and a total f only 28 itations twn 1984 and sarly any noti in thr
2008. In omparison' anothr popular ilount of th origin of lif, A. G. noti fom th gnral p
ains-Smitlt,sSue lus tl th origi of Lif, pulishd in 1985, had 95 iologist, lit in zr po;
itations during this sam tim priod. Although uninfluntialin th wold of baus of th novlty o1
..om-
sin' str is rvrd in rationist irls, whr adjtivslik ing an antivolution
plling'' and ..impotant'' ar irrvarialyusd to dsriit. only with privatorgn
dmi world; Johnson's i
SABLISFIING ID. on of th authors of str, historian and hmist guably put ID on th ma
harls Thaton, promotd th nw ID nrovmnt y organizing onfrns standard rationist argu
Th first ws hld in th sam yr that
of fllow hristin ntivolr.rtionists. rros.

work its way through th ourts'
on Was issud in th summr of
l antivolutionmovment would
pially yond th young.arth
th IR if it wr to xpand and
lutionism tht oth would hav
.oid th First Amndmnt pro-
l l d l n t l l i g n D s i g n w a s
irtd(and still dirts) a Txas
'n for Thought and this (FT).
downplay ilial litralism and
rm of spial rtion to plain
ris approah God th Dsignr
f volution to xplain omplxity
liation of The str of Lif's
v r i t t n y r h r s i n t i s t s s s o i -
]ntratsonly on allging wak_
n of lif, srupulously rlgating
rgarddas founding doumnt
Wasan xampl of a lass of phe.
tion through natural auss. This
lf inomplt knowldg. Th ori
lrolm,it was an unsolval on
ss.Natural arrsswr quatd
hiitd in th strutur of th first
oining of llular struturs. Suh
d his oauthors laimd, only by
in a sintifiplanation.
agntWas matrial (suh as in-
..yondth osmos''
dnt agnt
:piloguof th ook th authors
d to pand sin to inlud
lmpltviw of truth.
r world. An lnstitut for Sintifi
a fw rviws,largly in philoso-
ly 28 itations twn 1984 and
:ount of th origin of lif, A. G.
f Lif, ftlrishd in 1985, had 95
ough uninfluntialin th world of
..om-
rls.whr aditivs lik
d to dsriit.
| str, historian and hmist
,vmenty organizing onfrns
rst Was hld in th sam yar that
meric ntieuolutionism 77
str was pulish d, 1'984, and was titld ..Going yond th Naturalisti
Mindst: origin of Lif Studis.'' It was followd y ..Sours of Information
ontent in DNA'' in 1988. Othr grandly namd ID onfrns,undr diffr-
..Darwinism:
nt sponsorship, followd 1992 rought Sintifi Infrn or
PhilosophialPrfrn?,';1995,..ThDath of Matrialism and th Rnwal
of ultur''11996,,,ere ration''; and7997,..Naturalism, Thism, and th
Sintifi ntrpris.''
Som of th fas at ths onfrns would hav n familiar to anyon
who follows th antivolution movmnt' inluding a fw young-arth r-
ationists'suh as Paul Nlson, who had workd with th innapolis-asd
Bil.SinAssoiation and latr with Studnts for Origin Rsarh. h
lattr organization gan as a group of prdominantly young-arth r-
ationist studnts at th Univrsity of alifornia at Snta Brara and latr
morphd into an ID advoay organization with th odd nam Ass R-
sarh Ntwork. Most of th onfrn partiipants, howvr' wr assoi-
atd with an old-arth viw, suh as Hugh Ross, an astronomr who foundd
and dirts th Rasons to Bliv ministry, and youngr antivoluionists,
suh as mthmatiian/philosophr l/illiam Dmski' philosophr Stphen
. Myr, and thologian and iologist Jonathan \/lls.Ths onfrns, a
fw of whih inludd dfndrs of volution' wr a mans for th young
movmnt to dvlop its idas and, mor important, to hon its mssag and
stratgyfr th outside world.
vn in th ginning, ID advoats aimd thir fforts at th puli shool
lassroom(Sott and atzk 2007). Bginning in th arly 1980s Bull n-
ouragd th writing and markting of a ttook intndd as a supplmnt
to hrgh-shool biology txtooks. Part of th rason for th dlay in pulia-
tion was that Bull sought a slar rathr than a rligious pulishr; th lat-
tr would make th ook far lss salal in th puli shool markt (Sott
1989).Undr th auspisof FT, in 1'989Pival Davis and Dan Knyon
oauthord f Pds d Pople, a rif ook that in typial ID fashion
was long on ritiisms of volution and short on spifis of what might r-
pla it. FT attmptd to promot it to shool oards for adoption, ut th
ook was widly rjtdy oth tahrsand sintists.
In 1991 a law profssor from th Univrsity of aliforni at Brkly'
hillip Johnson, pulishd th first widly rd ID ook, Dri o Tril.
Although The ster of Lif's rigi and f Pds d People attratd
sarlyany noti in th sintifi ommunity, Johnson's ook rivd wid
noti from th gnral prss and Was vn rviwd y notd volutionary
iologist,alit in a popular sin journal (Gould 1992). This was likly
auseof th novlty of a pofssor from a major sular univrsity author-
ing an antivolution ook. Antivolutionism had prviously n assoiatd
only with privat organizations suh as IR that wr marginal to th aa-
dmiworld; Johnson's involvmnt ratd a flurry of mdia intrst and ar-
gualyput ID on th map. Th ook itslf onsistd of a rworking of many
standardrationist argumnts and was roundly ritiizd y sintists for its
rrors.
78 l ric tiuo luti o i sl

Th nxt major ID puliaion was Drxai's Blk Bo y Lhigh Univ- himslf as living irr
sity biohmist Mihl Behe (1996), pulishd y th Fr Prss. Dri,s prominnt lD thorists]
Blk Bo marks a high point in ID's effort to ast itslf as a sin. Th tion'' (Dmski1995,3
prmis of this ook was similar to that of ster: som phnomna in n- votlts' inluding in hi
tur a simply too ompl to plaind through ntural auss.Spili- (Bh2007).
ally, Bh pointd to moleular struturs suh rsth atrial flagllunr As PhillipJohnsonha
and prossssuh as th lood-lotting asad s ing too orplx ro rO thir hif onrrr
plainal through inrmntal natural sltion. Bh,s ook rivd tlrrough natural pross
som rviws in th sientifi litratur-whil Johnson's book did not- intrvntion is rquird
prhaps aus hr again Was th novlty of a professor from a seular uni- phnomna ar dsign
vrsity pulishing an ntivolutionism ook, this tim from a mainstram through natural pross
press' Drlui's Blck Box also rivd mor attntion from sholars than is tlrat it is possibl ttl
Johnson,s Dri o Tril us it attmptd to provid a sintifi argu- ompl phnomna of
mnt. Howvr' th rviws y sintists wr lmost uniformly ngtiv iologial phnomna,.
(oyn |9 9 Q D orit 1 997; Blaks tone 1997i vlir-Smit 19971 Thornhill y ration sin.rhr
and Ussry 2000). poaility and informz
B 7998, whn philosophr and mathmatiian \illiam Dmski pu- Dmski has ontn
lishd his first ook, Tbe Desig lferc, th asi strutur of ID had (..spifid''in advan
n stalishd.In be Desig Infere and othr puliations Dmski things thar hv hnil
prsnted a proaility-asd xplnation fo th furrdamntalida of ID: ( S I ) , i t i s l a i m d 'i d n
that som natural phnomna ar too ompl to plaind through nat- quir tl-lation of int
ural auss and thrfor rqui ..intllign''for thir xplanation. Invok- a n d o a u h o r s ' o l l t l l t
ing an ..xplanator hlter', (p. 7) of inrsing improbability, Dmski lif was too omplii1t
laimd to al to remov natural auss and han as potntial -pl- to idntify a lass f ph
nations of omplx phnomna, laving dsign as th rsidual planation. natural' that is, sintiI
Dmbski has had a diffiult tim onvining fllow philosophrs and proa- Mihal Bh's ida
ility thorists of his viw and has n quit unsussfulat prsuading i- is similarly usd to argu
ologists that his onpt of ompl speed informtil has any utiliry for Bo (7996)that rtin
undrstanding iologial phnomn (Fitlson et aL. |999; Shanks 2004; e p l a i n a l y n a t u r a ls
Wilkins nd lsrry 2001; Rosnhous2003; Pnnok 1,999,20\ lsbrry th tril flagllurn,-
and Shllit2003). thir many omponnts
ursor of an irrduil
TH ONTN oF INTLLIGNT DSIGN. In ontnt' Intllignt Dsign dif- natural sltion-thou1
frs fom ration sin primarily in its rlutan to mak spifi fat ats on On omponnt
.s(hras
laims' ration sin maks laims aout th ag of the eath a harm ralt.Bh i
(only a fw thousand yars old) and th origin of gographi faturs (ata- for irh addition of
strophi ation' suh as Grand anyon's ing formd y Noh's Flood), litlts hv tl b ssn
ID is mut aout th history of th univrse ausID intnds to a ig natural sltion arrn
tnt that will hous all antivolutionists'and ausinsid this ig tnt r misundrstandingof tht
popl with diddly inomptil idas (Sott 2001). ost of th ID advo- out a numr of ways t
ats ar old.arth rationists, ut thr ar young-arth rationists who pl struturs' vn s(
ar ativ supporters of ID. Similrly, as do th young-arthrationists,ID 1997; rr 1996).
advoatsallow for volution within ratdkinds, ut thr is no onsnSus
in th litratur of ithr an1pon what a ..kind'' is, th limits of gntivri- ID As SHOLARSHIP. B
ability tht rstrit volution outsid thsratd kinds, or what dtrmins ons' th is tlr isk tl
ths limits. Mihl Bh, a prominnt spoksprsonfor ID, on dsrid stutur that was disor

''sBlk Bo Lhiglr Univr-
;hdy th Fr Press. Dri's
.t to ast itslf as a sin. Th
\ 4 s t r s:o m p h n o m n ai n n a -
l through natural auss. Spifi-
i suh as th atrial f-lagllum
iad as bing too orpl to
sltion.Bh's ook rivd
whil Johnson's ook did not-
of a profssorfrom a sular uni.
lk, this tinrfrom a minstm
or attntin from sholars than
rptd to provid a sintifi argu.
w r l m r s tu n i f o r m l y n g a t i v
,; avalir-Smit1997; Thornhill
mtiian William Dmski pu-
;e, th asi Strutur of ID had
and othr publiations Dmbski
' for th furrdamntalida of ID:
pl to plaind though nat-
n'' for thir xplanation. Invok-
rasing improaility, Dmski
;sand halr as potntial xpla-
sign as th rsidual xplanation.
rg fllow philosophrs and proa.
.ritunsussfulat prsuading i.
1dinformtio,asany utility for
tlson t a|. 1'999; Shanks 24;
)03;Pnnok 7999,2001;' lsrry
[. In ontnt, IntlligrrtDsign dif-
:s rlutan to mak spifi fat
laims about th ag of th arth
ligin of gographi fturs (ata.
's
ing formd y Nah's Flood),
:rsbaus ID intnds to a big
rnd ausinsid this big tnt a
(Sott2001). Most of th ID advo-
] ar young-arth rationists who
do th yong-arth rationists' ID
td kinds, ut thr is no onsnsus
..kind'' is, th limits of gnti vari-
g ratdkinds, or what dtrmins
poksprsonfor ID, on dsribd
m r i c nt i euo lti oisnt 79
himslf as living in God-guidd volution (Bh 2000)' although othr
prorinntID tlroistshav dlrd that ..ID is no find of thisti r,olu-
tion'' (Dmski 1995, 3). In rany ways, Bhe stands apart from thr ID ad-
voats,inluding in his aptan of ommon anstry for living things
( B h 2 0 0 7 ) .
As Philiip Jolrrrsonhas rommndd,ID advo:rtstry to lirrritthmslvs
to thir hif onrn, whilr is whth natur n xplaind solly
through rraturalprosssor whthr intllignt-in prati, suprnatual-
intrvntionis rquird. This ida undrlis ID's or laim: som natural
phnomnaar dsignd y n intllignrathr tlrn havirrg om aout
throughnaturl prosss.Tlr sondary,though sintifiallyritial' laim
is that it is possil to distinguish intlligntly dsignd phnomna from
ompl phnomna of ntur:rl origin. Typially, ut rot alwys, ths ar
iologialphnonrna,vr oftn th sm phnornsltdfor srurirry
y rtion sine.2Th mthodology prposd involvs a onpt asd on
proailityand information thory dvlopd y Dmski.
Dmski hs ()ntnddtht low-proaility vnts of a partiular type
(..spifid'' in advan y sid information) ar assignalto th atgory of
things that hav n intlligntly dsignd. omplx spifid information
(SI)'it is laimd, idntifisa nwly disovrdlass of phnomna that r.
quir th ation of intllignfor thir origins. Harking ak to Thton
and oauthlrs'ontntioninTh strlt of Life,s origi that tl.rorigin of
lifws too ompliatd to plaind y natural us' Dmski uss SI
to idntify a lass of phnomna not only unxplaind ut unxplainal y
naturl'that is, sintifi,prosss.
ihal Bh's ida of irrdui1orrrplity(I) of biologizrlstuturs
is similarly usd to argu against volution. Bh limd in Drin's Blk
Bo (1996) that tain iohrnialprosssor llular strutursWer not
xplinly ntr.rrlsltion.H ontn]dthat som strutllrs' sr-ras
th batrilflagllum,rquir th assrnlyin final Opratingordr of all of
thirmany omponnts for thy an b funtional. By dfinition' any Pr-
ursorof an irrduily ompl strutur is nonfuntional. Bh's viw of
nturlsltiln-though not tht of volutionary iologists-is tht it or-
atson on omponnt of a strutrrrat tim, rathr lik adding harms to
a hamralt.Bh argusthat if natural slti<rn rquirs sltivvalu
for ah addition of ah omponnt of an I strutur' ut all t mpo_
nntshv to ssrldtor thr is ny sltivvaiu, inrmntal
natural sltiorrannot xplain irrduily omplx struturs. This is a
misundrstanding of th pross lf natural sltion,and ritis hav pointd
out a numr of ways tht natural sltion ould oprat to produ om.
pl strutrs'vn so-lld I ons (Thornhill and Ussrv 2000; Dorit
1997..rr 1996\.
IDAs SHOLARSHIP. BausID onfussr'lnsolvdprolmswith unsolval
ons'thr is t[rrisk that ID might disprovd l,an irrduily ompl
strutr
that was disovrdto hav a plar"rsil
naturl planatiorr.Howvr,
80 mri tiultilism

this dos not sm to an atual diffiulty, for ID proponnts an always al- ID artils in th si
low that thy w wrong aout th irrduil omplity of tht strutu doing sin y ID's
and om up with anothr. Ironially, thy sm to ignor strutus that ar not produ (som)n
irrduily omplx, suh s th mammalian middl ar or th avian wrist ural auss' th tho
ompl, ut whih in fat hav a wll-undrstood volutionary history (Gish- .!ha
atual rsarh.
tik 2004). From a sintifi standpoint, of ours, th inaility t rfut I y look lik othr than :
disproving laimd xampls is not a strngth ut a waknss. Sin pro- show, that natural pr
grsssy liminating planations that ar disprovd' not y proving plana- quat to aount for
tions in som solut sns. snario prsntd
Intllignt Dsign proponnts hav produd a larg numr of ooks thory hind ID dis<
sin th apparan of Th str of Life's rigi' th ulk of thm sin thr should no st
te 1996 puliation of Bh's Drlui's Blh Bo and Dmski's 1998 tur. Dmski hims
Th Desig Ifr. ost of ths ooks, howvr, must lassifid as th lak of progrssil
rligious or philosophial in natur rathr than sintifi;it is rvaling that ..d
mploying a vagu
most a pulishd y starian pulishs suh as IntrVarsity Prss, Bakr of thnologial inn<
Book Hous, nd Brazos Prss.his is urious' sin ID's laim to al al systms (Dmsk
to idntify intlligntly dsignd biologicl strutures would imply that its al phnomna will
adhrnts intnd lD to answr qstions that ar ing invstigatdin th will in som way sul
iologial sins.Th pla to do this would in sintifijournals, sp- out that iologial pI
ially in th filds of llular and molular iology, ut hr ID has n fatul.
asnt. vn Bh, an aadmi sintist t a sular univrsity (Lhigh) Although the sin
who has pulishd in mainstram sintifi journals on sintifi topis un- ration sin'thr
rltd to ID, dos not appar to pulishing or attmpting to pulish r- nals, primrily au
sarh that maks a positiv s for ID. Th most prominnt [D dvoats writtn for th gnr
appr to prfr to publish in popular ooks rathr than sintifi journals. tal xption. As a
Dmski has ommntd: ..I'v iust gottn kind of las6 aout sumitting Bh ar ook rviv
things to journals whr you oftn wait two yrs to gt things into print. philosophy or math
And I find I an atually gt th turnaround faster y writing a ook nd uss ID (.$ilkinsanr
gtting th ideas prssdth.y ooks sell wll. I gt a royalty. And th 2003)' Thr is a gro
matrial gts rad mor'' (MMurtri 2001' 8). ritiqu ID on sint
Normally, nw sintifi idas ar first psntd t aadmi onfrns Forrst and Gross 20
and, aftr profssional fdak and som gnral disussion, ar rfindto Young and dis 200,
th point whr artils ar sumittd to sintifi journals, aftr whih thy ussions among ID a
may or may not join th sintifi onsnsus.Aording to an artil in th vr (.g., www.pn
hroicl of Highr dutio, whn askd why h is not uilding his and Shallitt200;Wt
irrduil-lmplityida and th ID model at profssionalmtingsof ll For all th laims t
iologists and iohmists,..Mr. Bh rsponds that h prfrsothr vnus. ntrpris, vn a bri(
.I just don't
think that larg sintifi mtings ar fftiv forums fo pr- no on is applying tl
snting ths idas,' h says'' (MMurtri 2001, 8). stand or plain io
Again parallling ration sin,ID proponnts ontnd that thir viws 2004). Th urdn o
ar systmatially kpt out of sintifi journals austhy rjt th st- thir position is sin
tus quo. Yet unorthodoy is not routinly anishd from puliation (as th tionary biology app
historis of th rption of ndosyrnisisand puntutd quiliria wit- insights and undrst
nss),although unusual laims rquir mor than th usual amount f sup- nw fossil rmains r
port and tak long to stalish.A mor plausil rson for th darth o{ dvlopmntal biolog

lr ID proponntsan always al-
l omplit of tht strutur
m to ignor struturs that ar
r m i d d l a r o r h a v i a n w i s t
tood volutionry history (Gish-
Irs,th inaility to rfut I y
'h ut a waknss. Sin pro-
;provd'not y proving xplana-
ud a larg numr of ooks
's
rigi, th ulk of thm srn
lh Bo and Dmski's 1998
, howvr,must lassifid as
ran sintifi;it is valing that
uh s IntrVarsity Prss' Bakr
ious, sin ID's laim to l
struturswould imply that its
rat ar ing invstigatd in th
rld in sintifijournals, sp-
r ilogy, ut hr ID has bn
rt a sular univrsity (Lhigh)
j o u r n a l so n s i n t i f i t o p i s u n -
ring or attmptingto pulish r-
1 most prominnt ID advoats
(s rathr than sintifi iournals.
kind of las6 aout submitting
/ o y a st o g t t h i n g s i n t o p r i n t .
nd fstr y writing a ook and
sll wll. I gt a royalty. And th
,8).
rsntdt dmi onfns Forrstand Gross 2004; Shanks 20041K. R. Millr 7999iK. B. illr 2003;
gnraldisussion,r rfind to
intifi iournals, aftr whih thy
s.Aording to an artil in th
kd why h is not building his
ll at profssionalmtings of ll
onds that h prfrsothr vnus.
-ingsar fftiv forums for pr-
: 0 0 18, ) .
Oponntsontnd that thir viws
r r n a l s a u s r h y r j t t h s t a -
anishdfrom puliation (as th
is and puntuatd quilibria wit-
, r t h a n t h u s u a l a m o u n t o f s u p -
plausiblrason for th darth of
meri ntieuolutioism 81
ID artils in th sintifi litratur is th onstraints that ar imposd on
doing sin by ID's foundational laim, whih is that natural auss an-
not produ (sm)natural phnomena. Sin sin xplains through nat-
ural auss,th thory tht undrlis ID maks it rathr diffiult to do any
.lfhat
atual rsarh. would ID rsarh in iology (rathr than thology)
look lik othr than attmpting to show, as ration sinistsattmptd to
show, that natural prossssuh as thos involvd in volution ar inad-
quat to aount for th origins and histoy of lif? Thr is no historial
snarioprsntdy ID advoats' so thr ar no fat laims to tst. If th
thory hind ID disouragssintifi ativity and thr ar no fat laims,
thr should no surpris that thr ar no artils in th sintifi litra-
tur. Dmski himslf has ommntd, prhaps with som frustration, on
th lak of progrss in produing a sintifi foundation for ID and suggsts
..dsign-thorti
mployinga vagu framwork'' suh as omparing a modl
of thnologil innovation dvlopd y Russian nginrs with iologi.
al systms (Dmski 2002). Prsumably Dmski ntiipats that iologi-
al phnomna will rsml nginrd systms (dsignd),and that this
will in som Way support ID, in dfian of opious litratur that points
out that iologial phnomn ar organi in omposition rathr than arti-
fatual.
Although th sintifi ommunity has paid mor attntion to ID than to
ration sin, thr is not an xtnsiv anti.ID litratur in sintifi jour-
nls' primarily aus ID has n promotd almost xlusivly in ooks
writtn for th gnral puli, Dmski's Th Dsig Ifrce bing a no-
tal ption. As a rsult, most of th pulishd ritiqus of Dmbski and
Bh ar ook rviws, as itd arlir, although thr ar a fw artils in
philosophy or mathmatis journals-rarly in iology journals-that dis-
uss ID (Wilkins and lsrry 2001; Shanks 1999; Sor 2002; Pnnok
2003). Thr is a growing numr of books ing pulishd that spifially
ritiquID on sintifiand/or philosophia1grounds (Pnnok 1999,2I;
Young nd dis 204). On an find thnial analyss and vn livly dis-
ussions among ID advoats and ritis taking pla on th Intrnt, how-
vr (.g.,www.pandasthum.org; Matzk 2004; Gishlik 2003; lsbrry
ad Shallitt 2003; \Win 2002).
For all th laims that ID advoats mak for th sintifi validity of thir
ntrpris,vn a rif omputerized sarh of sintifi journals rvals that
no on is applying th onpts of SI or irrduil omplxity to undr.
stand o plain iologial phnomna (Gilhrist 1'9971Forrst and Gross
2004|. Th urdn of proof rmains on th poponnts to dmonstrt that
thir position is sintifilly valid and usful. Manwhil, rsarh in volu-
tionary iology appars to in a partiulaly produtiv priod, with nw
insights and undrstandings ing provided not only from th disovry of
nw fossil rmains ut also from molular iology, spially as pplid to
d v l o p m n t l i o I o g y .
82 Amri Atiuoltionism
.William
ULURAL RNWAL. Dmski has writtn, ..Intllignt dsign is minimalist dfinition f.
thr things: sintifi rsarh program that invstigats th ffts of in- nonthlss ontains }
tllignt auss;an intlltual mvmnt that hallngsDarwinism and its Intllignt Dsign rt
naturalisti lgav; and a wav of undrstanding divin ation'' (Dmski ural auss, known as r
1999,1'3). Although th sintifi rsarh program of ID has n unsu- ial naturalism. Philos
ssful, th ulk of th ffort xrtd y th ID movmnt is dirtd toward opposd to hristian t
te ultrI rLuIomponnt hintd at in Dmbski's sond laus: th f- ould roadndto l
fort to rpla ..naturalism''-in th ID worldviw' tantamount to athism- uralism ould aan
with hristian thism. As suggstd y oth th inadquay of its sintifi is privd as its stror
rord and th titls of its organizing onfrns, th ID movmnt is in fat (orland 19941Scott
primarily onrnd with th hristian rligion and its onflit with matrial- God (or ..intllign
ist philosophy-th philosophy that thr ar only matrial (mttr nd n- jo, is a ky gol of ID
rgy) phnomna in th univrs; thr is no suprnatural rality. histi sin is
To dfat sularism in modrn soity, th ID movmnt attaks sin ene and oprtion si
through volution. Th attak, signifiantly nough, is n.Jt primarily through mal,'' mthodologiall
sintifi vidn ut through philosophial and vn statistil argumnts' sintifi planations.
though ths argumnts ar flt to thin y philosophrs and proaility singular, unrpatal
thorists. To undrstand how th ID advoats rason rquirs a short di- aout whih supposd
grssion int th natrrr of sin as it is pratid y sintists today. triution of ausality t
Although philosophrs of sinradily argu aout th dfinition of si- ration sin.a
n and vn whether sin an lanly distinguishd from othr wys volution, thn, is a
of knowing, ths argumnts ar not vry important to th tual ntrprise aus of its matrialis
of sin as it is pratid y sintists' who tnd not to think muh aout mthodologial and ph
th philosophil nitis. It would rlativly asy to gt sintists (and as a thfat to hristia
proaly vn philosophrs of sin)to agr on a oupl of asi lmnts larly is on of th m.
that haratriz sin, vn if suh lmnts did not fully dmarat si- Drui on ril lvas tl
n from othr pistmologis. Sin is an ffort to undrstand th natural Johnson romots a l
or matrial world-th world of mattr and nrgy and thir intration. It takd as a mans t)
is, therfor,not apal of xplaining th phnomnon of rligion: the pur- rstalishmnt of l
portd xistn of a transndntal rality y dfinition not limitd to th mony of thisti sin
matrial univrs. Sin is in this sns a limitd ndavor: it is limitd to also n instrumntal
matrial phnomna. ommitmnt to any fa
Sintists would also agr that th ssn of sin is testing xpiana- a wa of avoiding th
tions against th natural world; an xplanation asd only on inspiration. of The str of Lif'.
rvlation, rativity' o imagination would not a sintifi planation, son onntrats on th
although a sintifi xplanation might gin that way. Only after onfirma- natural sltion)to x1
tion through tsting would it om a sintifi planation. Th famous
story of how Fridrih Kkul6 am to disovr th irular struturof n- So th qustion is: ..[
zene aftet draming of a snak hasing its tail illustrts this wll: it took you now s full.fld
vrifiation-tsting th xplanation against th natural world-to stalish important thing"-tl
a sintifi planation. Kkl6 did not stop with th dram. It is also likly Gt th Bil nd th
that sintists would agr that a major omponnt of tsting is holding r- not Want to rais th
tain variabls onstant in ordr to masur th fft <rfothrs. Of ours, mnt in suh a way tJ
sintistsand philosophrs of sin an and do dt othr fatursthat in a way that tnds tt
may or my not dfin sin, suh as th various vrsins of falsifiation trating on, "Do you r
and thir appliation, ut th dsription prsntdhr would suffi as a it on its own?'' nd r

i writtn' ..lntlligntdsign is
lt invstigarsth ffts of in-
t hallngsDrwinism nd its
.ding divin ation'' (Dmski
logrmof ID has n unsu-
D movmnt is dirtd toward
)mbski'ssond laus:th f-
lviw,trrntmountto athisn-l-
th inadquay <lfits sintifi
lsr th ID rovrntis in fat
'n and its onflit with mtrial.
only nratrial(rnattrand n-
suprnaturalrality.
, I D n l o v l n t t t a k s s i n
nough,is not primarily through
and vn sttistialargtrmnts'
'y philsophrsand proaility
lts r2tsonrquis a short di-
tidby sintiststody.
rgu aout tl-rdfinition of si-
.distinguishdfrom othr ways
I p o r t n t o t h a t t | a l n t p i s
l tnd not tl think muh aout
ivly asy to gt sintists (arrd
'on a oupl of basi lmnts
n r s d i d n o r [ u l l y d m a r a t s i -
ffortto undrstandth natural
nrgy and thir intration. It
hnomnonof religion: th pur-
y dfinition not limitd to th
imitd ndvor:it is limitd to
; of sin is tsting plana-
t i o n a s d t r l v n i n s p i r a t i o n ,
not a sintifi planation,
l that wy. only ftr onfirma.
:ntifixplanation. Th famous
/r th irular strutur of n-
til illustrtsthis wll: it took
t natural world-to stalish
with th drm. It is also likly
ponntof tsting is holding r-
th fft of otlrrs. of ous.
Ld do dat othr faturs that
vrious vrsions of flsifiation
.sntlhr would suffi as a
mericn tiuolutionism 38
minimalist dfinition for virtually all sintists.But this miniml dfinition
nonthlssontains hratrististhat ar opposd by ID advoats.
IntlligntDsign rationistslivthat th rstrition of sien to nat-
ural auss,known aS nthodologi1 naturalism, lads popl to philosoph-
ial naturalism.Philosophial naturlism is, of ous' th nmy ausit is
opposd to Christian thism' If a powrful ultural ntrpris lik sin
ould roadndto inlud God's intrvntion' and if mthodologial nat-
uralism ould aandond, thn philosophial naturalism would los what
is privdas its strongstsupport. Th stalishmntof ..thistisin''
(orland 1994; Scott 1998), whih prmits th invotion of th hand of
Gd (or ..intllign'')whn natural xplanations ar indquat for th
jo, is a ky goal of ID ationism.
Thisti sin is asd on th onpt of a dihotory twn origi sci-
e and opertion sitl, th lattr ing ID proponnts'trm for th ..nor-
mal''' mthodologially matrialisti sin usd in th v:rst mjority of
sintifiplanations. origin sin is dfind s th sin usd to plain
singular, unrpatabl vnts (th origin of lif, for xample)' hypothss
about whih supposdly ar untstal and thus r.rnfalsifiabl.Thrfore' at-
triutionof ausality to God is aptalin origin sin,though not in op_
ration sin.a
volution, thn, is a stalking hors for an attak upon sin, whih -
aus of its matrialisti as (ID rationists do not distinguish twn
mthodologialand philosophil naturalism in any pratial sns)is sn
as a thrat ro hristiarr thisnr. Th ID advoat who nraks ths links most
larly is on of th most prolifi ID authors, Phillip Johnson, whos 1991
Drluilt o ril was th first ID puliation to attrat wid puli noti.
..wdg'''
Johnson promots a stratgyhe alls th whrin volution is at-
takd as a mans to hallng naturalism, thus paving th way for th
rstalishmntof histin snsiiiity in Amrian ultu and the hg-
mony of thisti sin (Johnson 1997i Forrest and Gross 2004). H has
also n instrumntai in promoting a puii Prsnttionof ID that voids
ommitmntto any fatual laims, suh as a youn8 or old ag of th arth' as
a way f avoiding the pitfalls of ration sin.Instad,as did th athos
of he ster of Life's rigi and f Pds nd Pople for him,
John-
son onntratson th supposd inadquay of natural auss (speially
nturalsltion)to plain th origin arrd divrsity of living things:
So th qustion is: ..How to win?'' hat's whn I bgan to dvlop what
you now s full-fldgdin th ..wdge'' stratgy:..Stik with th most
important thing''-th mhanism and th uilding up of information.
Gt th Bil nd th Book of Gnsis out of th date ausyou do
not Want to rais th so-alld Bil-sindihotomy. Phrs th argu.
mnt in sh a way that you an gt it hrd in th sular aadmy and
in a way that tnds to unify th rligious dissntrs.That mans onn-
trating on, ..Do you nd rator to do th rating' or an natur do
it on its own?'' and rfusing to gt sidtrakd onto othr issus,whih
84 mricn Antiuolutionisnl

popl ar always trying to do. Thy'll ask, ..W.hat do you think of of th univrs at one til
Noah's flood?,' or somthing lik that. Nvr it on suh qustions - ar ford to support tht
aus thy'll lead you into a rrklsswastland and you'll nv gt out idn gainst volutin
of it. (Kushinr 2000, 40) a frqurrtlyrworkd lil
suh as gps in th foss
Baus it dos not prsnt an altrnativemodl to volution, and eus law of rhrmodynamis
of its rlutan to mk fat laims, th ontnt of ID thus dvolvs to th imossiility of dvlop
ritration of argumntslong ago raisd y th young-arthrationists:gaps dom variation and natu
in th fossil rord, th inadquay of natural seltionto produ volution- living organisms'' (IR
..prolms'' of th amrian
ary hang, th plosion, and s on. As is th i d n a g i n s t v o l u t i o n
..<rntrivd
s with Y, ID appars to rting a dualism''' to quot who ar puzzled that un
L u. rkss, in whih if volution an shown to untru, [D is apparan of amrian
thus stblishd. sltiv mhanisms i
thoy o rsons to rj
uidence gist euolution Tahing th ..vidn
Aftr th deurds dision struk down qual-tinr lws, Wndll Bird, who mnt laus prolm gl
had argud Louisiana's position fore th Suprm ourt, gamly advisd stratgy is thrfor fav
his fllow rationists that ration sin might still lgally taught if it who ar oprating in th(
wr taught with a sular purpos (IR 1987). His prsptiv was not for xampl, has om tt
shard y judgs in distrit and appals ourts who intrpretd d,rdsas ID ut rathr to tah th
not distinguishing twn rtion sin and rationtsl:.drds as
n itd y fdral distrit nd appals ourts in disionsthat hav struk
down attmpts by tahrs in lllinois and alifornia to bring ration sin Th Futur of Antit
into th lassroom.
A dissent to do,rds,writtn y Justi Salia and signd y Justi Rhn- Sott and Branh hav t
quist, suggstdan altrnat approah that was quit ompatil with xtant th priods of Amri
..Th popl of Louisiana, inluding thos who ar hist- ontrovrsy today as thr
ration sin:
ian fundamntalists,ar quit ntitld' as a sulr mttr' to hav whatvr 282). t any givn tim
sintifi vidn thr may against vlutin prsntdin thir shools, o f t h r a t i o n i s tp s i t i o
iust as r. Sops ws ntitled to prsnt whatvr sintifi evidnth 1. volution is wak o
was for it,, (dturds v' guillrd,64). Th ICR sizd upon this immdi- aandoning in inr
atly aftr te drds dision was pulishd. hoing Salia, it nour- 2. volution is inhr
..In th mantim' ..
agd tahrs to tah th vidn against volution: 3. It is only fair to
shool boards and tahs should strongly nouragd at last to strss o r l n t l I i g n tD s i g
th sintifi vidns and argumnts against voltion in thir lasss (not good pdagogy to ..
iust ar8umnts agains som proPosd volutionay mlranism,ut against ..waknsss''or ..
volution pr s), vn if thy don't wish to rogniz ths ils vidnsand did.''
argumnts for ration (not nssarilyas arglllnts for a partiular dat of
ation' ut for ration pr s)'' (IR 1987). Ths thr thms w
Th link twnation sinand ..vidnagainst volution'' is asy trovrsy in th futu. I
to find. Proponnts of rtion sin ontnd that thr ar only two possi- studis of antivolutiln
ilitis: rationism' onsisting of thir spifi vrsion of hristian spial stongsr motivation for
ration, and volution, whih also inluds all th otlrr origin storis from 1980s nd arly |990s,
othr rligions and ultus.Thrfor, to thm it is logial to infr that vi- rtionism in favor of t
dn gainst volution is vidnfor rtionism. Pratiill onsidtions ral puIi and, spi
ar aiso involvd: thr r no rdil data that suPport th spial ation Both ration sin

sk, "'S7hat do you think of
r it on suh qustions -
land and yu'll nvr gt out
rodl to vlution, and aus
tnt f ID thus dvolvs to th
young.athrationists:gaps
sltionto produ volution-
xplosion' and so on. As is th
..ontrivd dualism,'' to quot
rn shown to untru, ID is
rl-tim laws,.tJ7ndllBird, who
iuprm ourt, gamly advisd
night still lgally taught if it
987). is prsptiv was not
ts who intprtd drds as
and rationism; drds has
rts in disions that hav struk
i f o r n i t o r i n g r
alia and signd y Justi Rhn-
rasquit ompatil with xtant
inluding thos who ar Christ-
ular mattr, to hav whatvr
tion prsntdin thir shools,
'hatvrsintifi vidn thr
IR sizd upon this immdi.
;hd. hoing Salia' it nour-
..In th mantim'
t volution:
gly nouragdat last to Strss
st volution in thir lasss (not
l t i o n a r ym h a n i s m , u t a g a i n s t
rogniz ths as vidns and
'gumntsfor a partiular dat of
7\.
idn against volution'' is asy
nd that thr ar only two possi-
;ifi vrsion of Christian spial
all th othr origin storis from
lm it is logial to infr that vi.
tionism.Pratial onsidrations
that support th spial ration
Amri Atieuolutionism 85
of th univrs at on tim, in its prsnt form, so ration sin advoats
ar ford to support thir viw with a ngativ argmnt' that of allgd v-
idngainst volution. Th ontnt of ration sin primarily onsists of
a frquntly rworkd litany of argumnts prsntd as disproving volution,
suh as gaps in th fossil rord, th supposd inompatiility of th sond
law of thrmodynamis with a gradually inrasing omplity of lif' th
impossiility of dvloping phylum-lvl ody-plan diffrns through ran-
dom variation and natual sltion, and th ..vast information ontnt of
living organisms'' (IR 1'987).It prhaps gos without saying that suh ..v-
idnagainst volution,' gneratslank looks from volutionary ilogists,
who ar puzz|d that unsolvd aras in sin suh as th origin of lif, th
apparanof amrian ody plans, or th rlativ rol of sltivand non-
sltivmhanisms in volution ar onsidrd waknsss in volutionary
theory or rasons to rjt th infrn of ommon anstry.
Tahing th ..vidn against volution'' avoids th ovious stalish-
mnt laus prolm gnratd y ID, whih is ..\/hois th dsignr?'' Th
stratgyis thrfor favord among th mor sophistiatd ntivolutionists
who ar oprating in th arly twnty-first ntury. The Disovry Institut,
for ampl, has om to urg shool oards not to rquir tahrs to tah
ID ut rathr to tah th ..strnsthsnd waknsss',of volution.
t i o n s i n
Th Futur of Antivolutionism
Sott and Branh hav rfrrd to thr thms that hav run throughout th
thr priods of Amrian antivolutionism and ontin to illustrat th
ontrovrsytoday as th ..pillars of rationism'' (Sott and Branh 200'
282). t any givn tim in this history, ths thr argumnts hav n part
of th rationist position:
1. volution is wk or unsupportd sin that sintists ar
aandoning in inrasingnumrs.
2. volution is inhrntly antirligious.
3. It is only fair to ..alan''th tahing of volution with rationism
or IntlligntDsign. A vaiant of th ..fairnss''argumnt is that it is
good pdagogy to ..alan'' th tahing of volution with
..waknsss'' ..vidn
or against'' volution ..and lt th hildrn
did.''
Ths thr thms will ontinu to shap th rationism,/volutionon-
.W
trovrsyin th futur. an illustrat thm y xamining som rnt as
studis of antivlutionism. Although rligion ovrwhlms all ls as th
stfongstmotivation for antivolutionism' aus of lgal disions of th
1980sand arly 1990s, rationists hav downplayd th sond pillar of
rationismin favor of th first and third, at last whn addrssing th gn-
ral puli and, spially, puli offiials suh as shool-oard mmrs.
Both ration sin and ID proponnts hav hammrd away at th ida of
86 A m eri t iuolutioism

volution as wak sin and hav strssdth importan of fairnss' sp- Indd, onsrvativso1
ially in trms of tahing ..vidn against volution.'' that thy did not look k
Anothr trnd has n th ..morphing'' of Y to ID to ..vidnagainst t h s t a n d a r d s( u n n i n g
volution.,' In a numr of ommunitis, popl who ar young-arth r- D u r i n g h s p r i n ga n
ationists bgan to idntify thmslvs as ID supportrs' thus avoiding th trs to th ditors argur
stigma of ration sin, whih is at st onsidrd a minority viw, if not Inlusion of volution .
a fring prsptiv'in oth rligion and sin.IntlligntDsign, on th othr vlop qustions for th s
hand, is lss wll known to th gnral publi, and aus of th vaguenss in thos xams' it ws
with whih it is prsntd (making no laims aut th ag of th arth, fr rooms. Tahers and s
xampl), it appars lss marginal than ration sin.Latr' as a ommu- hav a prominnt pla
nity that is dating volution duation oms fanriliar with ID and its standards tht ws sub
sintifi strility and duious onstitutionality, th antivolutionists oftn th National Sind
rtrat to a fallak position of ..vidnagainst volulion,'' usually por- Aadmy of Sins,ar
traying it as a ompromis that should aptal to ll. Morphing of Sin (AAAS) Bnh
this sort ourrd during th wll-puliizd Kansas stat sin standards bn influntial in shap
ontrovrsy of 1999. NSS and th Bnhma
inlud it throughout th
In midsummer 1999
As sUDY: KANSAS
standards ommitt'sd
As a rsult of duation rform movmnts that bgan in th 1980s' most l r g | y n w r i t t n b y
Amrian stats hav stalished duational standards for history, math- ation for Mid.Amria,
matis, sin, and other aadmi sujts. Aftr th fdral No hild Lft of St. Louis. Arams's st
Bhind At of 2001 mandatd priodi high-staks tsts for studnts, stat or apabl th wor
standards took on vn mor importan. Tst topis would asd on th ludd muh aout how
standards; if volution wr inludd in th standards, thn it would in- puli harings oftn ml
ludd in th tsts. Stats that did not alrady hav standards or framworks draft of th standards t
gan to dvlop thm in th 1990s; th vast majority of thm inlud volu- ompomisdraft thtit
tion, though in varying dgrs of omptny (Lrnr 2000). any stats draft ompromisd btl
that had ignord volution prviously now rquir that th topi taught. thos who wntd the ,A
A good dal of th antivolution ativity of th past fw yars has n di- is suh s th ig ang
rtd t th stalishmnt of stat sin duation standards: rationists standards and adding s
oppos th inlusion of volution in th standards,or try to gt ID addd, or of sien.'
try to gt ..vidn against volution'' inludd. Muh as th rturn of vo. A loud hu and ry r
lution to th urriulum in th mid-1960s gnratda rationist aklash' paprs' tlvision, and r
so also has th standards-asd rquirmnt that volution taught gnr- tlvision talk shows, ;
atd its own aklash. Kanss Wizrd of z ir
Kansas undrtook to writ sin duation standards in 1999. 27- Kansas suffrd at th l
prson ommitt of mastr tahrs and sintists was appointd y th th nt shool-oard l
Kansas Dpartmnt of duation and th Kansas Board of duation to draft at majority swiftly abar
standards that thraftr wnt out for puli ommnt' wr rvisd, and adoptd th final draft 1
wr ultimatly sumittd to th oard of duation for approval. This pro- th nw oard took offi<
dur was prolongd, ontntious, and ittrly fought. A onsrvativ lo Th Kansas ontrov
of fiv shool-oard mmers had om to powr in th 1996 state shool- rationism, th lim tt
oard ltion and lashd with th fiv modrat shool-board mmrs Proaly baus th
ovr many issus. Whn th sin duation standards am up for onsid- arth rtionists' and t
ration. it was an asy ordition that volution wld a ontntiousissu. ludd in th sinsta

e importanof fairnss,sp-
rolution."
/ t o I D t o ' . v i d n a g a i n s t
lpl who ar young-arth r-
supportrs'thus avoiding th
rsidrda minority viw' if not
. IntlligntDsign, on th othr
, and ausof th vagunss
aout th ag of th arth, for
on sin.Latr, as a ommu-
oms fanriliar with ID and its
ity' th antivolutionists oftn
3ainstvolution,'' usually por-
'ptablto all' orphing of
Kansas stat sin standards
tlrat bgan in th 1980s' most
l standrds for history, math-
Aftr th fdral No hild Lft
-staks tsts for studnts, stat
st toPis would asd on th
stndards, tlrn it would b in-
.hav standardsor framworks
rnajority of thm inld volu-
ry (Lrnr 2000). Many stats
quir that th topi b taught.
th past fw yars has n di-
l u t i o n s t a n d a r d s : r a t i o n i s t s
lards, or try to gt ID addd, or
Jd' uh as th rturn of vo-
nratda rationist bklash,
that volution taught gnr-
tion standards tn 1999. ^ 27-
intistswas appointd y th
rsasBoard of duation to draft
i ommnt, wr rvisd, and
luationfor approval. This pro-
ly fought' A onsrvativ lo
powr in the 1'996 stat shool-
Lodratshool-oard mmrs
n standardsam uP for onsid.
o n w o u l d a o n t n t i o u si s s
mri Atiuolutionism 87
Indd,onsrvativson th oard lt it known rathr arly in th pross
that thy did not look kindly on th inlr-rsionof volution in th first daft of
the standards(unningham 1999).
During th spring and summr of 1999 Kansas nwspaprs wr full of lt-
trsto th ditors rguing for and against volution in th sinstandards.
Inlusion of volution was impotant: th standards would b usd to d-
vlop qustions for th stat gr:rdutionxms. If volution Was not inludd
in thos xams) it was a saf t that it would not taught in th lass-
rooms. Tahrs and sintists thrfor Wr vry onrnd that volution
hav a prominnt pla in th sin edutiorr standards. The draft of th
standardsthat ws sumittd fo puli ommnt in April 1999 paral|e|ed
th National Sinduation Standards (Nss), produd y th Nationl
Aadmy of Sins, and th Arnrian Assoiation for th Advanmnt of
Sin (AAAS) Bnhmarks for Sin Litray, two doumnts that hav
n influntial in shaping sin standards aoss th ountry. Both th
NSS and th Bnhmrks trat volution as an importurntsintifiid and
inlud it througlrout th sien urriulum.
In midsummer 1999 shool-oard mmr Stv Arams ountrd th
standardsommitt's draft with ir rplamnt standards doumnt that had
larglvn writtn y Tom Willis, th had of th ration Sin Assoi-
r i o n f o r i d - n l i a ' y o u n g . r t h r r i o n i s to r g n i z a t i o no p r a t i n gr l u t
of St. Louis. Aranrs's standards inldd svral provisions that any sintist
or apal tahr would onsidr highly qustional and of ours in-
luddmuh aout lrow volutiorrwas unrlial sin.Aftr a summr of
puli harings oftn markd y ittr ontrovrsy' the oard took th final
draft of th standards ommitt nd th Arams standards and produd a
ompromisdrft that it aptdas th stat standardsin August 1999. This
draft ompromised twen thos who wantd volution to e taught ad
thoswho wantd th Abrams draft y rmovirrgvolution nd rlatd top-
is suh as th ig ng, rrtinntal drift, i.rndradiomtri dating from th
standardsand adding som languag from Arams's draft aout th natur
of sin.
A loud hu nd ry nsud,not only in Kansas' ut also in national nws-
paprs'tlvision' nd radio. Kansas irmth butt of joks ln iat-night
tlvision talk shws, and ditorial artoonists had a fild day with th
Kansas Wizrd lf oz tonogrphy of Dorothy and Toto. Th ridiul that
Kansas suffrd at th hands of th national prss stung suffiintly tht in
th nt shl_oadltion th onsrvativslost thr sats.Th mod-
atmajoity swiftlv andond th ompromis standards and on a 7- vote
adoptd th final draft prdud y th standards-writing ommitt whn
th nw board took offi in Fruary 2001.
Th Kansas ontrovrsy illustirtsquit larly th first of th pillars of
rationism'th lim that volutiorris wak or inadqiratsintifi thory.
Proaly aus th most ativ antivolutionists in Kansas Wr young-
arthrtionists,and ausration sin had no han of ing in-
. luddin th sinstandards,th third, or fairnss,pillar did not ntr into
388 meri Atieuolutionism

th ontrovsy.Howvr' tim nrarhson, nd shool boards hang om- pulishr offrings,1

psition: although in th 2000 ltion modrats prvaild, in 2002 th ..sin
provid any
oard on again was dadlokd with a 5_5 ratio of onsrvativsto mod- aldwll first popor
fats. Th sin duation standards wr lft alon during tlrat trm, ut ooks (Rosn 2003t
th prsn of a 64 onsrvtiv majority aftr th 2004 ltions mant psal to promot ta
that th sin standards rvision that took pla in 2005 was highly on- rommndd a vid
tntious. rationist-orintd shool board mmrs Supportd standads JonathzrnWlls's bo
that did not dirtly all for th tahing of rirtion sin or ID, ut for th vido, Ulokig th
'h
tahing of lD_inspird ritiisms of volutior.r. standards wr passd in YC diatri y Jon
Novmr of 2005 (..Antivolution Standards Adoptd in Kansas'' 2005) l u t i l ( 1 9 9 9 ) .A u i r
and wr widly ritiizd y loal tahrs and sintistsnd national si- !itnsss, Lif: ot
n and duational organiztions, inluding th AAAS and th National towr Bil and Trzr
Aadmy of Sins(..posing t Flaws of th Kansas SinStandards'' Instrutionirllnat
2006). Th nt yar, howvr' Was an ltion yar, and two onsrvativ ommunitis, must l
oard mmrs lost thir sats (..Th Pndulum Swings in Kansas'' 206).In tors. To giv thir
ear| 207 th ronstitutd, mr modrat shool oard rsindd th from loal univrsit
2005 standards nd adoptd standards that nithr dnigratdvolution nor mitt rportd to th
inludd som form of rtionism (..volution Rturns to Kansas'' 2007). Rosvill urriulu
Kansas is also an mpl of th morphing of young.arthtionistsinto sin. In Sptl
ID rationists. A numr of antivolutionists who ativly opposd te 1999 th valuation
Kansas standards hav ast thmslvsas ID supportrs.John alvrt is a whthr or not to us
good xampl: h organizd th IntlligntDsign Ntwork (IDNI), an or. might possily rsul
ganization tht hs sponsord arrnual onfrnspromoting ID in Kansas thy had n rjt
and in som othr stats.lvrt hs om a iruit ridr for ID, traviirrg Crationists on th
to ohio, Nrv xio, Gorgia, Wst Virginia, nd Montana to promot i. volution rsotl
Th Disovry lnstitut' howvr, has om to disourag th promotion of als ritiizing vllu
..vidn against
ID in th lassroom in favor of tahing volution.'' Th tahrs ould not
Disovry Institut has promotd this approah sine te 22 Ohio sin studnts ould go to
duation standards ontrovrsy' during th ontrrtious2005 rvision of th protstd that thv h
Kansas stndards.and lswhr' support th uriul
want to wast mon
unrlatd to th n
As sUDY: RoSEV|LL' ALlFoRNIA
sor ntrs wr l
Although stat o:rrd of dation ontrovrsisnd antivolutionism lgis- ldwll nt att
lation tnd to gnratmor Prss ovrag'th majority of arrtivoiutionisrn lr alld Quality Si
inidnts our t th loal shool-oard lvl' A ontrovrsY in Rosvill, ..strngts
irrg of an<
alifornia, similarly illustratsth pillars of rtionism, as wll as th mor- to poliis that hv
phing of Y into ID into vidnagainst volution. QS ontinud ovr
Rosvill is a ommunity of approimatly 92,000 loatd 20 rnils north- My 10, 2004, in v
ast of Sarmnto in north nrrl alifornia. A sustantial onsrvativ tw hours. A ptiti
Christin ommunity thr tnds to lt shool-oard mmrs who shr opposing th poliy
..ultur-war''ontrovrsisin Rosvill
thir viws. Thr hav n many ppl who tstifid
ovr th past svral yars' inluding s duatin as wll as vllution. Th In Jun 2004 th
most rnt volution ontrovrsy gan in th sr'rmmrof 2003 whn high- rtionism in Ros
shool iology txtbooks wr ing rviwd for adoption. A loal itizn, haps aus of or
Larry ldwll, omplaind that th ttooks, all stndd ommrial arguing ov rirti

and shool oards hang om-
ldratsprvaild, in 2002 th
5 ratio of nsrvativsto mod-
: llt alon during that trm, ut
. ftr th 2004 ltions mant
< pla in 2005 was highly on.
l mmrs supportd standards
rtionsinor ID, but for th
ln. Th standardswr passd in
rrds Adoptd in Kansas'' 2005)
; and sintistsand national si_
ing t AAAS and th National
lf th Kansas SinStandards''
j t i o n y a r ' n d t w o o n s r v t i v
lum Swingsin Kansas" 2006). In
:t shool oard rsindd th
nithrdnigratdvolution nor
:ion Rturns to Knss'' 2007).
g of young-rth r t i o n i s t si n t o
ts who ativly opposd th 1'999
ID supportrs.John alvrt is a
Dsign Ntwork (IDNI), an or-
rnspromoting ID in Kansas
rr iruit ridr for lD, travling
i n i e ,a n d o n t n a t o p r o m o t i t .
to disouagth promotion of
vidnagainst volution.'' Th
ah sin the 202 ohio sin
onttious2005 rvision rfth
)RNIA
,rsis;rndantivolutior-rismlgis-
:,th majority of antivoltionism
lvl. A ontrovrsy in Rosvill,
lf rationism,as wll aS th mor-
volution.
ly 92,000 lotd 20 mils north.
o n i . A s u s t a n t i l o n s r v a t i v
shool-oardmms who shar
r-war''ontrovrsisin Rosvill
duationas wll as volution. Th
n th summr of 2003 whn high-
wd for adoption. A loal itizn,
:tooks,all standard ommrial
A m ri Ati uolu titli sm 189
pulishroffings,Prsntd on-siddtratmntof volution nd did not
provid ny ..sintifi altrntivs.'' Aordirrg t<l a nwspapr aount'
aldwll first proposd tht ID e inludd in th urriulum nd in th tt-
books (Rosn 2003a). Rathr quikly, howvr, aldwll harrgd his pro-
posal to promot tahing ..idasthat ountr volution'' (Rosn 2003). H
rommndd a vidotap promotd y th Disovry Institut asd on
Jonathn!ells'sook los of uolutio (22)plus anothr lD-proroting
video, Ulokig th ster oi Life (2002). Arrlthr itizn sumittd a
Y diatri y Jonathan Sarfati at Answrs in Grrsis cal|ed Rfutig uo-
Iution(1999). A uiquitous ration sinook pulishd y th Jhovah's
Witnsss, Life: Ho Did It Get Hr? B uolutio lr retioi (Wth-
twr Bil nd Tat Sorty 1985) was also propos1.
Instrutionalrnatrialsproposd for lassrootn us in Rosvill, :rsin lros
ommunitis'lllst rviwd y a ommitt of tahrs and drninistra-
tors. T giv thir opinion mor rdiility, th tahrs askd sintists
fom loal univrsitis to rviw aldwll,s suggstdmatrials. T.h om-
mittrportd to th shool oard that th matrialswr unsuital for th
Rosvill urriulunr and, rfltingtlr sintists' valuations, sustandard
sin.In Sptrrrr2003 th ord, stymid y th unitd fronr prsntd
by th valuation ommitt, votd to lt th individual shools did
whthror not t use th matrials.Thrsonsidrdthis an nd run that
might possily rsult in th use of th ooks in th lassroom' vn though
thyhad n rjtdy th valuirtionommitt'
rtionistson th oard lso rommndd tlrt shlol lir:rrisst up
evolutionrsour ntrsthat would hav ooks, vidos, and othr matri-
als ritiizing volution. Th ord mmrs apparntly rasond that if
tahrsould not omplld to tah ..vidnagainstvolution,'' at last
studntsould go to th lirary to gt antivolution information. Lirarians
protstdtht th hd littl nough spa in thir liris fr matrials tht
spportth r.rrriulum-th linritd purpos of a shool lirary; thy did not
wnt to wast mony or spa on what thy onsidrd a suprf-lttous issu
unrlatdto th nds of tahrs for urriular support. Th volution r-
sourntrsWr nvr stalishd.
aldwll nt attllptdto hav th oard pass il poliy h draftd,whih
halld Quality Sindr-ration(Qs). It would hav rquird th th-
ing of ..strngthsnd waknsss.'of volution- hoi of wording similar
to poliisthat hv appard in som othr omnrunitis.ontovrsy ovr
QS ontinud ovr th wintr and ulminatd in a ontntious mting on
May 10, 204, in whih mmunity mmrs datd th poliy for ovr
two hours. A ptition signd y 28 of te 2 sin thrs in th distit
opposingth poliy was suritrdto th oard. In addition, tlr rrrajorityof
poplwho tstifidat th shool-oard mting opposd QS.
In Jun 2004 th shool oard votd 3_2 against QS, nd th issu of
rationismin Rosvill smd to hav n didd in th ngtiv. Pr-
haps ausof onrplaints from Parnts tht th ord ws Wasting tim
arguirrgovr rtionism whrr ilportant issttsof funding, ovrrowding,
 9 0 A m ri ti uo luti li sm

and othr rad-and-Llttronrns wr rrot ing addrssd,rh tssuws that this is whr th ratit
not brought r.rpagin, and in th N<lvmber2-004ltion mor modrat th mo obvious prsslo
andidats prvaild, putting an nd to th urrnt Wav of antivolutionism sign and toward th lss l
in Rosvill. Som tahrs' howvr, rport that it is mrly a mattr of against volution.''
..vidnagainst voltion'' riss in
tir until th issu of rationism nd
Rosvill irgin,at last as long irs rligiotrslyonsvtivshool.oard rnm- PRED||oNs FoR
rs kp gtting eltd. In January 2005 Larry aldwll sud th distrit
and som offiials on th grounds that his onstitutionl rights wr irrfringd Prdition is unrtain,sp
during th ontrovrsyin Rosvill.Dfnding thmslvs,th distitlaird to hv said, ut som ft
on th ontrary that aldwll's proposals had rivda full hring, and, in ism,/volution ontrovrsy
fat' an undu amount of tim irnd nrg Was spnt on aldwll.s issus. distrit-ourt judg larn
A judg sidd with thrn in Nor,mr 2007, and ldwll lost his sLlit trit struk down txtbo(
(Rosnhall 2007). y th Co ounty, G<
. . T h i s t x t o o k o t i n sn
fat, rgarding th origin ol
oRPHlNG oF RA|oNlsMs
with n opn mind, studi
Kansas and Rosvill illustat tlr morphing phnomnon tht has n tak- P a r n t si n t h o m m u n i 1
ing pla for svral yars. ommunity mrnswho ar idntifid with or 2002, ut th as took a l
hold Y positions astthmslvsas ID advots.In truth, rhis is nor a th shool oard hd a rl
lngthy jotrny, aus all young-arth rationists r supp()rtrsof th Christianity) in rquiring tl
dsign viw, although not all ID supportrs ar young-arth rationists.In furthrmore, undr Gorgi
Rosvill, for xampl, oth Y and ID rsorrrswr offrd to th shool tional aus it rquidt
distrit. Th lgal advlntagof ing idntifid wit positior-r'lD, that has Th judg struk down th
not alrady n struk down y th ourts is lar. Yt ID is not without its i n g r I i g i o n .I n t h d i s i
waknsss:th phras itlligetdsig arris th ilpliation of a dsigner, sholarly rtils' as wll a
.,onol
an agnt, and it is no srt tlrirttl-risagnt is th hristian God. Prhps for fat'' langug was
this rason, th Disovry Institutativists' fom approimatly th last half m p l o y dh y a n t i v o l u r i o n
of 2001, hav n arguing not fr th inlLlsionof ID in th urriulum but gling out volution from
f o r t h i n l t l s i o n o f . . v i d n g a i n s t v o l u t i o n ' ' ( o r . . s t r n g t h s : r n dw k . would hv th fft of pr<
..ritial ..tah is rationism. Th distrit
nsss of volution''' analysis of volutiorr,'' or simpl th
ontrovrsy''). Tahing ID is mor likly to run afoul of th stalishmnt pals vaatd th ruling an<
l:1Lls than thing ..vidnag:rinstvolution,'' altl-roughth nt fft of sideration or rtrial. Th gr
nouragilrgstudntsto apt spial-rationtholog is onrmon to itlrr not onrn th judg'srulil
approah. i n g v i d n .I n d i s u s s i o
Parallling th IR's ditum that ..vidnagainst volution is vidn agrd to rtry th as.It h
..limittions plaintiffs' lgal tal riv
for ration sin.''Wiltim Drski hs writtn that on volv-
aility y ratrial rlhanisms onstitut vidn fo dsigl-t'' (Dmski prt witnsss' whih strn
2002). Talring studnts that volution is wak or unsupportd sirrthat ruld in thir favor. Th pll
annot xplin th divrsity of lif nourgsth aptanof rationisnr: tion of som of th attorn
if volution dt>snot plin living things, y dfault,rationismdos. shool ord poliy rqui
..vidnagainst nation of fators was suffi
Suh morphing of Y to ID to volution'' has ourrd
in ir numr f ratitlrrism/volution ontrrvrsisovr th pst fw yars, The sttlmnt was ver fa
inluding ontrovrsis ovr th writing of sit-tduation standads in agring nvr again to dis
Kansas, Ohio, ./stVirginia, Nw io, and Minnsota and in shool- orally, or in any othr for
distrit ontrovrsisin Dary. ontana. d Dovr. Pnnsylvania.It is lar gia stat standards, whih

t ingaddrssd,th issu was
r 2004 ition mor modrat
l r r n tw a v o f n r i v o l u t i o n i s m
t that it is mrly a mattr of
n againstvolution'' riss in
:onsrvativsho1-boardmm-
-arry aldwll sud th distrit
lstitutiOna1rights wr infringd
g thmslvs, th distrit laimd
d rivda full haring, and, in
Was spnt on aldwll's issus.
l07, and aldwll lost his suit
; phnonrnonthat has n tak-
nrswho ar idntifid with or
d v o t sI.n t u t h , t h i s i s n o t
.ationistsar suppotrs of th
r young-arthrationists. In
oursWr offrd to th shool
[d with a position, ID, that has
is lar.Yt ID is not without its
risth impliation of a dsigner,
s th hristian God. Prhaps for
from appoimatlyth lst half
rsion of ID in th urriulum ut
..strngthsand wak-
;tion'' (or
..tah th
:volution,''or simply
o run afoui of th stalishmnt
ution,'' although th nt fft of
ion thologyis ommon to ithr
n against vlution is vidn
..limitations on volv-
lrittnthat
vidnfor dsign'' (Dmski
ak or unsupportd sin that
] st h p r n o f r a t i o n i s m :
y dfault,rationism dos.
againstvolution'' has ourrd
ovrsisovr th past fW yars'
,f sinduation standards in
, and Minnsota and in shool-
nd Dovr, Pnnslvania. It is lar
Amricn Atieuolutionist 391
that this is whr th rationism/volution ontrovrsy is hadd: away from
th mor ovious xprssions of rligion or ration sin or Intllignt D-
sign and towrd th lss lgally vulnral stratgy of tahing ..vidn
against volution.''
PRDI|oNs FoR (NAR) FUURE
Prdition is unrtain, spially aout th futur, as Yogi Brra is supposd
to hav said, ut som rnt vnts may rshap th futur of th ration-
ism/volution ontrovrsy in th Unitd Stats. In January 2005 fdral
distrit-ourtjudg larn oopr tn Slm u. obb out School Dis-
trict struk down a txtook dislaimr ordrd insrtd into iology ooks
y th o onty, Gorgia, Board of dution. Th dislaimr rad:
..This ttook ontins matrial on volution. volution is a thory, not a
fat, rgarding th origin of living things. This matrial should approahd
with an opn mind, studid arfully, and ritially onsidrd.''
arntsin th ommunity had sud th distrit and th oard of duation in
2002' ut th as took long tim to om to trial. Th plaintiffs argud that
th shool oard had a rligious purpos and fft (of promoting starian
hristianity) in rquiring that th dislaimr plad in txtooks, and that
furthrmor' undr Gorgia law' th shool oard's ation was unonstitu-
tional aus it rquird th xpnditur of stat funds to promot rligion.
Th judg struk down th prati ausit indd had th fft of promot-
ing rligion. In th dision th judg took notie of law.rviw and othr
sholarly artils' as wll as tstimony that pointd out that th ..thory' not
{at''languag was ..on of th latst stratgisto dilut voltion instrtion
mploydy ntivolutionists with rligious motivations'' (Slm, 1308). Sin-
gling out volution from all sintifi thoris for spial ngativ tratmnt
would hav th fft of promoting th favord altrnativ to volution, whih
is rationism. Th distrit appald th as. Th Fifth iruit ourt of Ap-
palsvaatd th ruling and snt th s ak to th Distrit judg for ron-
sidrationor rtrial. Th grounds for rturning th ase to th lowr ourt did
not onrn th judg's ruling pr s, ut rathr a mssy ourt rord and miss-
ing vidn.In disussions among th judg and th two lgal tams, it was
agdto rtry th as. It gan to look unpromising for th dfenswhn th
plaintiffs' lgal tam rivd prmission to ropn disovry and bring in x-
prt witnsss,whih strngthndthir as for a judg who alrady had
ruld in thir favor. Th plaintiffs' lgal tam was also uttrssdy th addi-
tion of som of th attornys and witnsss who had sussfully dfatd a
shool oard poliy rquiring ID in Dovr' Pnnsylvania; prhaps th omi-
nation of fators was suffiint to nourag th distrit to sttle out of ourt.
Th sttlmntwas vry favoral to th plaintiffs, howver, with th distrit
agringnvr again to dislaim vlution, whthr with txtbook stikrs, or
orally,or in any othr form. Th distrit also was rquird to follow th Gor.
gia stat standards, whih urrntly rquir th tahing of volution, and to
92 Amri Antiuolutioism

rfrain from ising or rdating matrials On vollltion from instrtional rligious ida. Th pl
rnatrils.This lattronsrraintwas stimulatdy an arlirpratiin th dis- dfinitionalhartr
trit in whih th pages on volution in ttl.looks routinly wr t out to allss nd thus tsta
prvnt studnts from rading thm. posd that aus ID
Had th o ounty distrit lost on appal' othr ..thory'not fat'' poli- thws no pdagogi
is and dislaimrs would hav n nouagd, and I prdit w would At th nd of a long t
hav sn an aundan.Dislaimrs ar popular with shool oards aus did not mrly strik d
thy provid th apparan that th shool ord is ..doing somthing'' qustion of whthr ID
aout th tahing of volution without rquiring urriulum hange or th that th shool oard u
othr and pns of rviw and purhas of nw instrutional matrials. ligious viw' ut .|udg
Thy satisfy a onsrvativ hristian onstituny without spnding mu mnts for and agair-rst t
..in th h
tim or mony' that h did so
In Dmbr 204, a month aftr th Slm as Was trid' a lawsuit was and othr rsourswh
fild y parnts of hildrn in Dovr, Pennsylvani' protsting a poliy of its ing th pris qustion
oard of duation that would hav rquird th tahing of ID and a variant Although ID suppor
of ..vidn against volution.,' Th Dovr rsolution, whih latr was in- ritiizd te Kitzmille,
srtd into th urriulr-rm,rad: ..Studnts will mad awar of gaps/pro- rtly) dlaingID was
lms in Darwin's Thory and of othr thoris of volution inluding, ut not quird th judg to ult
limitd to ID. Th origins of Lif is not taught.'' In Novmr 2004' amid ould onstitutio
ommunity rumors of a possil lawsuit' th Dovr Board of duation is- iudg's dision on wh
sud a ..larifiation'' that statd that th ID ttook of Pds d Pople witnsss for oth sid
would mad availal to srudntsas a rfrnfor th topi of ID. on- widly usd by sintt
fusingly, th larifiation also statd that tahrs wr not to tah ration- Was that th dfnswi
ism or IntlligntDsign. ut th judg disagr
In thir omplaint plaintiffs argud that ID is a rligious onpt indistin- shool lassroom.
guishal from rationism' and that its tahing would violat th stalish. Th witnsss for th
mnt laus. Th tril took pla ovr a si-wk priod in th fall of 2005. p r s u d r h j u d g o f r
Th Disovry Institut pulily distand itslf frorn th lwsuit, stating that irduib1 omplxit1
..although of th llod lottirrg
w think disussion of intlligntdsign should not pohiitd,
w don't think intllignt dsign should rquird in puli shools'' thy wntd and mort
(..Lading Intllignt Dsign Think Tank'' 2004). Although originally a num- Shool Board disior
r of Fllows of th Disovry Institut agrd to tstify as prt witnsss sin-whih will
on half of th dfnse, all ut two withdrw from th as, ut not until it of ID.
Was too lat to add nw prt witnsss. But th two who rmaind, If th Dovr Ara S
Mihal Bh and Sott Minnih' wr th ID proponnts with th st si. th ountry would har
ntifi rdntials, and thus th most qualifid to dfnd th sintifi rdi_ gardlss of th hor
bility of ID. tahing of [D y st
Th ky lgal issu in Kitziller u. Dour Ar S'hool District 'vlas viwd as zr fornr of
whthr thr was ir valid sul rason to th ID. Th ovious rligious ouag th ringing
onnotations of lD would not hav n suffiint groDnds to strik down diffrnt nam. Howr
th poliy had thr n a valid sulr purpos and fft for tahing it. p a l d l l d t h r f o i s
Th dfns,thn, had to dmonstrat that ID was a valid sin,and that ID proponnts might
tahing it as an altrnativ to volution would provid studnts with an ity of an ID poliy' wi
llnt ritial thinking xris. vr, b:tttsof th l
Th plaintiffs had to onvin th judg that ID was ot valid sin,and matrial irr th trial r<
thrfor th only rason to tah it was unonstitlltionallv to Dromot a stantially diffrnt out

on volution from instrutional
c y an ariir prati in th dis-
books routinly wr ut out to
..thory,not fat'' poli-
al, othr
rragd, and I prdit w would
'ular with shool ords aus
..doing somthing''
ol oard is
uiring urriulum hang or th
' of nw instrutional matrials.
ituny without spnding muh
lJtiC?Sewas rrid, lawsuit ws
,lvania' protstinga poliy of its
r h t a h i n go f l D n d a v a r i a n t
. r s o l u t i o nw
' hih lt was in-
vill mad awar of gaps/prob-
s of volutioninluding, ut not
ught.''[n Novmr 2004' amid
l Dovr Board of F-duationis-
' ttook of Pds d People
frnfor th topi of ID. on.
h r sw n o t o t h r a t i o n -
D is a rligious onpt indistin.
:hingwould violt th stalish-
.wk priod in th fall of 2005.
slf from th lawsuit, stating that
dsignshould not prohiitd,
rquird in puli shools''
004). Although originally a num-
.edro tstifyas xprt witnsss
w fom th as, ut not until it
:s. But th two who rmaind,
ID proponntswith th st si-
id to dfnd th sintifi rdi-
|ouer r Shool Distrit was
l tah ID. Th ovious rligious
uffiintgrorrnds to strik down
urposand fft for tahing it.
: ID was a valid sin, and that
would provid studnts with an
,hatID was not valid sin, and
unonstitutionallyto promot a
mri Atieuolutioism 9
rligious id' Tlr plaintiffs olltndd that ID did not mt important
dfinitional hrirtristisof sin (suh as ing rstritd to natural
usSand tlrus tstal)and tlr:rtits fat lairns wr inorrt. Thy pro-
posd that as ID was not :l sin and its fat laims wr rfutd,
thr was no pdagogil rason to tah it in a sin lass.
At th nd of a long trial' th plaintiffs wr rwardd with a dision that
did not mrly strik down th Dov poliy, ut that also wighd in on th
qustion of wlrthr ID was sintifi.Thr ws plnty of vidn showing
that th shool board unonstitutionally intndd to promot a strian r-
ligious viw, ut Judg John . Jons III additionally onsidrd th argu-
mnts for and against th lim that ID was valid sin.Judg Jons wrot
..ir-r
that h did so th hop that it may prvnt th ovious wast of judiial
rrdotlrr rsourswhih would oasiond y a susqunttril involv-
ing th pris qustion whih is for us,, (Kitzmiller,75).
Although ID supportrs from th Disovry Institut (D.!7olft al. 2006)
ritiizd the Kitziller dision as unnssarily (and in thir viw, inor-
rtly)dlaring ID was not a valid sin,th lgal thoris of oth sids r-
quird th judg to ul on prisly this pint: if ID was valid sin,thr
ould b a onstitutionally admissil sular rason for tahing it. Th
judg'sdision on whthr ID was sin ws rlativly straightforward:
witnsss for oth sids tstifid that y th standard dfinition of sin as
widly usd y sintiststoday, ID did not qualify as sin.Th diffrn
was that th dfnswantd to hrngth dfinition of sinto inlud ID,
ut th jirdg disgrd thr sul.ra dision should b md in th high_
shool lassroom.
Th witnsss for th dfns did not far any ttr in thir ttmpt to
prsuadth judg of th validity of th laims of ID (suh as th supposd
irrduil omplity of th atrial flagllum and th unvolvaility
of th lood lotting mhnism). In th nd, th plaintiffs got vything
thy wantd and mor: a dision not only striking down th Dvr Ara
Shool Board dision, but a dision valuating ID as not qr-ralifyingas
sin-whih will itd in any futur attmpt to rquir th tahing
of ID.
If th Dovr Ara Shool Board lrad Won th as' shool distrits around
th ountry wold hav n rrouragdto impos similar ID poliis, re.
gdlssof th hortations of th Disovry Institut not to rquir th
tahing of ID by statut. Int[lignt Dsign is widly-and auratly-
viwd as a form of rtionism, and a lgally sussfulpoliy would n-
ourag th ringing of rationism into th lassroom, alit undr a
diffrnt nam. owvr strong a dision, te Kitzliller ruling Was not ap-
paldand tlrrfois prdntonly in th iddl Distrit of Pnnsylvania.
ID proponnts might attmpt to find anothr vnu in whih to tst th lgal-
ity of an ID poliy, with a diffrnt st of fats and diffrnt judg. How-
vr, aus of th rdth of th Kitzmillr dision nd th walth of
matrialin th tril rord' it is doutful that anoth trial would hav a su-
stantialiydiffrnt outo1.
94 mricntiuolutilism

Th most popular antivolutionist stratgy in th futur, though' will

dirtivs from shool oards or stat oards of duation for tahrs to
..balan'' AKNoWLDG
volution with th tahing of ..vidn against volution.''
It is not lar whthr judgs will psuadd,as Was Judg oopr in th I t h n k G l n n B n ha n
Slm as, that ..vidnagainst volution'' is a od phras for rationism; o h m n u s r i p .w h i h
this rquirs an appriation of th history of th rationism/volution
ontrovrsy. Baus rationists liv and loudly prolaim that thr ar
Nos
only two altrnativs, rationism and volution, thy nourag tahrs to
tah that volution annot xplain th divrsity of living things. \/ithvolu- 1. ration sinhad
tion out of th pitur, thy liv that studnts will onlud that living of Judg !illiam Ovrto
things wr spially ratd. During 2008 thr was a flurry of ..Aadmi only rr,voaltrnativs:vo
ration sin advoat
Frdom Ats'' introdud into svral stat lgislaturs(Florida, issouri,
vidn for rationism;
Alaama' ihigan' and South arolina) that would prmit and nourag altrnativ of ration y
tahrs to tah that volution is an rronous ida. Boilrplat for suh 2. In kping with th
..Aadmi
Frdom Ats'' is providd y th Disovry Institut on a w- divirr tion in th rlr
sit (Aadmi Frdom Ptition 2008). Rathr t'an diretizg tahrs to fi n-tuning argumntsa(
s t l p o f t h i s d i s u s s i o nt
tah th ttook vrsion of volution and thn ritiiz this viw, suh ills
a n d o n t l n . JS t n g r2 0 0
prosrib ations y shool distrits that might pnaliz th tahr for do-
3 . m p l si n | u d D
ing so. Rathr than ovrtly proposing that rationism b taught, th fous- Hunrr200l; andWik
or misdirtion-is on th aadmi frdom of tahrs to tah ..strngths 4. [n truth, origin si
and waknsss'' of volution (as in th South arolina ill, sB 1386 a s t r o n o m y ' g o l o g y ' i o I
..full h . l o g yt:h l l i g h a n g . t
[..Antivolution Lgislation in South arolina'' 2008]) or th rang of
th origin of humns.
sintifi viws'' (as in Florida's SB 2692 f.,ntivolution Bills Dad in
Florida'' 2008]). But what suh ills propos is to prott tahrsfrom on-
squns of violating th First Amndmnt. l/hthr suh ..gt ut of jail |BLIoGRAPHY
fr'' ards would onstitutional rmains to tstd, ut at th tim o{
this writing (July 2008) non of ths ills hav pssd. A a d m i F r d o mP t i t
.!ill .admifdm
judgs always strike down suh poliis? Lgal xprts notd that y
Altrs, B. J., arrd . . Nt
th end of th first trm of Prsidnt Gorg \W.Bush' ..all of th 13 iruits duation. uolutior
will hav a majority of Rpulian judgs'' (arp t aL.204,25). As Bush's , \ n t i v o l u t i o nb i l l s d . l di
sond trm oms to an nd, ..PrsidntBush has namd 294 judgs to th ay 3. http://www.
fdral ourts' giving Rpulian appointsa solid majority of th sats,in- _i l ls_dd_in_fl_s
Antivoiution lgislation
luding a 6".to-40o edge ovr Dmorats on th influntial U.S. appals
duation, ay 15.
ourts'' (Savag2008' A11). An analysis of disions of judgs appointd y 5 3 5_ntivolution_l
th last ight prsidntsshows that Prsidnt Bush's first-trmappointsar ntivolution standards:
partiularly onsrvativ in ass involving ivil lirtis' among whih is- d u a t i o n ,N o v m
sus of sparation of hurh and stat ar found. ..Only 28 prnt of th _ntivolution_stnd
Bush ohort votd on th liral sid of issus prtaining to Bill of Rights Bh,. |996, po'*|n.'
Nw York: Fr Prs
and ivil rights mattrs, thus giving th prsidnt th lowst sor of any
2000. [untitlda
modrn hif utiv,' (arp et a|,204,26).It is not unfair to onlud .org/gi/lttrsl 288l'
that th lgal limat favoring th stlishmnt laus ovr th fr xris 27. Th F.dg,
laus in dutional affairs has waknd. It may that th lgal saf- Nw Yok: l-rPrs
guards that hav until now disouragd th inlusion of various forms of Blkston, N. W. 1997.
Bo: h Biohmi,,
rationism in th puli shool sin lassroom will b mor diffiult to
no.4:445-447.
mustr by th sintists and tahrs and parnts who support volution
airns-Smith, A. G. 1985
duation. Univrsity Prss.
 meri ntieuolutionism 395

in th futllr, though, will

i of duation for tahrs to AKNoWLDGMNs
.vidnagainst volution.''
.!sly
d, as was Judg Coopr in th I thnk Glnn Branh and lsrry for hlpful ommnts on and orrtions
to th manusript,whih sustantiallyimproved it.
s a odphrsfor rationism;
of th rationism/volution
loudly prolaim that thr ar Nos
n, thy nourag tahrs to
.With
ty of living things. volu- l. rtion sin had arlir n typifid y a ..ntrivd dualism,'' in the words
lents will onlud that living of Judg !illiam Ovrton in L u, Arkss, whih assumd that thr wr
..Aadmi only two altrntivs:volution or spial ration. With only tw altrnativs,
r Was a flurry of
ration sin dvoats argud that vidn against volutiot-twas ipso fato
g i s l t u r (sF l o r i d ' M i s s o u i ' vidnfor rtionism; ID likwis rlis on disproving volution to Support th
t would prmit and nourag altrntivof ration y an intllignt agnt.
ous ida. Boilrplat for suh 2. In kping with th ig-tnt stratgy' ID also wloms thos who s rom fr
Disovry Institut on a w- divin ation in th ralm of osmology; disussionsof th anthropi prinipl and
fin-tuningrgumntsr not unommon in thir puliations. It is yond th
hr than directig tahrs to
sopo{ this disussionto orrsidrths;s anson 2003 flr rprsntationpro
n ritiiz this viw, suh ills nd on and Stngr 204 for an tndd argumnt gir-rst.
ht pnalizth tahr for do- 3. ampls inlud Dmski 1998, 1999; Johnson 2002; orland 1'994;
ltionism tught, th fous- Huntr 2001; and.Wikr 2002.
..strngths 4. In truth, origin sinrdusto thos issusin th historial sinsof
of tahrsto tah
iouth arolina ill, SB 1386 astronomy' gology, iology, and anthropology that hav impliations for hristian
..full rang of thology:th ig ng, th origin f lif, th rigin lf th ..kinds'' of aimals, and
l'' 2008]) or th
th origin of humans.
[..AntivlutionBills Dad in
is to prott tahrs from on-
.Dhthr ..gt out of jail
suh |LIoGRAPHY
o tstd, ut at th tim of
Aadmi }.rdomPtition. 2008. Disovry Institut.http://www
r passd.
.aadmifrdomptition.om/frdom.php (ssda 22, 2008|,
s?Lgal prts notd that y Altrs,B. J., and . . Nlson. 2002. Prsptiv:Tahing volution in highr
. . a l lo f t h l i r u i t s
W. Bush, d u a t i n . u o l u t i . l 5
6 , n o . ] 0 : l 8 9 1 - 1 9 0 l .
) a r p t a | . 2 0 0 4 , 2 5 ) . A s B u s h ' s ntivolutionills dd in Florida. 2008. Ntional ntr flr Sinduation,
h has namd 294 judges to th May . http://www.nsw.org/rsours/nws/2008lFLl739'antivolutiln
_ills_dad_in_fl_5_3_2008.sp (assday 23, 2008).
l s o l i d m a j o r i t yo f t h s a t s ,i n -
Antivolutionlgislationin South arolina. 2008. National ntr for Sin
n th influntial U.S. appals duation, ay 1 5. http://www.nsw'org/rsours/nws/200 8/S/
isionsof iudgs appointd y 53'5_ntievolution_lgislation_in_s_5_15_2008.sp (assdMay 22, 2008).
Bush's first-trm appointS ar Antivolution standards adptd in Kansas. 2005. Nationl ntr for Sin
ivii lirtis' among whi is. Ju tin. N<mr 17. http://...n;srr.g/su;s/nrr/]00 5 /KS/3 ]6
.ound...only 28 prnt of th _ntivolution-standards_adopt_1 1_10_2005.asp (assday 15' 2008).
Bh,. 1996. Dri's Blk Bo: Th Biochmil hllge to uolt.
us prtaining to Bill of Rights Nw York: Fr Prss.
: s i d n tt h l o w s s o r o f n y 2000. [untitld artil]. lie Si gzin, http://www.sinmag
26). It is not unfair to onlud .org/gi/lttrsl288l 5467 1813#165(assdJuly 7, 2000).
L n t l a u s o v r t h f r r i s 2007. Th dg of uolutkl: Th Srh for t Limits of Driism.
Nw York: Fr Prss.
. It may tht th lgal saf-
Blakston,N. !. 1997. Argumntum ad ignorantiam: A rviw of Drlui's Blk
inlusion of various forms of
Bo: Th BiohmiI hIlg tl uolutkl. Qurtrl Ruit,tlof Biolog 72,
'sroom will mor diffiult to no. 4: 445-447.
parnts who support volution airns-Smith, A. G. 1 985 . Su ltts to th rigi of Lif. amridg: ambridg
Univrsity Prss.
96 mer i tit, o lt i.ltsr

rrrp, R. A., K. L. N1anning,and R. Stidhar.2004. h dision-mkinghavior H t t n t r , . G . 2 0 0 1 . D . l

of G-lgW. BLrsh.sjudiial iIrtn.rnts, 88, nr. 1: 20-28.
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valir-Smit,. |997. Th lirrr.] iohmist.rltr]sin ollgd uoltio 1:r>s lf uolutitl. 200
12: 162_|6. Produtions.
oyn, .|.^. 1996. God in th dtaiIs.Ntur 383:227-228. Institut for ration Rr
ration musum unptd sllss. 208. Florid tsptist Witss, Aril 17. mning. lR llp
http://www.floridaaptistwitlrss.om/8688.artil (:rssd y 15, 2008). .htm.
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1. Th dision.makinghavior
d i t u r 8 8 , n o . 1 : 2 0 _ 2 8 .
.ds itl F'olog d uolutil
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8.
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i,sTrnstils3: 1_8.
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in th Unitd States.nul

..thistl
sholrshi,''irnd
N,ltkI ntr for Si
w.org/rsours/artils/6 149

Rports of th Ntill ntr for

dutio..!7stprt,T: Grnwod

i s m : h n g s n d t ' n t i n u i l i s .

lsign in sinlassrooms.
e s1 4 ( S u p p l .\ ) : 8 6 6 9 - 8 6 7 6 .
:al nalysisof IntlligntDesign in
-2.98.
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y rirsning.lntrtkl J ou'rnl

s? In . Young nd T. dis' ds.,

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| 'B i l l l g20 3 : 1 1 1 _ 1 1 6 .
irtor.DVD. 67 minuts. Lir

l Did lt Gt Hr? 8y
7athtowrBibl and Trat Soity

A ritiqu of Willirrn-r
lolats:
)rigisArhiv (updatd Aril 23,
qs/nfl/(assdJanuary 24, 200 5I.
nsisFlood: T.he Biblil Rord
NJ: Prsytrinnd Rformd.
m Hdoisfs.Downrs Grov, IL:

r'antgsof thft vr toil: T dsign

g d P h i l r l s o p h l | 6 : 7 7 1 - 7 2 4 .
,tDsig Fils: Sitifi
nsrvikN . J : R u t g r sU n i v r s i t y
ALPHABlAL GUlDt
Adapttio|| Cl,||

This ook, pulis

.!7illiams,
is on o]
thugh no-Darwi
lishd paradigm
ommittd to th t
ious imprssions.
th ologist Vro
Soil Behuilur
grou ovr th in
possiility of grou
ought to alwys pl
vidual fftsand l
In making his u
groundraking p
(1964a,1964) ,
(ants, s, and w
known as kirr s
hans by raising
prodution.
Anothr topi a
progrss' thn mu
.!7il
Rus 1996).
progrss' pointing
ity) ar oftn vr
tks dvlopmn
rdily ompl
pl is ttr.A jt
propllr ngin,
lattr.
Th main thrus
and genrally suff
sorn Was rsrv
ltion. . H. Wa
similation, wh
.\/illiams
flis. h
dpttion nd Nturl Selection (Gorg C. Williams)
This ook, pulishd i 1966 y th Amrian ihtlryologist Gog '
Williams, is on of th grat dunking works irr th history of sin.Al-
though no.Darwinism, or th syrrthtithory of volution, was th sta-
lishd paradigm in volutionary studis, Williams (who was dply
ommittd to th tlrory itslf) flt that it was inftd y a numr of falla-
ior.rsimprssilns.Ivlost signifiant of ths Was the lif, promulgatd y
.Wynn-drvards
th ologisr Vro . in iml Disprsiotl in Rltiotz to
Soil Bhuior (|962\, hat stlrntimsnaturl sltion an favor th
group ovr th irrdrvidual.Although h did not want to dny th outright
possiilityof group sltion,Y/illiams flt that on gounds of simpliity on
ought to always prfr th hypothsis that group fftsr ausd y indi-
vidual fftsnd tlrirtsltionthforrnust nefitfirst th individual'
In making his as, Wiiliarns was on of th first tO mak rfrnto th
groundraking paprs of th tlrn-graduat studnt \/illiam amilton
(I964a, 1964), r,vhohad shown how th strility of hymnoptran rvorkrs
(ants,s, and w.asps)ould plaind as a rsult of what has om
known as kin sltion. Th workrs re improvillg thi rprodutiv
hansy raising frtil rlativsinstad of putting fforts into thir owt] r-
prodution.
Anothr topi attakd with rlish y \/illiamsws that of voltionary
progrss'thn mrrh favord y popl lik Julian Huxly (7942; s lso
Rus 1996). \Williams was withring in his ritiqu of various notions of
progrss,pointing out that popula ritria of improvmnt (lik ompl.
itv) a oftn vr1' rrrislading.p:rt from th fat tlrat gnrally lro on
taksdvlopmntinto aount (th livr fluks rht afft shp hv an in-
rdilyompl lif history), it is y no mans ovi<lusthat th mor om-
plxis ttr.A jt nginis larly suprilrto a gasolin-fuld,motor.drivn
propllr ngin, d yt in prinipl th formr is lss omplx than th
lattr.
Th mairr thrust of th ook, holvv' is tlrrtni1turalsition is th main
and gnrallysr-rffiintpiarrationof volutionary hng.n rtiular
sornWas rsrvdfor thos who would rpla or othrwis supplmnt s-
ltion.. . Waddington (1957\ had intrldud th notion of gnti as-
similation,whr lr was al to sit-nulatLararkian-typ ffts irr fruit
flis.Williams hal no rjtirlnto th rnpirilfindings, ut thought tlrt in-

40
gssiz

frns out th irradquay of sltion as suh ar simpl not wll tkn.

For him, and for his rany nthtlsiasti rzrdrs' a strippd_down' pur Dar-
winism is not only asthtially mor attrativ' it is sintiIially suprior-
a point lost on th philosophy of iology ommunity, whih has spnt a
h,ppy nar-half ntury wriring paprs and ooks rfuting ky linrs f
\filliars's work.

BlBLIoGRAPY

amilton, W. D. 1964zr.Th gntialvluti<lt-t f s<lirl

hvitlurI. Jorl oi
Tboretil Billog 7 : 1-| 6' \o{
|964. Th gntial volution of soial haviour II. Jourl lf Thortil
Biolog 7: 17_52,
Huxly, J. S. |942. t,'olutil:h odr S'thsis.Llndn: Allrr rrd Unrvin.
Rus, . 1996. old to : Th opt of Progrss i ullutir Birl|lg,
mridg, A: Harvard Univrsity Prss.
Sor, ., d. [1984l |993' ptul lssues i uolutitlr Bklklg.2nd d.
mbridg, A: IT Prss.
.!?addington,
Louis Agasstzws ms
. H. 1957. Th Strteg lf th Gns. Lndon: Alln and Un,tlr. spimn,lopom s1
Y/illiams, G. ' 1966. Adpttion d Nturl Sltio. Printon, NJ: Pinton d r a n ri n w h i h h s w t
Univrsity Prss. tht h had md from r
Wynn-drvrds, V. . 1962.'Aitttl Disprsi<lttin Rlti<lttto SoiI Bht''itlur. o r r s p o n d d a l l.t
d i n u r g h :O I i r r a n d B o d _.R.
20-21.

Agassiz,JanLuis Rdlphe$8a7_t873\
Born in Switzrland in 1807, l,ouis Agassiz as a oy Was a kn naturalist.H o lssifitio (|8,
studid iology at svralGrman univrsitis,arning an advand dgrin aross mryoni for
..On !
narural history \n 1829. H soon arnd a rputatioll as a brilliant rsarhr' th istn of
thanks to ooks dsriing spis of fislr' living and fossil' and to his arful blans for h a
.silliam
studisof Alpin glairs(Agassiz 1833_1843' 1840) (sfigur).Bginningin Paly, whos
1837 h ssmldvidn for surprisirrgnw thory aout an I Ag, makr or nginr Wh
whn a thik lyr of i had rvrdnorthrn urop. Aftr moving to th pupos. Agassiz.sGod
Unitd Statesin 1846,e amon of Amria's lading sintists,founding ral variations on a fv
th usurn of omparativ Zoology at Harvard in 1859. Agassiz rgudthat
A harismati lturr' Agassiz hamd th puli as wll as his studnts ation and nnot har
with dsriptions of rnt disovris in mryllogy' palontology, and jtd th iblial sto
anatomy. Lik Rihrd Own in Lndn, Agassiz mphasizdth pattrn of hallngd whn h n
rsmlansalld hornolgv. Althouglr it had long n known that tlr dos him no rdit'
strutur of an animal's organs is latdto thir funtion, not until th nin- an opinion wlomd
tnth ntury did hiologists onfront th fat that divrs raturssm to Darwin's o th '
b modifitions of ommon luprint or typ. Vrtratsar on sllh prtation of narr.rr. Pt
typ' austh sam layout of ons and oth organs an trad in a win argud that hom
fish, a frog, a at, and a humn. Agassiz imprssdhis audinswith unfa- ommon ansto,wh
miliar ampls of tlr sam prinipl:tht s,spidrs,and lostrsar 2rlso n t r a l s l t i o n .B f
honrologous to otl anothr' s ar otopuss,snails, and lams, or stirrfish, a P r d v o l u r i o n ,i f
jllyfish, sa urhins, and orl polyps. F.ollwing Gorgs uvir, h alld t o B r a z i l i n 1 8 6 5a n d r
thsfour goups vrtrats,ilrtiLllats,mollusks, and radits.In his ssay his mind. H did in 1

; sulrr simply not wll takn.
lrs,a strippd-down.pu Dr-
i v ,i t i s s i n t i f i a I lsyu
ommunitv, whih has spnt 1
d ooks rfuting ky laims of
of soiirl hviur |. Journl lf
haviour7I.lourl l| Thortil
/:esfu.Londorr: Alln and LIn,irl.
Prgrssi utlltitlr Biolog.
u olutionr tsio llg. 2nd d.
:res. Lndon: Alln nd Unwin.
iIti,Printon' NJ: PriI-rton
l i Rltion tl Soil Bbuiour.
t7-1873)
as a oy Was a kn naturalist.
is'arningan advand dgr in
putationas a rilliant rsarhe'
iving and fssil, and to his arful
l 3 , l 8 4 0 ) ( s f i g u ) .B g i n n i r l gi n
n8 nW thory out n I Ag'
hrn urop. Aft moving to th
rria'sladingsintists,founding
a r v a r di n 1 8 5 9 .
th puli as wll as his studrrts
mbryllogy'palorrtologyJ nd
\gassizmphasizedtlr pattrn of
it had lorrg n known that th
thi ftrntion'not Llntil th nin-
.atthat divrsraturs sm
) typ. Vtrtsr on suh
d othr organs an b trad in a
prssdhis audins with unfa.
s,spidrs,and lostrs ar also
ss' snails' and lirms, or starfish,
lowing Gorgs uvi, h alld
rllusks,nd rdiats.In his ssa1'
Agssiz 40''
r i o r -
,t$5st
.$r'
f:..
*il.r.
f.:r
Louis Agassiz was a mastr at ronstrutingfossil fish. lris Dartilrlar
s.pimn'lopom spin(,sum,ws giving him diffiultyand th.n h hd
dramin whih h sw th full form. .Whnh vntullyompard th skth
that h had mad from mmory with th atul mdddspeimn.th two
orrspondd atly.It is dsridin Agassiz 183_184J' vo|.4. tnijides.
-.R.
20-2r.
o lssifction (1859) Agassiz argud that homologis, whih tnd
aross mryoni forms and ak thrugh th fossil rord, ar vidne of
th istn of ..one Suprm Itllign'' who hd onivd ths rsm_
blnsfor h ratd thm. This agumnt is quit distint from tht of
William Paly, whos Ntrl Tholog (1s02) omard God to Wth-
makr or nginr who uilds mahins that ar lvrly adaptd to thi
purpos.Agassiz's God is lik musiin or arhittwho njoys playing
sv_
ral variations on a fw asi thms.
Agassiz argud that ah spis was th produt of a listint at of
r-
ation and annot hang, ut h ndrstood gologial tim and firmly r-
jtdth ilial story of ration. His onpt
of spis was svrly
hallngdwhn h nountrd popl of Afrin ds.,'t, and his rspons
doshim no rdit. insistd that human ras ar atually distint spis,
an opinion wlomd y slav ownrs'
Darwin's on tbe rigin of Spcies was a srious thrat to Agassiz's intr-
to
prtationof natur. Pointing to th sam phnomna itd y Agassiz,
Dar-
win argud that homology and iogography are vidn of dsnt from
a
ommon anstor, whil th adaptdnss of form to funtion is viden for
naturai sltion. Bfr long Agassiz's students and rost of his ollagus
aptdvolution, if not natural sition, ut Agassiz nvr did. Travling
to Brazil in 1855 and to th Galjpagos Islands in 1872 did nothing to hang
hls mind. H did in 787.
106 lndr

|BLloGRAPY dr hypothsizd tha

Agassiz,L. 1833-1843. Reherhs sur les PoissosFossils'Nuhtl: Imprimri vnt this sort of or,
d Ptitpirrr. groups. In onnti
184.tudssur ls g!cirs.Nuhatl:Jnt t Gssmann. (1983) ida that hur
1859.ssayo lssctio'London:Frthought Pulishing onfrs in ompl
ompany/Longman,Brown' Grn,Longmans,& Rorts,and Trunr& thought-out thoris
o. Rprint(. Luri' d.),amridg,MA: arvardUnivrsityPress,1962.
(Origirrlly hairlssnss in adults
pulishdin 1857 as Part I of L. Agssiz,otributils to the
Nturl Histor of th UitdSr/s.Bosto:Littl, Brwn.) nan and its impli
L u r i , . | 9 6 0 . L o u i s g s s i z : A L i f i n S i e . h i g : U n i v r s it yhoi f g o omination of higl
Prss. group-grouP ompt
.!.
Paly, [1802]1819.Nturl Tholog(olletdWorks,vol. 4). London: onfidn and an rr
Rivingt;n.
mntion a fw. Ths
Winsor,. P. 1991.RdigtheShpof Ntur:otprtiu Zooklg t th
Agssiz useum. l-riago: Univrsityof hiago Prss. -lvl.P.W. sarh by human }
Rihard Alandt
Amrian Assoiati
Danil Giraud lliot
Alxandr, Rihard D. (b. |929\
th Darwin Awrd iI
Rihard D. Alandr was traind as an ntomologist at ohio Stat Univr- ial Soity (|986l,,
sity. Soon aftr riving his PhD '1954, h r<loka position at th Univr- Bhavior Soity (2(
sity of N4ilrigan usltm of Zoo|<lg,whr h rmaind flr th rst of his o f S i n st n | 9 7 4 ,
arr (h am Profssor mritus in 2001). arly in his ar h on- ation for th Advanr
ntratd his rsarh fforts on Ommuniation in th singing insts-th th Animal Bhavio
rikts, katydids, and iadas. was th first iologist to us inst songs havior and volutior
to rogniz spis' and h mad numr of mthodologial innovations tifi Award io 27
fo th study of inst songs. H dsrid426 new spisand gnraof in- artils' and has tra
Sts nd h dsrid th longst and most ompliatd hyrid zon vr portant ontriutior
known in th tru katydid, Ptropbyll mllifoli' H also was a pionr in
th study of havior and mad important ontriutions to th undrstand- ILIoGRAPH
ing of how havior volvs. H forrsdon inst songs' maring havior, Alxndr,R. D. 1979
.Washington
aggrssion,and trritoriality in his studisof inst havior.Vith a numr Prss
of ollagush dvlopdimportant thortialidas aout th volution of 1.987.Th Bio
1 9 9 'H o D i
usoiality in mammals, idas that wr onfirmd with th disovry of u-
Spcis.Ann Arbo
soiality irr nakd mol rats. Pulition1.
In th arly 1970s, Alexandr turnd his attntion to th volution of hu- H u m p h r y ,N . 1 9 8 3 .
man havior. In his ook Dru-,iism zdHm Affirs (1979) h showd id. oiord: O
how ultur an sn as th produt of soial intrationamong individual ott, D., and R. D. Al
human ings who hav volvd to mximiz thir inlusiv fitnssin ns- Grllid).Philad
tral nvionmnts.In this ook h rviwd larg amount of anthropologi-
al data to larify and support his thory. |n The Biolog of orl Sstms
(1987) h prsntdth thory that th human snsof morality is a prodrrt
Altuism
of a history of intrgor.rpomptition in human groups' whih favrs any Th istn of altr
trait that hlps humn ings to form latgr and ttr-unitdsoial gups. profound haling
suggstdthat a basi fat of human xistn,as with ll sexually rpro- tions for altruism h:
duing organisms' is onflit of rprodutiv intrst among individuals. As and vn onomists
groups om largr rrd onsist mostly of gntiallyunrlardpopl, pr- illuminatd th und
vnting onflits from r:rkingu th group orrrsmor diffiult. Alxn- modrn volutionf

s F ossils. Nuhtl: Imprimri
Jnt t Gassmann.
rrthoughtPublishig
rns,6{ Rorts, and runr &
: Harvard Univrsity Prss, 1962.
Agassiz, otri|utios to th
r: Littl, Brown.)
hiago: Univrsity of Chiago
td lorhs,vol. 4). London:
re: omprtit' Zoolog t th
_'P.w.
LigoPrss.
tomologistat ohio Stat Univr-
h took a position at th Univr-
:r h rmaind for th rst of his
2001).arly in his arr h on-
ation in th singing insts-th
first biologist to us inst songs
r o f m t h d o l o g i a li n n o v a t i o n s
426 new spis and gnra of in_
)st ompliatd hybrid zon vr
rcllifoli'H also was a pion in
: ontriutionsto th undrstand-
cn inst songs' l-tlatinghavior,
of inst hvio. .With a nmbr
.tialidas about th volution of
lnfirmd with th disovry of u-
s attntion to th volution of hu.
Humn ffirs (1979) h showd
oial intrationamong individal
rizthir inlusiv fitnss in ans-
d a larg amount of anthropologi-
Io Th Biollg of orI Sstems
man snsof morality is a produt
humn groups, whih favors any
r and ttr-unitdsoial groups.
istn,as with all sually rpro-
:ivintrst among individuals. As
'f gntially urrrlatd popl, pr-
p omsmo diffiult. Alan-
ltruism 407
dr hypothsizd that morality as a form of indirt riproity hlps to pr-
vnt this sort of nflit and allows th formation of largr, ttr.unitd
groups. In onntion with this ida, h pandd Niholas Humphry's
(1983)ida that human intllignhas volvd ausof th advantagsit
onfrs in omplx soial intrations. Alandr has dvlopd wll-
thought-out thoris aout a nunrbr of human traits. hs inlud rlativ
hairlssnssin adults, fmal orgasm' sxual dimorphism, ologial domi-
nan and its impliations for soiality, soially imposd monogamy, th
omination of high patrnity onfidn and multi.mal soial groups'
group.group omptition as a form of pla, th assoiation of low patrnity
onfidnand an mphasison utrin kinslrip, and ross-ousitlmrriag, to
mntion a fw. Ths thortial idas ar th foundation of mrrh urrnt r-
sarhby human havioral ologists.
Rihard Alandr has rivd a numr of important honors: th
Amrian Assoiation for th Advanmnt of Sin Award (1961), te
Danil Giraud lliott dal y th National Aadmy of Sins (197I),
th Dwin Award in Inst Bhavioral ology y th Florida ntomolog-
il Soity (\986), and Distinguishd Animal Bhaviorist y th Animal
Bhavior Soity (z00z). H am a mmbr of th National Aadmy
of Sinsin 1974, and has n ltd a Fllow of th Amrian Assoi-
ation for th Advanmnt of Sin, th ohio Aadmy of Sins, and
th Animai Bhavior Soity. H was ltd prsidnt of th Human B-
havior and volution Soity in 1995 and rivd its Distinguishd Sin-
tifi Award in207. H has pulishd four ooks and ovr 130 sintifi
artils,and has traind 32 studnts, most of whom now ar making im-
portant ontributins to th volutionary study of havior.
|LIoGRAPHY
Alxandr,R.D. 1979' Dtliism d Huntn ffirs. Sttl:Univrsitof
WashingtonPrss.
1987' The Biolog of orl Sstems.Hawthorn,NY: Aldin d Gruytr.
1990.Ilot,tlDid ums uolu?Refltionso th Uiqul Uiqu
Spis.Ann Aror: Univrsityof ihigan N4usumo|,Zoo|og,Spil
Puliation1.
Humphry,N. 1983. osiot.tsss Regind:hptersin th Dulopmtof
id. xford: Oford UnivrsitPrss'
-be
ott,D., and R. D. Alandr.1'983. Austrli rihts(orthoptr:
Grllide). Phildlphia:Aadmyof Ntural Sinsof Philadlphia. -w. t.
Altruism
Th istnof altruism in animls and humans has posd on of th most
profundhallngsin volutionary iology, and th volutionary xplan-
tions for altruism hav inflund psyhologists, soiologists, philosophrs,
.!hil
nd vn onomists. volutionary thry and empirial studis hav
illuminatdth undrstanding of altruism, it rmains an ativ sujt in
mdrnvlutionary iology and psyhology.
408 ltruism

Th prolm of ltruism is dividd into two sparatut rltd issus:i. (1998) argu that' und(
ologial altruism and psyhologil altruism. In iologial altruism, an or- ltion. If th avrag1
ganism prforms n altruisti at whn th at nfits anothr organism at a thn th avrag fitnss
ost to itslf. Bnfits and osts ar masurd in trms of rprodutiv fitnss. fitnss onsquns of
Suh altruism is wll doumntd in many spis. Vrvt monkys nd t a i n p d o m i n a n l '
prairi dogs hav sntris at th outskirts of olonis that alrt mmrs of suvrsionfrom wirhi
th olony of approahing pdators,oftn at th ost of oming th tar- twn groups' Althou1
gts of thos prdators. Psyhologial altruism, whih is th olloquial sns workd out and th pll
of th trm, is intntionally prforming an ation at on's own ost in ordr studis of flour tls
to nfit anothr. Th nfits and osts for psyhologially altruisti ats, sltion modl ar m
howvr, nd not hav anything to do with rprodutiv fitnss. An xampl ratly xplain naturl p
of a psyhologilly altruisti at wold to plant a slow-growing tr, not xplin how altru
knowing full wll that you will not nfitfrom its shad or fruit' mutation in bhaviora
Although iologial and psyhologial altruism ar sparatprolms, th- for thy om nur
orists hav offrd iologial undrpinnings for psyhologial altrism. Th A sond rson fo
prolm of altruism, oth iologial and psyhologial, is at th ntr of pat of lWilliam Hamilt
grounding a thory of morality within iology. I will disussth two kinds of has om to known.
altruism in turn. havior is gntlallya
Altuism sms to a prlm for volutionary thory whn looking at s- tivs, with whor thy
ltion from an individual prsptiv (i..,sltion that ous on th lvl of altruisti gns througl
organisms). If altruisti organisms liv among slfish organisms' th slfish r- \/ith this xplanation,
ganisms will always mor fit than thir altruisti ountrparts. ovr tim, Frm gn's-yviw
altruisti individuals will sltd out in favor of slfish ons. For ampl, fi hrslf for th livs
sntry monkys lowr thir fitnss y aling othrs to th prsn of prd- of hr gns.But th
ator. ovr tim, slfish monkys, ons that do not tak thir turn on wath' fo th ost of slf-sar
would produ mor offspring and vntually liminat any altruisti gns. Kin sltion appar
harls Darwin' tnTh Descet of n (7871)' notid this prolm and m o t h r s d f n d i n g h i
..although high
hintd at a solution: a standard of moality givs ut a slight planation is that it r
or no advantag to h individual man . . . an inras in th numr of ut largr familial rla
wll_ndowd mn and advanmnt in th standard of morality will r- than distant ons' Whi]
tainly giv an immns advantag to th group,, (p. 1'66).Darwilr ontinus sual or olfatory us' tJ
by suggsting thet groups that ontain mor altruisti individuals will out. fftiv prois for rl
ompt groups omprisd mainly of slfishindividLrals.From this pssag,it that r los y'' will
appars that Drwin Was suggstingthat sltion an our at th lvl of Suh aggrgationn o(
groups' a radial dpartur from his or afgumnt aout th importn of if disprsal from th ho
sltionamong individuals. But how an w pli
volutionary iologists hav vaillatd in thir aptan of this so-alld rlatd if group sltio
group sltion argumnt. Darwin's influn gav it grat rdn for d- tion was artiulatd by
ads. It fll from favor in th 1960s undr th influn of Gorg C. iprol altruism. A<
IWilliams's writings (e.g., 1966| aout sltion and adaptation aus
toward othr organism
group sltion for any attribut am to onsiderd a iologial rarity. This planation dos
on of th main prolms with the group sltion argumnt is that omp- stats in animals. A sta
tition within a group will always favor th slfish individuals, so that vn if havior of vampir a
groups of altruists ar mor fit as a group than groups of slfish individuals, out fding; howvr, v
th groups of altruists will suff suvrsion from within. Rntly, lrowvr, individual nds. Vam
.Wilson
group sltion nrodls hav rivd a spiritd dfns.So arrd bats who hav rntly

o sparatut rlatd issus:i-
r. In iologial altruisnr, an or-
:t nfitsnothr oganisrn at a
in trms of rprodutiv fitnss.
y spis.Vrvt monkys and
olonisthat alrt mmrs of
at th ost of oming th tar-
lm' whih is th olloqtrial sns
tion at on'S own ost in ordr
lr psyhologiallyaltruisti ats,
produtivfitnss. An ampl
to plant slow-growing tr'
'om its shad or fruit.
'uism sparatprolms, th-
for psyhologialaltruism' Th
'syhologial,is at th ntr of
4y.l will disussth two kinds of
'onary thory whn looking at s.
:ltionthat ours on th lvl of
g slfishorganisms, th selfish or-
ltruistiountrparts.ov tim,
avor of slfish ons. For ampl,
] othrs to th prsn of a prd-
do not tak thir turn on wath'
l y l i m i n a t a n y a l t r u i s t i g n S .
( 1 8 7 1 ) ,n o t i dt h i s p r o l m a n d
lard of morality givs ut a slight
. . an inras in th numr of
r standard of morlity will r-
oup'' (p. l66). Drwin lntinus
'r altruisti individuals will out-
i n d i v i d u a l s F. r o m t h i s p s s a g ,i t
ltion an our at th lvl of
rgumntaout th impotan of
t thir aptanof this so-alld
1 gav it grat rdn for d-
:rdr th influn of Gorg .
ltion and adaptation aus
onsidrd a iologial rarity.
sltion argumnt is that omp-
slfishindividuals, so that vn if
than goups of slfish individuls,
n from within. Rntly, howvr,
.Wilson
riritd dfns. Sor and
Altruism 409
(|998) argu that' und rtain onditions, altuism an promotd y s-
ltion. If th avrg fitnss of groups prdominatd y altruists is gratr
than th avrag fitnss of groups prdominatd y slfish individuals, th nt
fitnssonsquns of ing an altruist may positiv. Groups an main-
tain a prdominantly altruisti population and dfnd thmselvs ginst
subvrsion from within if thr ar priodi migrations of individuals -
twn groups. Althorrgh th mathmatis for group sltion hav n
workd out and th plausiility of th thory onfirmd with primntal
studis of florrr tls, it is lss ovious that th rquirmnts of th group
sltion modl ar mt oftn in natur and thus that th modl will au-
ratly xplain natural phnomna. Furthrmor' grouP sltion modls an-
not plain how altruists, whn initially rar (as would happn through
mutation in havioral traits), avoid ing liminatd by slfish individuals
fothy om nutnrous nough to at a group sltiv advntag.
A sond rason for th lips of group sltion xplanations is th im-
pat of William amilton's famous pprs (|964a,1964) dsribing what
has om to known as kin sltion. amilton argud that if altruisti -
havior is gntially asd, thn organisms that wr altruisti toward rla-
tivs,with whom thy shr gnti matrial, would promot th sprad of
altruisti gns through th nfits to th rlativs that rry thos gns.
With this xplanation, group sltion was not ndd to plain altruism.
From a gn's-yviw it would b sltivly nutral for a mothr to sri-
fihrslf for th livs of two hildrn aus ah hild possssson-half
of hr gns.But th nfit of saving thr offspring mor than ompnsats
for th ost of slf-sarifi.
Kin sltion apPars ommonly in th biologial world (just think of
mothrsdfnding thir young). On potntial prolm for th kin sltion
planationis that it rquirs organisms to idntify not only family mmrs
but lgr fmilial rlationships, eus losr lativs ar mor valual
than distant ons. \ihil many organisms n idntify thi kin through vi-
sual or olfatory us, th kin sltion argumnt works vn if othr us ar
fftivproxis for rlatdnss. A huristi rul lik ..shar with organisms
that ar los y'' will work whn family mmrs tnd to aggrgat in spa.
Suhaggrgation an ou vn in animals without advand soial systms
if disprsal from th hom sit is limitd.
But how an w plain altruisti havior twnorganisms that ar not
rlatdif group sltionis not th answr? A proposd solution to this qus-
tion was artiulatd y Rort Trivrs (|97|) and has om to known as
riproal altruism. Aording to Trivrs, an organism havs altruistially
toward othr organisms in th ptation that othrs will hlp it in rturn.
This xplanation dos not prSuppos a onsious dision o vn mntal
statsin animals. A standard ampl of iproal altruism is found in th
haviorof vampi ats. Vampir ats an surviv only a day or two with-
out fding;howvr, vampir ats that find pry an fed in xss of thir
individual nds. Vampir ats will oftn shar mor, and mor oftn, with
batswho hav rntly shard with thm.
410 ltruism

Riproal altruism has th advantag ovr kin sltion in xplaining al-

Alvaz, Waltr (b.
truisti bhavior toward unrlatd individuals. on qustion, howvr, is
.Walter
whthr riproal altruism is rally altruisti. It ould e argud that th al- Lvarez, th gol
truist is not paying a ral ost ut only dlaying its rward fr hlping othrs varez and grandson of r
(imagin a snario whr two businsssooprat with ah othr in ordr was orn otor 3, 19
to maimiz thi own utility). This prolm and sris of mor ompli- arlton ollg io |96,
atd fats of altruism, spially riproal altruism, hav bn plord tation on th strutu o
xtnsivly y thortiians using gam thory, and gam thorti trat- arhologial gology' v.
mnts hv om a fertil ara fo dvloping nw idas about altruism ing to th study of mass .
and making nw prditions aout havior in ntur. (Plass th alpha- Univrsity of CaIiforni
..Gam honors, inluding th G
til ntry thory'' fo a mor thorough tratmntof th issu.)
Biologial altruism an usd s a foundation for studying psyhologial ltion to th Natinal
altruism. Th major diffrn twn thm is that altruisti traits manifst Arts and Sins.In 19
thmslvsas psyhologial traits in humans, as opposd to ing prsntd makr namd a minor
as haviorl traits in th nonhuman iologial world. Darwin suggstdthat attriuting th nd-rt
.!7idspr
psyhologial altruism, or morality mor gnrlly,ould plaind by th rstrial ojt.
motion of sympathy, whih may hav iologial roots. Similar mphasis on xtnsiv tsting and or
th role of motions in thial havior an found in th work of th ohi- xplanations for vnt
losoph Dvid um. strophism.
Howvr, on nd not hav iologial xplanation of psyhologial al- A l v a r z ' s s i g n a l o n t r
truism. orality, for eampl, may a produt of rationality and not iol- trial auss for xtintr
ogy. Furthrmore' thr ar numr of qustions aout psyhologial altruism Rathr, h gav th th<
that ar not iologial but philosophial in natur. For ampl, dos psyho- allowd arth and plan
logial ltruism ist t all? goism, th viw that all hviors hav slfish susidiary lrypothssg
roots' is dirtly opposd to psyhologial altuism. In this viw, individuals First pulishd in 19
who fd th starving masss ar slfish aus it givs thm plasur and anomlous dta ollt
.W..
slf-satisfation to do so. Howvr, Aristotl (among othrs) notd that sim. 1980; s lso Alvar
ply aus a feling of stisfation ompanis ..ltruisti'' ats, it dos not long rgardd th tin
follow that th at Was prtormd in ordr to ahiv th dsird {lin. erthlss,in gologialti
nt rapidity ould hav t
|BLloGRAPHY at th nd of th rta
Darwin, . 1871.Th Dsntof . [-ondon: sdimnt would rprs
John urray.
amilton,\W.D.1,964a. Th gntialvolutionof soilhaviourI,Jollml of masuing th amount .
ThoretiIBiolrg7: 7-16. oundary layr. Iridir'lm
|964. Th gntialvolutionof soil haviourII.Jourl ofThoreticl most ntirly asntin t
Biolog 7: 1'7-52.
rain of osmi dust fro
R o s n g^,. \ 9 9 2 . A l t u i s m : T h o r t i oln t x t sI.n I ] . F . K l l r a n d . A . L l o y d ,
K-T oundry sdimn
eds.,K1,ords in uolutiorBkllog,19-28. amridg,A: Hrvrd
UnivrsityPrss. sdimnttin. Sampls
Sor,., and D. S. Wilson. 1998' Uto otrs: The t,olutiod Pshologof bio. ltaly. showdan i
Usel|ishBhuior.amridg,MA: arvard UnivrsityPrss. slowd sdimntation. l
Trivrs,R, L. t971, Th volutionof riprolaltruism.Th Qrtrl Reuit,l, of an xtratrrstrialimpa
Biolog46: 5-57.
Alvarz and his grou
\illims,G, , 1966. Adpttiod Nturl Sltil. Printon,NJ: rinton
UnivrsitPrss. -1.2. and ln ihel, stru
gologial vidnthat
th nd of th Crtaot
ags of trrstrial and r

r kin sltion in plaining al-
rls. on qustion. howv. is
:.It ould argud that th al-
ng its rward for hlping othrs
)pratwith h othr in ordr
n and a sris of mor ompli-
Ll altruism, hav n xplord
'ory, nd gam thorti trat-
lping nw idas aout altruism
in natur.(lass th alpha-
ough tratmntof th issu.)
ation for studying psyhologial
r is that altruisti traits manifst
i, as opposd to ing prsntd
:al world. Darwin suggstd that
rally,ould xplaind y th
'gial roots. Similar mphasis on
found in th work of th phi-
planation of psyhologial al-
ldut of rationality and not biol.
ions aout psyhologial altruism
atur.For ampl, dos psyho-
w that all haviors hav slfish
ltruism. In this viw, individuals
:aus it givs thm plasur and
(among othrs) notd that sim.
..altruisti'' ats, it dos not
tnis
ahiv th dsird fling.
Johnurray.
of soialhaviour|. Jourl of
haviour1I.Jourl of ThoretiI
: x r sI. n . F . l l r n d . A . L l o y d ,
-28. amridg,A: Harvard
h uolutiod Pscbolog of
LrdUnivrsityPrss.
altruism.Th Qrtrl Reuieu,,of
*J' n.'n..Ii.
Printn,
Slctio'
lurez 41 1
lr|varcz,.Wltr(. t9 40)
!altr,|varez,th gologist son of Nol Priz_winning physiist Luis Al-
.lfaltr
varez ad grandson of wll-known alifornia physiian . Alvarz,
was orn Otobr , 1940, in Brkly, California. H arnd his BA from
arlton ollg io |962 and his PhD from Printon ln 1967 with a dissr-
tation on th strutur of th Ands ountains. |arez workd in orogny'
arhlogial gology' volanis' ttonis, and palomagntism for turn-
ing to th study of mass tintions.H hs spnt th ulk of his arr at th
Univrsity of alifornia at Brkly and has rivd numrous awrds and
honors, inluding th Gologial Soity of Amria's Pnrose dal, and
ltionto th National Aadmy of Sins and th Amrin Aadmy of
Arts and Sins.In 1985, th plantary sintistsGn and arolyn Sho-
makr namd a minor plant for him. |arez is st known for his thory
attriuting th nd-rtaous mass xtintion to th impat of an tratr-
.\Widsprad
rstrial ojt. rspt for how th thory hs hld up undr
tnsivtsting and orrtion has olstrd aptan of othr nongradual
planations for vnts in volutionary history and promotd noata-
strophism.
Alvarz's signal ntriution lis not in originating th ida of xtratrrs.
trial auss for xtintion-suh suggstions go ak hundrds of yars.
Rathr, h gav th thory a logial strutur and vidntiary as that has
allowd arth and plantary sintists as wll as iologists to tst th many
subsidiaryhypothss gnratd.
First pulishd in 1980 ln Scie, th Alvarz argumnt grw out of
anomalous dat olltd to tst diffrnt hypothsis (L. !. Alvarz t al.
.s7.
1980;s also |varz et a|. 1984|. Th palontologial ommunity had
long rgarddth tintion of dinosaur linagsas a gradual pross. Nv-
erthlss,in gologial tim thir dmis appard rathr arupt. This appar-
ntrapidity ould hav n plaind by unusually slow sdimntatron rats
at th nd of th rtaous. trfsdimntation ws vry slow, a thin layr of
sdimntwould rprsnt a larg sli of tim. To tst this, Alvarz proposd
masuringth amount of iridium prsent in th rtaous-Trtiary (K-T)
oundary layr. Iridium' a vry havy lmnt of th platinum group, is al-
mostntirly asnt in th arth's rust; th only sour is th tiny ut stady
rain of osmi dust from spa. An inras in th prsn of iridium in th
K-T oundary sdimnts would indiat a slowr than usual rt of arth
sdimntation.Sampls takn in |979 from th posd K-T layr nar Gu-
io, Italy, showd an iridium spik far in ss of anything attriutl to
slowdsdimntation. Aordingly, |varez invstigatd th possiility that
an tratrrstrialimpat had ontriutd th iridium.
Alvarz and his group' inluding Luis Alvarz and hmists Frank Asaro
z. and Hln ihl' stuturd thir argumnt in thr parts. Thy mustrd
gologialvidn that an astroid or omt had indd struk th arth t
thnd of th rtaous.Thn' thy assmldth as that numrous lin-
agsof trrstrial and marin organisms had indd om xtint at that
412 Amphibis

tim. Having stalishd th two vnts and thir synhrony, thy laoratd
th ologial impliations-that a larg impat hd ausd a dust loud to
nirl th arth, utting off photosynthsis and making it old and dark for
a suffiintlv long piod so that virtuallv all land animals ov 50 pounds
did' along with som 7 5" of marin organisms-and drw th infrn
that th impat had ausd th tintion of a larg fration of spis 65
million years ago' inluding th tintion of dinosaurs.
Alvarz's framing of th issus providd a road asis for analysis in di-
vrs filds. His original artil has n itd thousands of tims in hundrds
of sintifi journals from iology and gology to mdiin and nular
physis. Th original hypothsis has bn strngthnd y furthr disovris,
most dramatially by dirt vidn of th impat providd y th Chixu.
lub ratr and its uniqu hmial signatur. Furthr, th thory has apturd
th popular imgination, inrasing intrst in sintifi invstigation and
promoting undrstanding of th unity of sin. 1arezhas himslf writtn
a popular aount of th disovry of th tratrrstrial impat and its im-
pliations for lif on arth in . Re.ld the rter of Doom (1,997).

B|LloGRAPHY
Th astrnspadfo
Alvarz, L..v.'.!(/.|varz,F. Asaro, and H. V. Mihi. 1980. xtratrrstrialaus notid inhaitantof t
for th rtaous-Trtiary tintion'Si208: 1095_1108. plosivly in th spri
Alvarz, \/. 1997. T. R d th rtr of Doon. Printon'NJ: Printn grow rapidly and mt
UnivrsityPrss. fd on insts nd ott
Alvarz, !.' . G. Kauffmann' F. Surlyk, L. !. Alvrz, F. Asaro, and H. V. ihl. intl sandy soil ttl wa
1984. Impatthoryof msstintionsand th invrtratfossil rord. is part of an old lina
Siee 22:,l',l' 5 -1'1'4 1'. -P.NI.P,
Stats,whlstadpol
dsrtin s littl as 10

Amphiians
Limd vrtbrats aros mor tan 37 5 million yars ago in th Dvonian Most salmandrs,
priod and thy ar gnrally alld amphiians. This was a vry divrs as- Although amphiian
smlag of aquati and trrstrial animals, ranging from th siz of a sala- ntr watr.
mandr to largr than an alligator. Som of thm vn ame limlss. But Frogs (Anura), wit
by th nd of th Palozoi priod and th grat Prmian tintion, aout tail, and wll dvlop
300 million yars ago' most of th linags Wr tint. A fw thrivd in th sudivisions. Many 1
arly sozoi priod for thy, too, disappard. Rnt amphiians and adults, thy ath pr
th amniots rprsnt th living dsndantsof this arly radiation. Thr ar toris, ut th most g
thr vry diffrnt kinds of living amphiians. Prhaps th most familia and in ponds in th spri
rtainly th most numrous and widsprad ar th frogs, whih ar found qualitis of th mal
on vry ontinnt pt Antartia and vn on a fw oani islands (s whih ar gnrallyl
figur). Th salamandrs ar wll known to inhaitants of th north rmprat tiv tadpol, a larva
zon, ut only on linage has ahivd any suss in th tropis. In ontrast' sason' during whih
th rlativly unfamiliar and srtiv ailians ar rstritd to tropial r- alga or vgttion.I
glons. Tadpols may liv m.
Rnt amphiians diffr dramatially fom ah othr in strutur and way any frogs hav no
of lif. All hv moist skins, howvr, and rspiration is largly utanous. that dvlop dirtly
 mphibins 41

thirsynhrony,thy laoratd
pat had ausd a dst loud to
and making it old and dark for
'll land animals ovr 50 pounds
rnisms_and drew th infrn
lf a larg fration of spis 65
f dinosaurs.
a road asis fr analysis in di-
l thousandsof tims in hundrds
ology t mdiin and nular
ngthndy furthr disovis,
impat povidd y th hiu-
Furthr, th thory has apturd
;t in sintifi invstigation and
n.Alvaz has himslf writtn
tratrrstialimpat and its im-
l rtr of Doom (|997).

h astnspdfoot toad, Scphiopus bolbrookii, is ommorr u littl_

ihl. 1980.xtratrrstrialus notidinhaitantlf th southastrnUnitd Stats'Spadfoottoads rd
r c 2 0 8 . . 1 0 9 5 _ 1 1 0 8 . plosivlyin th spingin phmralponds aftr havyrains;th tadpols
,z. Printon, NJ: Printn grow ra;lidlyand mtamorphosin aout thwks.Th juvnilsand adults
fdlrrinstsr-rdtlthrsmll nimals in th littr layr of forstsarrdurrow
Alvarz,F. Asaro,and H. V. ihl. into sandy soil to wait th nt opplrtr-rnity
to mrrg and rd.This spts
fossil rord.
d th invrtrat is art of an old linagof frogs with rprsntativs in th southwstrnUnitd
-..P.
Stats,r.l'lros
tdpolsan o-rplt dvlopmntin phmralpools in th
dsrtin s littl as 10 dys.

rillion yars ago in th Dvonian ost slmandrs, a fw fogs, and on ilin lak lungs ntiely.
ians. This was a vry divrs as- Although amphiians ar assoiatdwith moisr haitats' many spisnvr
nrwatr.
' ranging from th siz of a sala-
f thm vn am limlss. But Frogs (Anura), with mo than 5,500 spis,hav vry short odis. no
) grat Prmian tintion, aout tail, nlwll dvllpdlims, spiallylong hind lims tlrt hav for main
Wrxtint. A fw thrivd in th sudivisions.ay frogs ar apal of prodigious laps. arnivorous as
rppard.Rnt amphiians and adults,thy ath pry with projtil tongus. Fogs display divrs life his-
:sof this arly radiation. Thr ar toris'ut th most gnral nd familiar is on in wlrih rh ssongrgat
rns.Prhaps th most familia and in porrds in th spring, witlr fmals slting mals as mts asd on th
rd r th frogs, whih ar found qualitis of th ml all. Mating involvs xtrnal frtilization of ggs,
:vnon a fw oani islands (s whih ar gnrallv Iaid in lustrs lr strings. ggs dvlop into th distin_
inhaitantsof th nrth tmprat tiv tadpol, a lrval forrrr uniqu to rogs.Tdpols typially liv for on
sussin th tropis. In ontrast, sson'during whih tim thy onsum primary produtivity in th form of
ilians ar rstritdto tropial r- algaor vgtation'Howvr, thr r many varitions on this lif lristory.
adpolsmay liv more than on yar for nrtamorphosinginto froglts.
lffIaChothr in strutur and wa ny frogs hav nl larval stag at all ut ly larg ggs in small numrs
l rspiration is largly utanous. that dvlop dirtly into miniaturs of th adult. othr frogs hav tadpols
414 Amphibis

that may roodd in spial ompartmnts tht form in th skin of th of th skin and lads to dt
bak, in spializd pouhs on th ak, in th voal sas of mals, or in l i n s i n s p i sl i v i n g i n 1
divrs ologial settingssuh s aroral romliads.A fw frogs ar vivip. California, whr pidm
arous' with ggs dvloping in th fml rprodutiv trat' whr thy r- idly, and thr is no know
ivnourishmnt. pliatd in th dlins
Salamandrs (audata), with mor than 560 spis'rsmlanint am- fators inlud ffts of
phiians mor losly than do frogs or ailians. Thy ar gnralizd in tion from pstiids and
strutur' with a ody of modrat lngth, a wll-dvlopd tail, and two thse fators and othrs
pairs f lims, with th hind lims ing only a littl longr than th for- many parts of th plant
lims. ads of salamandrs ar rlativly smllr than thos of frogs, ut zlmnt ovr th dlins.
oth groups hav wll-dvlopdys and llnt vision. Salamandrsof- Today's living amphiia
tn gathr in ponds to rd ut mls hav no alls. Larva ar arnivorous jor group an trad a
rathr than hrivorous) and mtamorphosis is far lss dramati than in to dliat, thy wr
frogs. Som spis rmain in prmanntly larval or smilarval stat and trouling that amphibran
som lrva om vry larg. ost lads f salamandrshav som vari- should faing xtintlo
tion of th asi lif histoy, ut th most sussfulnd numrorrssalaman-
drs, in th family Plthodontida, hav dirt dvlopmnt' and it is only |BLIoGRAPHY
ths that hav sussfully invadd th tropis (rstritd to tropial Amr- Dullman,.W.., and L. Tru
ia). A fw spisof salamndrsar viviprous. Hill.
C a i l i a n s ( G y m n o p h i o n a ) a r l i m l s sa n d x t r m l y l o n g a t ,w i t h i - Huthins, N4.'!. . Duilm
lopdi.2nd d.V
thr a vry short tail or no tail. Th had is aout th sam diametr as th
Lnnoo, M. 2005.mphibi
strongly sgmntd trunk, and th ys ar inonspiutlus or invisil. A Spis.Brkly:Univ
uniqu snsory organ, th tntl, is formd from omponnts of th nos Stuart,S. N., J. S. hanson,J
.!7
and y and provids nvironmntal information to ths mainly urrow- Fishman,nd R. !(.
ing animals. Som ailins hav aquati larva ut m()st hav ithr di. xtintionsworldwid.j
.Wlls,
r t d V l o p m n to r a r v i v i p a r o u s .A m o n g t h m n y v i v i p o u ss p i s K. D. 207.be col
of hiagoPrss.
a r f o r m s t h a t a r l u s i v l y q u a t i . T h r a r a p p r o i m t l y 1 7 0 a -
ilian spis.
Amphibians play imprtant ut gnrally undrappriatd rols in
Andrson, dgar (18
osystms.Frogs ar important prdators as adults, ut thir larva an b
important onsumrs in equati systms.Rmoval of tadpls from strams dgar Andrson is rgard
an quikly ld to ovrgrowths f alga and forration of larg algal mats. tionary iology during th
Although salamandrs ar rypti and lss ovious than frogs, in som intlltual traditions, Anr
osystms thy an trmly numrous. As arnivors, thy ontriut in gntis and ytology t
to rgulation of trrstrialfood ws. Mmrs of th soil ommunity' a- systmatis.H is stkn
ilians ar major onsumrs of arthworms, ut thir ologial rol is lss natural sttings and for
wll undrstood than that of frogs and salamandrs. hyridization, a pross;
Amphiians hav rivd attntion ausof onrns aout vidnt d- hyridization and ak
lins and tintions lf populations as wll as spis throughout th l949 with th puliatio
world. A rnt assssmntof th sttus of amphiian spis aross th tio. Te ook was widl
glo found that a highr proportion (mor than on-third) Wr at risk of ontriution to plant gn
tintion than for any of th othr vrtrat taa. spially trouling volution.
Was th finding that a larg prntagof th dlins Wr from unknown Andrson, orn in For
.Whil
auss. most amphiian dlins doutlss ar rlatd dirtly to d m i n i s t t o ra n d a p i a n i
habitat dstrution nd modifiation y humans, th ar also othr implr- th }IihiganAgriultural
tant fators. On inftious disas,a hytrid fungus that ttaks th kratin Andrson was thr. Fro
 Aderso 415
s that forrn in th skin of th of th skin and lads to dhydration, is rsponsilfor wll-doumnted
d-
th voai sas of mals, or in linsin spisliving in prottd ars in ntral Amia,
Australia, and
l m l i d sA. f w f r o g s a r v i v i p - alifonia, wh pidmis hav n rordd. Th disas
sprads rap-
'rodutivtrat, whr thy r- idly, and thr is no known Way to stop it. Gloal limat .h".'g.
lso is im-
pliatd in th dlins of many frogs in tropial montan
forsts. Othr
0 s p i sr' s m l a n i n t a t n - fators inlud ffets of introdud spis (.g., for
sport fishing), pollu-
:ilians.Thy ar gnralizd in tion from pstiids and frtilizrs, and synrgisti intrations
of som of
wll.dvlopdtail, and two ths fatos and othrs suh as ultraviolt adiation (UVB).
Howvr, in
ly rr littl lorrg than th for- many parts of th plant amphiians rmain aundant, adding
to the puz-
mallr than thos of frogs, but zlmntvr th d1rns'
tllntvision. Salmandrs of. Tday's living amphiians ar rmnants of a vry anint linag;
ah ma-
l t l i l s ' L v r r n i v < l r o u s jor group an trad ak out 200 million
yars. Altho,.gh ,.,y appar
;is is far lss dramti thn in to dliat, thy wr ruggd survivos of many *tin.tio.,
vnts. lt is
y larvirl r smilarval stat and troulingthat amphiians that lived happily with th long-tint
dinosaurs
l [ s a l r n d rhse v s o m v r i - should b faing xtintion on or wath'
;ssflnd nnroussalatnan-
t dvlopnt' and it is only BlLloGRAPHY
lis (rstritdto tropial Amr- .W.
Dullman, ., and L. Tru. 1986.Biolog of Amphibin's.Nw
York: Grw.
fous. Hill.
,nd xtrmly lorrgat,with i- Huthins,M., ). . Dullman,and N. Shlagr.20.Grzimek's
niml Li|
clopedi.2nd ed.o|. 6, phibins. Farmlngtonills, I;
a b o u tt h s a m d i a m t r a s t h Gl Group.
Lannoo,. 200.'. mphibi Dclines:Th onseruti-l Sttusof (Jitd Stts
inlrrspiuousor invisil. A Spis.Brkly:Univsityof alifonia Prss.
:d fror omponllts of th nos Stuart,S. N.' J. s. hanson,N. A. o' B. . Young,A. s. L. Rodrigus,
D. L.
ation to ths mainly urrow. Fishman'nd R. W. Y/allr.2004. Statusand1rndsof amphiin
dlinsand
larva but most hav ithr di- xtintionsworldwid.Sice6 1783-1786.
Wlls,K. D.207. h olog d Behuiorof mphibin.s'hiago:
n g ,r h n l r r yv i v i p a r o u s s p i s Unrvrsity
of hiagoPss.
r ar approximatly 170 a- D.B.w.

:ally Lrndrappritdrls in
dgar(ts97-|969)
Andrson,
s adults, ut thir larva an
movl of tadpols from strams dgar Andrson is rgardd as on of th lading sintists
of plant volu-
rd formtin of lag algal mats. tionry iology during th twntith ntury. A produt of
numr of divrs
ss ovious rharr frogs. in sonl intlltuaitraditions, Andson ws al to synthsiz nwr
dvlopmnts
Ls.As arnivors' thy ontriut in gntisand ytology to hlp a.ddrssmor traditional prolms
in plant
r r s [ t h s i l o m m u n i t y , a - systematis. H is st known for his invntiv studismasuring variation
-i't.og..,,'u.in
;, ut thir ologil rol is lss naturl settings and fo his artiulation of th onpt
of
mandrs. hyridizatin' a pross y whih nw gnti matrial is introdud
through
usof onrnsaout vidnt d- hybridization and akrossing. Muh of this work
was formalizd in
wll as spis throughout th 1949 with the publiation of smll volum tit\ed Itrogrssiu
bridiz-
of mphiian spis aross th tion. The ook was widly rd and immdiatly .og.'i,.d
as an original
than n-third) wr at risk of ontrition to plant gntis that ould shd light o' , g..'","l
thory of
:rattaa' spially toubling volution.
:hdiinsw fronr unknown Andson, on in Forstvill, Nw York, ws th son of
an duational
loutlssr rlatd dirtly to administrator and a pianist. His fathr am a prfssor
of dairy sin at
n n s .t h r r I s o o t h r i m p o r - thihigan Agriultual ollg and movd his family to
ast Lansing whn
id fungusthat ttirks tlr krtin Andson was thre. Fronr n arly g, Andsn ws fasintd
y plnts.
416 ndrson

H was a kn studnt and in 1914 ntrd Mihigan Agriultural ollg' ILloGRAPHY

whr h studid otany and hortiultur. In |9|9 he ntrd th gradut Andrson' . 1'949.ltrlgr
program at Harvard, working at th Bussy Institution under th suprvision Klinman'K. 1'999.is ow
of notd agriultural gntiist dward Murray ast. His rsarh Was on th thory in th 1940s./
gntis of slf-inompatibility in Nicotin' th toao plant, ut h ws Smoovitis,V. B. 1999.d1
eds',AtrirtNtio
also fond of aras lik onomi otany and gravitatd to Oaks Ames' th
UrivsityPrss.
notd onomi otanist at Harvard and prt on orhids. Aftr gradua. Stins,G. L. 1978.dga
tion, Andrson aptd a position as gntiist at th issouri Botanial moirs 49 -23.
Gardn and was appointd assistant profssor at Washington Univrsiry in
St. Louis. |n 1929 h aptd a National Resarh Fllowship to study in
Britain's John Inns Hortiultural Institution, working with ytogntiist
AnimI Specisn
. D. Drlington' statistiian R. A. Fishr, and gntiistJ. B. S. Haldan.
Aftr rturning from Britain, h was inflund y Harvard gntiist A. J. Cain's imI Sp
Karl Sax. first omprhnsiv dis
uh of Andrson's rsarhwas spnt dvloping novl visual and quan- ring of th so-lld mt
titativ mthods to masur gographi variation in natural populations, th iogography' taxonom.
st known of whih is th idogranr. Anderson is also known for his d- ohsiv tatmnt of s
taild studis of th pattrns of variation in th ommon lu f|ag, Iris ursi- and th rols of natural
olor, a|ong th ississippi dlta and what th pattrns rvald aut t r n s o f h a r a t rv a r i t
mehanisms of spiation in plants. This study rivd reognition in th ooks; howvr, it was l
ear| 1940s, an intrval of tim important in th history of volutionary i- thsis that mad ain's l
ology, baus it fuld th movmnt known s ..nw systmatis'' (or prsntd in this slim v
biosystmatis),whih sought to ring intrdisiplinary insights from gn- oaks and from s urlri
tis and ology to mor traditional taxonomy. As a rsult, Andrson in tras an ar fr
mrgd as on of th ladrs sking gnral prinipls of plant volution. ussing traditional ida
H was invitd to giv th prstigiousJsup Lturs at olumia Univrsity tion, and taxonmi rar
along with avian systmatistrnst Mayr in 1941. Andrson Was to provid
th plant sid of th nw systmatis.Although h gav th lturs,h did
not onrplt th rquird mnusript for a ook asd on thm. Th om-
pltion of this manusript would hav plad Andrson within th irl of
arhitts of th volutionary synthsis of twntith-ntury volutionary i- . o:
ology. Th rol of otanial arhitt vntually fll to his los frind, G.
Ldyard Stbins.
Andrson's failr to omplt th manusript was part of his rrti pr-
forman pattrn. H was nononformist who fquntly ngagd in varid
.#
intrsts that omptd fo his tim. did mak notal ontribuions to
aras in hortiultur and vn am th dirtor of th Missouri Botanil
P. itnotis
Gardn. In th 1940s, h inrasingly turnd to qustions important to un-
drstanding hyridization in maiz, with his intrstsditd toward o-
nomi otany.
A talntd individual with a strong prsonlity, Andrson lft his imprint
on th history of plant volution. For th lattr part of his lif h was
plagud with m<ltionalpolms for whih h ws hospitalizd. H was
marrid to Dorothy Moor' a fllow otnist, and had on hild. H did in T h g r a d u a l v a r i a t i nt l
lvland Avnu Gathous, loatd on th grounds of th Missouri Botanial rkn s on trf th sr
Gardn. i
g o g r a p h i a vl r i t i l n

c ihigan Agriultual Collg,
In t91r9 h ntrd th graduat
l n s i t u t i o nu n d t h s u p r v i s i o n
:ray ast. His rsarh was on th
' th toao plant, ut h was
rd gravitatd to Oaks Ams' th
pt on orhids. Aftr gradu-
ltiistat th issori Botanial
.!7ashington
ssor at
Rsarh Fllowship to study in
.ion'working with ytogntiist
., and gntiist B. S. Haldan.
J'
rflnd y Harvad gntiist
dvlopingnovl visual and quan-
riation in ntral populations, th
tdrsonis also known for his d-
n th ommon lu flag, Iris uersi-
yhat th pattrns rvald aout
study rivd rognition in th
in th history of volutionary bi-
..nW systmatiS'' (or
known as
rdisiplinary insights fm gn-
txonomy. As zr rslt, Andrson
rral prinipls of plant volution.
. p I , t u r as t C o l u m b i a U n i v r s i t y
ln 7941'.Andrson Was to provid
hough h gav th lturs, h did
. a o o k a s do n t h m . T h
ad Andrson within th irl of
twntith.ntury volutionary i-
ntually fll to his los frind, G.
rusiptwas pat of his rrati pr-
st who frquntly ngagd in varid
did mak notal ontriutions to
: dirtor of th issouri Botanial
rnd to qustions important to un-
r his intrstsdirtd toward o.
:sonality,Andrson lft his imprint
th lattr part of his lif h was
lhih h was hospitalizd. H was
anist,and had on hild. H did in
h grounds of th Missouri Botanial
Animal Soisand Thir volution 417
B|LIoGRAPHY
Andson', |949, ItrogrssiuHbridizatkl.Nw York: Wily.
Klinman,K. \999. His own synthsis: orn, dgarAndrson,and volr-rtilnry
thoryin th 1940s.Journl of th istor of Biolog 2:293-320.
Smoovitis,. B. |999. dgarAndrson.In John A. Garratyand Mark . (Jarns,
eds.,meriNtiol Biogrph,vol. 1,452453. Oford: Oford
UnivrsityPrss'
Stbbins,G. L. 1978.dgar Andrson. NtillAdm lfSiesiogrphi:l
emoirs49:-2. -\1B.s.
Univrsity in
Aiml Spiesnd Their uoltio (A.J. ain)
A. J. ain's niml Spcies d Thir uolutiltt (1954) offrd orr of tlr
firstomprhnsivdisussionsof th natur of spisftr th initial flow-
ringof th so.alld modrn synthsis.ain drw ids from palontology,
iogography' taxonomy, and population gntis. H omind thm into
ohsivtatmnt of spis onpts' lassifiation mthods, spiation,
and th rols of natul sltion,history, nd gnti drift in rating pat-
trnsof haratr variation. Sorn of thstopis had n ovrd in alir
books;howvr, it was th omination of topial radth and sarlsssyn-
thsisthat mad ain's tratmntstand apart. Th mpls and as studis
psntdin this slim volum ovrd a staggring rang' from ohids to
oaks and from sa urhins to shws (sfigur).
in trs an r fronr lzrssialidas to novl ons. gins by dis.
ussingtraditional idas of iologial nomnlatu, mthods of lassifia-
tion, and taxonomi rank. ain thn turns to th natur of spis, whih h
o m -
P. ictrotis
hegradulvariationthat n snin adjoiningpopulationsof organismsis
takeas one of th stpisof evidnfor gadualvllution.This pitr'rr,
of
gogphialvritionin Roslls'is tkn from irirr |954,61 .
418 Atonouis

dsrisas olltions of populations that display variation in many hara- in volving tolranto I
trs aross th spis'gographialrang. ain disussshow this variation t i a l r p < l d u r i vis o | t i
is promotd and maintaind in som ass y diffrnt sltion prssurs in mtal-ladn soil (Anton
diffrnt loations and in othrs y gnti drift. H argus that th uiquity was a striking dmnstr
of intraspifi variation of this typ oviats th lassial dfinitions of gn flow twn popt
speis; in fat, he argus that no singl dfinition of spis is adquat fo dnts prformd lgant
th normous natural divrsity of organisms. Cain disusss fiv typs of gntially divrs offsp
spies that rflt fiv diffrnt spis onpts' from th iologial spis in a spatially varial an
onpt hampiond by rnst ar (1942) to what ain alls agamospis, 7984). Whil thoris f<
olltions of similar individuals that rprodu asually. H also disusss suh an advantag,thr
mods of spis formation, offring on of th first attmpts to link diffrnt isly whih ologial l
typs of spis with diffrnt mods of spiation. Th finl haptr of th third, far.rahing pro;
ook is a prsint disussion of sympatri spiation, spiation without a ours of an inftious
gographial sparation twn forms. Th ovrall argumnt is that a attl of pathogn and h
spis is nithr a olltion of idntial individuals nor a fixd ntity' ut pathogns' hosts, and th
rathr a dynami st of populatins that ar wapoints on a sris of volu- tions aross th landsp
tionary rrajtoris. w o t h m s m g f
If th ook's mssag is obvious to th modrn radr, it is aus th strong for in natur t
book mad so prsuasiva as. ain's influn would tnd from oiogial physiologial strssof h
gntiists to systmatists. Svrl topis raisd in th ook, suh as th adaptatiln-mutations 1
adquay of th iologial spis onept or th inadquay of rtain mth- mon mong plnt sp
ods in systmati infrn for apturing th rlationships among sts of orhr agnts of slti
volving spis' would rmrg rpatdly ovr th nsuing yars. Although plants to prdatls and
ain's onis tratmnts of ths topis is oviously not informd y th molds diversity, on mus
walth of data now at our ommand' his tratmnts rmain among th most of sltion an vary and
luid and ritial disussions of many important qustions onfronting volu- mov radily. Th intr
tionary biologists who think aout what a spisis and how spisar r- mrgs in spa and, l
latd to on anothr. Ths thmes ar mr
tionary iology ar intr
B|LIoGRAPY In addition to ths r
thord a numr of high
ain, A. I. 1954. iml Spcisd Thir uollttio.Londn: Huthinson's
UnivrsityLibrary. portant topi, posd n
ayr, . 1942. Sstemticsd th rigi of Spis.Nw York: olumia dirtions. Ths paprs
UnivrsityPrss. -I,. (Antonovis 1976), te d
(Lokhart et aL. 7996),
among intrating spl
Antonovis, Janis (. 1942) ology has inflund rnnt
Whil th study of plnts has shapd many fats of volutionary iology. influn is visil throug
fw sintists hav usd plants to illuminat so mny important issus as has
Janis Antonovis. Antonovis has ld a sris of rsarh programs in natural BlLloGRAPHY
plant populations that hav unovrd som of th myriad ways in whih
^ I , ' . ' ^ - | ^, . l t 7 ,'- l r
natural seltion shaps th faturs of organisms nd the divrsity of thir ! r r . r l r . r

dynmis lf nthr-st
populations.
uirll. llurl of
Thr ontributions stand out among many. His arly work, prformd Alrtnovis' J. 1976. h n
.!0'als,
with his dotoral advisor at th Univrsity Collg of North Anthony B ltil Grd 6: 2'
Bradshaw, and ollagus,showd how populations of plants on min tailings, 2003. Towrd t-l

isplayvariation in man hara-
s ow this vrition
i n d i s u s s h
y diffrntsltionprssurs in
rift' I-Iargus that th uiquity
lts th lassial dfinitions of
nitiorr of spis is adquat for
rs. ain disusssfiv typs of
pts' from th biologial spis
o w h a t a i n | l sa g a m o s p
Luasxually.H also disusss
h first attmpts to link diffrnt
iation. Th final haptr of th
spiation,spiation without a
.h
ovrall 2lrgumnt is that a
dividuls nor a fixd ntity, but
: waypoints on a sris of volu-
odrn radr, it is baus th
n would xtnd from ologial
:aisdin th ook, suh s th
r th inadquayof rtain mth-
t h l t i o n s h i p s m o n g s t s o f
ovr th nsuingyars. Although
; oviously not informd y th
atmntsrmin among th most
tant qustionsonfronting volu-
spisis and how spis ar -
|tion, London: Huthinson's
leies. Nw York: olumia
ry ftsof volutionary iology,
so many important issus as has
s of sarhprograms in natural
m of th myiad ways in whih
ganismsand th divrsity of thir
nany. is arly work, prformd
.Wals,
ollg of North Anthony
ulations of plarrts on min tailings,
tonouis 419
in volvingtolranto havy mtal dpositsin th soil, lrad also volvd par-
til rprodutivisoltion frol losl adjantpopultions tht wr not on
mtal-ladnsoil (Antonovis and Bradshaw l970; Antonovis 2006). This
was a striking dmonstrationof inipint isolation in th fa of onsidra1
gn flow twn populations' In arrothrprojt, Antonovis and his stu.
dnts priormd lgant primnts to show that individuals that produd
gntilly divrs offspring through sual rprodution had gratr fitnss
in a spatiall,vvarial and ot-rrptitivnvironmnt (Antonovis and llstand
i s ' 1984). Whil thois for th origin of sxul rprodr.rtionhad postulatd
suh an dvntag' thr was littl' if any, mpirial work that addrssd pr-
islywhih ologial faturs of th nvironmnt wor"lldfailitat it. In a
third, far-rahing projt' Antonovis and his studnts hav shown that th
ours of an inftious dis:rsis dtrmind y a alan twn th loal
attlof pthognr-rdhost in an individual population and th movmnt of
pathogns' hosts, nd th gns for viuln nd rsistan among popula-
tions arossth landsap('g.'Alxandr and Antonovis 1988).
Two th-lsmrg from Antonovis's work. First, rrtural sltion is a
strongfor in natur that mrgsfrom a vrity f ologial ftors. h
physiolgialstrssof havy mtals in th soil may sm an ovious agnt of
adptatioll-mutations that onfr tolrn of havy rtals r not om-
mon among plant spis-ut Antonovis and his ollagus hav found
othr agnts of sltin, from soil minral profil to rowding of othr
plants to prdatos and pathogns. Sond, to undstnd how sltion
moldsdivrsity,on must undrstarrdth spatial sal n whih th prssurs
of sltionan vay and th spatial sl at whih individuals (or thi gns)
mov radily. Th intrplay f ths two sals dtrmins how divrsity
mrgsin spa nd, ultimatly, th as With whih spei:ltion ours.
Ths thms ar mrad y Antonovis's thsis that logy and volu-
tionaryiology ar intrtwind disiplins.
In additiln to ths rs:rrhprograms' Arrtonovis hs auth<lrdor oau-
thord a numr of highly influntial ssays' ah of whih xamind an im-
portant topi, posd nw qustions aout it, and inspird nw rsarh
dirtions.Ths paprs irrludd disussions of adptiv gnti varirtion
-1.r. (Antonovis |976I, th dynamis of inftious disassin natural populations
(Lokhart t al. ] 996), and th potntial for riproal gnti influns
mongintratirrgspis (Antonovis 200). is appro:rhto vlltionarybi-
ology has inflund innumral othr sintists for ovr forty yars and that
irrflunis visil tlrroughout th disiplin.
|BLIoGRAPHY
Alarrl,
. N,t.,ndJ. Arltlnovis.
198B.Dis:rssprdnd popul:rtion
dynamisf nthr-smutinftionof Sillb ausdy th furrgusUstilgo
uioltl.
JourIof rllog76:9|-14'
Antonovis,
J.1976. Th ntuf limitst ntualsltion.
ls of th issouri
BotilGrd6: 224_247.
-. 2003.Towrdmmunitygnoris? olog84: 598-601.
120 Arhaoptryx

2006. vlLrtion in losly adjnt plan plpLrltions: Long-trr r-

sistn of prrprodr:tivisolation t mln' lundry' Hcre dit 97:
_37.
Antonvis, J., and A. D. Brdshaw. 1970. vlution in losly djar"rt pl:rnt
poulations. 8. linal pattrnst a min oundrr'.Hrditl'25: 49_62.
Antolovis,J., ad N. . llstand. l984. L.prilntlstudisof th volrrtionary
signifian tlf sual rprodution. l . A tst lf tlr frquny-dpr"rdnt
sltionhypothsis.ullutil38: 103_11.5.
.l
Lokhlrrt, A. B., P. H. Thrall' ndJ. Antorrovis. 996. Suaily rr;rnsmittddisass
in nimals: olr;gialnd volutitlayimpliatilns.Bitll,.lgilRuittls,f th
mbridgPhilosophilSit7 |: 415-471. - . 1 I ,. t .

Arcbeoptr
rhopteryr (anint ird, or Grman Uruogl) is th gnri naln givn
to 10 skltal spimns (as rvll as:r sondry flight fathr disovrdin
1860) of th arlist known ird' from ltst Jurassi lithogrphi limston
dsits kn.,vn as th Slnhofn Limston, loatd lust to th nrtl.i of th
ity of unih in southrn Grmny. Dating to approxinratly 150 rnillion
yars old, th dposits Wr in a hyprslin lilg()on that was llsdoff to
opn sa y orl rfs.Althlugh th disovryof th skltonsspnndsom
h most famous fossi
145 yars, th two most famous spimns ar th first two disovrd, th
disovrdarly in th
L o n d o r r n d B r l i n s p i r n sd
, i s o l ' r di n 1 8 6 l a n d 7 8 7 7 , r s p t i v i y . spinrn,''distlvrdi
Th last rovrd speimns ar th Solnhofn (th largst at 1096 largr
t h a n t h L o n d o n ) , d i s o v r di n a o l l t i o n i n 1 9 8 7 , a n d t h f i n a l s p i -
mn) th Thrmopolis spimn' dsribd in 2005. Althtrgh onsidral
vaition xists, th first svn spilr1nsar grlrallv rfrred tO as a sll.l-
g1 spis' Arhopter1, lithogrphic, w1|eth 199 ampl (Aktin- rPtilian fingrs with
Vrin) is distintiv in many faturs nd is rfrrd to as a nW spis' of modrn trunk-lim
Arhoptr bt,ri.[t is th onl1's1-limn tl show n ossifidstrnum' beoptrl ma,v hv
and th proportions of th legs ar qllit diffrnt from th othrs,lrly in. rchoptry also p
diating aroral haits. arly rtaous mir
Th lbrat]Brlin spimn (s figur) has n lld vrital to rvrsd as a lrallu
Rostta ston of volution aus it is omplt ard utifull1,prsrvd it w;rs lss t hom or
with outstrthd wings. It illustrats n almllst prft intrmdiat - bra had a pir of t
twn two major lasss lf vrtrats:irds and rptils. rhlptr sarum ar quit rpti
fulfills th Dwirrin xptation of :r missing link twn mjor vrt- ag l-riitd by rrro
brt groups. Its lliptial wings in profil ar similar to tl-rosof modrn irds. h skull lf r
woodland ird, with modrn flight fathrsvirtully idntil to thos of liv- tilian tth, ut with
ing irds' vn dlwn to hir tirostrutur' as shown y ltron mi- innr r wr qr.ritsi
rosopy. Fth sttltur' on ill'u'ntd,rrrrind r'rrlrngd in any ability.
major way for 150 million yars. Tl-rprimary and sondary flight fathrs Although rblp
ar nhord to th wing s in rnodrn irds, nd th flight fthrs hiit fossil irds hav n
asymnrtrivans that prodr"rdindivilual airfoils' indiating arldynami t o t h 1 9 7 0 s .M o s t o
fur-rtion.Mst palontologists liv t,atrheopterj' was a pal al- rsnt various typs
though primitiv powrd flir, and th sapula and orid ar primitiv boloris, whih sport
ut join at a 90" angl' as in mldrn irds. Th avian wings nd with thr muh lik tht o[ th l

t oulations : Long-trm pr-
n oundary. rdit 97:
ion in loslyadjrrtplant
ndar.Hrdit 25 49_62.
m n t x |\ 1 t l L l i os[ t h
of th frquny-dpendnt
1996.Suallytrnsmittd disass
itins. BitiogiI Ruil'l,s lf th
'1.
,uogel)ls th gnri nam givn
.dary flight fathr disovrd in
;tJurassilithographi limston
l, loatdjust to th north of th
lg to approximatly 150 million
l lgoo that was losd off to
:ry of th skltonsspannd som
ar th first two disovrd' th
in 1861 and 1'877, rsptivly.
rofn (th largst at 10o largr
on in 1 987, and th final spi-
in 2005. Although onsidral
r r g l r r a l l y f r dt
whil th 199 ampl (Aktin-
, i s f r r dt ( ) a 5 J n w s p
mn to show an ossifid strnum'
.frntfrom th othrs, lrly in.
,u)hs n lld a vrital
mplt and autifully prsrvd
lmost prft intrmdiat b-
,irds nd rptilrs. rhopter
i s s i n g I i n k l r w nm j o v r t -
ar similar to thos of modrn
; virtually idntial to thos of liv-
tur' as shown y lton mt-
d, rmaind unhangd in any
nary and sondary flight fathrs
:ds,and th flight fathrs hibit
r1airfoils, indiating arodynami
t rhoptry,ws a apabl al-
:apula and oraoid ar primitiv
s. Th avian wings nd with thr
Arhaoptry 121
v o l u t i n r y
-rT
Th most fmous fossil of thm all, rheoptr lithogrpbi, A,rst
disovrdrly in th 1860s in Solnhofn,Gmany.This is th ..Brlin
spimn,''disvrdin 1877.
s s i n -
rptilianfingrs with sharp' highly rurvd laws that losly mimi thos
i s . of modrn trunk-liming mammls and woodpkers, indiating that Ar-
choptermay lrav n a trunk limbr. Thr is rnt vidn that
rchlptrxlso possssdhandlg wings, s ar prsnt in th hins
arlyrtaousmiroraptors. Th Uruogl's ft ar irdlik, with th first
torvrsdas a hallu and highly rurvd pdal laws. A apal prhr,
it was lss at hom on th ground. lts long rptilian tail of 2 audal vrt-
bra had a pair of til fathrs oming off ah vrtra. Th trunk and
sarumar quit rptilian. lt lakd th inding uninat prosssof th ri
aghiitd y modrn irds, ut th puis was rtrovrtdas in modrn
irds.Th skull of rchoptery,Was oth rptilian and avian, having rp-
tiliantth' ut with an avian qudrat and rnil kinsis. Th rain and
innrar wr quit similar to modrn irds, indiating a sophistiatdflight
aility.
Although rbeoptryx rmains th oldst known ird, more Mesozoi
fossilids hav n disovrdin th last two dadstharr from th 1860s
to th 1970s. Most om from th l-owr rtaous of hina, and thy rp-
rsntvarious typS of primitiv irds, inluding th rchoptr-|ike l-
holoris'whih sports a long tail with 27 auda|vrtr,nd an anatomy
muhlik that of th famous Grman Ilruo'!l.
422 ristotl

|LIoGRAPHY Th surviving orpu

lzanorvski, ^.202. Arhaoptrygida(Uppr Jurssi of Grmany). In L. M. on a first ntury B..
.Witm,
hipp and L. . eds., sozli Birds: blu th Hds of dat to th lat fiftn
Di<lsrs'129_|59. Bkly:Univrsity f aliftlrni Prss. subjts, ut for th pu
Fduia' ^. |999. h rigill d uolutil of Birds. Nw Havn, T: Yal whih onstitut 25u%
Univrsity Prss.
Fdui, A., L. D. Martin' and S. arsitno. 2007 ' rhopter 207: Ql
V]is?Th uk 124: 7.]-380. ARlsoL's P}
I-ongrih,N. 2006. Strutur urrrdfuntion of th hindlim fathrsin
Archeoptr lithogrphi. Plobiolog 2: 4\74|' Aristotl was l to
ayr, G., B. Pohl, nd D. S. Ptrs.2005. A wll-prsrvdArhoptrx spilnn dal of thought to th
with thropod firturs.Si 310: 1483-1486.
from th suprfiial arr
Zotl'' Z.2004. Tlr rigin and rly volutilnlf irds: Disovris,disputs arrd
-.F. ganized sintifi undr
prsptivstrom fossil vidn.Ntt;rtuissshfte 9\:45547|.
of th goal to ahiv
d Postrilr nltis
Aristotl (84_22B...) ts and proposirins
Aristotl is proprly ognizd as th originator of th sintifi study of dg of th ssntialnat
lif. H onsidrd it a ntrl onrn of th thortial study of natur' His first prinipls. Ths dt
nations of :rl1th oth
tratiss Histor of nimals, o tbe Prts of ni?.-lls,and tb Gert-
Th sond ook of
tion lf imls (and a nurr of mor spilizd studis) povid a tho-
goal. Plato had formu
rtial dfns of th propr mthod for iologial invstigation and th
rsults of th first suh investigation.Thr Was nothing of sirilar sop and eno: ither you kno
sophistiation until th ightntlr ntury. Irl 1837 th grt nromist unnssry,or you do
q u i y i s i m p o s s i l A. i
Rihard Own dlard that ..Zoologial Sin sprang from his [Aristo-
the Posterior lti s,
tl's] laours' W may almost say, lik Minrva from th Hd of Jov, in a
stt ()f nol nd splndid raturity'' ()wn 1992' 91I. Bfo amining Thr h argustha
inquiry tht dos not l
this rmarkal irhivmnt.a fw words about its rtor r in ordr.
mation to dirt furthr
familiar phnomna lik
LIFEAND WORK
ut w an s his r
Aristotl was born in Stagir on th northrn Agan oast in 384 ... His anatomy and dvlopm
fathr, Nimahus' Was physiian to King Amyntas of adon, and his ri imlium), n the
moth Was Of walthy family from tlr island of uoa. H was snt at th Grrttio 'f ltir
th ag of 17 to Atlrns' whr h str-rdidin Plato's Adrny for 20 yars' will rfr to tlrsby tht
until Plato's dath in 347. B thn lr had dvlpd his own distintiv
philosophial idas, inluding his passion for th study of natur. H joind a
|oLoG|AL ING
philosophial irl in Assos on th lstof Asia Minor' ut soon movd to
th nary islarrd of Lsos whr lr mt Thophrastus' 21Ongmn with F r A r i s o t I 's y s t n l a t
similar intrstsin natural sin.Btwn th two lf thnr thy oigirratd srvs th gol of usa
th sinof iology, Aristotl arying out a systmtiinvstigationof an- and funtions is fundarn
imals, Thophrastus doing th sam for plants. ati prsnt:rtionof inf<
In 4 Philip II of Madon askd Aristotl to tutor his son Alandr, ut th first instan w shl
y 35 h had rturnd to Athns, now undr Alxandr's ontrol. \/ith ll animals. Aftr w l
Thophrastus h foundd a shool known as th Lyum. H hadd th auss. This is th natur
Lyuirr until Alndr tlr Grt's dth in 2. With anti-donin ir.rquiry (histori) aout
flings running high in Athns, Arist<ltlrtird to his mothr's irthpla. q u i r i s o u t w h i h r h i
H d i d t h r i n 2 2 . ' ' station'' (H I. 6,497

of Grmany).In L. N4.
]r Jurassi
Birds:bluth Hds of
of aliforniaPrss.
f Birds.Nw Hirvn,C: Yal
207:Qu
007.rhoptr1'
h hindlimbfathrsin
32 417_41.
l|-prsrvdArhtl!rspimn
,-1486.
disputsand
of irds:Disovris,
issshft9\: 45547 \ . -A.F .
riginator of th sintifi study of
th thortialstudy of natur. His
rs of imIs, and th Gr-
;pializdstudis) provid a tho.
rr biologial invstigation and th
r Was nothing of si.rilarsop and
uy. In |837 t grat antomist
rl Sinspran.from his [Aristo-
linrva from th Had of Jov, in a
)wn l 992.9l ). Bforxmining
ls aut its rator ar in ordr.
hrn Agano;lst il1 84 ... His
.ing Amyntas of Madon, and his
Lisland of uoa. H was snt at
d in Plato's Aadmy for 20 yars,
had dvlopd his own distintiv
Lfor th study of natur. H joind a
t of Asia Minor' ut soon movd to
t Thophrastus' a yolrng rnan with
n th trvo of thrrr thy originatd
o u t a s y s t m t i i n v s t i g a t i o no f a n -
plants.
;totl to tutor his son Alxandr' but
w undr Alandr's ontrol.
)Wn as th Lyum' hadd th
lath in 323. With anti-Madonian
l rtirdto his mothr's irthpla.
ristotl 42
Th surviving opus of Aristotl drivs from mdival manusripts asd
on a first ntury ... dition. Th first printd ditions and translations
dat to th lt fiftrrthntury. Ths works ovr a rmakal rng f
sujts,ut for the purposs of this ssay I will fous on his animal studis,
whih onstittlte25D of th xisting opus.
ARlsoL,s PHlLosoPHY oF sIN
Aristotl was al to aomplish what h did aus h had givn a grat
dal of thought to th natur of sintifi inquiry. ow dos on pogrss
from th suprfiial and unorganizd stat of vryday prin towrd or-
ganlzedsintifi undrstanding? To answr this qustion' you nd onpt
of th goal to hivd'and Aistotl dvlopdsuh a onpt in his Prior
d Postrior rzltis. Th goal of irrquiry, h argud, was systm of on-
ptsand propositions organizd hirarhially, ultimatly rsting on knowl_
dgof th ssntial naturs of th ojts of study and rtain othr nssary
firstprinipls. Ths dfinitions and prinipls form the asis of ausal pla-
nationsof all th othr univsaltruths abrrtth sujtof study.
Th sond ook of t Posterior Aaltis disusss how to ahiv this
gal. Plato had fornrr.rlatda famous paradox of inquiry in his dialogu
o: either you know th oit f y<luinquiry, in whih as inquiry is
unnssay' o you don't knw th ojt of your inquiry' in whih as in-
quiryis impossil.Aristotl rminds us of this prdo in th first haptr of
thePostrior Anltics, ut his full ply is in ook II.
Thr h argus tht prptual xprin givs us a grasp of th targt of
inquiry that dos not ount as knowldg, ut dos provid suffiint infor-
mation to dirt furthr inquiry. His xampls in th Postrior ltis are
familiarphnomna lik thundr' tipss,and sasonal shdding of lavs,
ut w an s his nrthod at work in his s'stmatiinquiris into th
anatomyand dvlopmntof animls Portrd i Histor of Aimls (isto-
ri imliur)' on the Prts of Aimtls (De prtibs imliun), and n
th Grtiott lf imls (De gertion imlium)' From hr on w
will rfrto ths y th shorthand H, P, and G.
BloLoG|AL INoUlRY AND BIoLoG|AL KNoWLDG
For Aristotl, systmatis-th organization of irrformation aout animals-
srrvsth goal of ausal planation, nd xplanation y rfrnto goals
andfuntionsis fundamntal.Th following rmark, intoduing his system.
ati prsntationof information aout :rnimals' strsssth first point: ..In
thfirst instan w should grasp th attriuts and th diffrns prsnt in
.With
all animals. Aftr w hav dn this w should attmpt to disovr thir
auss.This is th rratural way to arry out th invstigation, ginning with
inquiry (histori) out ah thing; for it oms apparnt from suh in-
quirisaut whih things and from whih tliings thr ought to dmon-
s t r a t i o n '(' | ' 6 . 4 9 | a 7 _ 1 4 \ ,
424 ristotl
.W. .!
gin with a study of th similaritis and diffrnsthat w osrv in hav spokn, th
the animal kingdom; only ftr this informrion is systmtiallygathrdand how many thy ar, .
organizd ar W in a position to pursu ausl dmlnstrations.H gins y anothr. Nt w nt
outlining kinds of similarity (in form' in kind, y analogy) that on firrds first off in loodd r
among animals and thn prsnts te for ctgorisof differen h will us: loodlss that th gr
in parts, ativitis, ways of lif, and hartr. This disussion pr<lvidshis Now all that ar f
framwork: ooks I-IV disuss similaritis and diffrns among parts' windpip, positiond
V-VIII among tivitis and ways of lif, nd I among trits of hratr. too in all of th fu.
Among parts, Aristotl distinguishs twn uniform and nonuniform parts they [eah of th krn
(tissusand organs, as w would say) and twn trnal and intrnal parts, parts. Put gnrally,:
.,gratkinds.''
moving systmatially through thsatgorisin nin groups of xhal hav a lung, v
Th ..loodd kinds'' (orrsponding to our vrtrats)ar irds, fish, sophagus and wind1
taans' four-lggd liv-aring animals, and flur-lggd gg-laying ani- as of th lung nit
mals; th ..loodlss''ar th hrd-shlld(tstaousmlllusks),soft-shlld larly. . . .
(rustaans),soft-odid(phalopods),and instdanimals (insts).Aris- Not all th lood
..division and orrlation'' t-lstalish rltionships of
totl uss a pross of do any othr animal
..kinds''
similarity and diffrn at various lvls of gnrality,using th aov 56a12)
as an id in this prss.H nots that many animls (inludinghuman ings)
..gratkinds.'' Taxonlmi tidinss is not his primary goal;
do not fall into his In his rviw of wht l
ths atgoris help in idntifying lli of systmati tlrrlations, and ths t r ad diffe r entit i l
will aid in th disovry of th ausl rlationships among animal diffrntia. bn disussd.H is no
Primary among ausal rlationships ar thos that xplain a part's diffrntil first lists th kinds tht l
attriuts y rfrnto th funtionl dmands <lfan animl's way of lif. izs to a oxtnsiv or
.!7hy
th Prts of AimIs presentsAristotl's attmpt to provid suh ausal organs and lung.
undrstanding.It gins with a dfnsof th propr mthods for iologial at that lvl; this o
study. It argus for th priority of tlologialxplanatin (planatilnsthat baring animals' and fot
idntify what a fatur is for) ovr matrial nd ffiintausal planation aus of th fish, whil
(what th featur is md from and how it is prtldtld)and for onditional ntwork of organs dos
cessit, whry matrials and prosss ar plaind y ing shown to This disussion is r
nssaryfor a goal. (Noti this rinforsth pririty of tlology.)This ity. In disussing th snl
viw of nssityand plantion prsuppossthat animals irr unifid om- ar th sam [in snaks
posits of bodily parts with funtional apaitis,whih Aristotl rfrsto as a n d n o w n s st h v i s
..Form'' in iologial ontts rfrsto th
thir matrial nd formal natus. pip is vry llng nd th
asi apaitisof nutrition, rprodution' l<ltlmltion, and ognition, whih (whals and dolphins),t
A r i s t o t l r e f r st o a s t h n i m l ' ss o u l . m a l s ( 5 0 5 3 0 )a n d d i s
This first ook also dfnds a mthod of multifatoril division, whry of animls ar in fous
many univrsal diffrntia r dividd in pralll in ordr to haratriz gnralizationswithout
orrlatd variations within kinds, and it also dfnds a mthod for stab- It is qually imprt
lishing th kinds that ar prsupplsdy this mthod. P II-IV attmpt oth usI pltions on
to stalish auss and to provid ausl plantins tht paralll th first ar givn fo thm. ri
four ooks of H. Th third of ths lrg wlrks is G, whih provids ar nssryand says n
ausl planations for th sujtmattr ognizdirr H V_vII. ths parts. The lgtl
essec' sl ctrl to r
N o w o n t r s tt h d i s
E MEHoD A WoRK

Th following quotation from initiatsAristotl's disussion of th intr- Not all animals hav u
nal organs of loodd animals: ruhicb tlte nk is nt

rnd diitrnstht w osrv in
tion is systmatiallygathrd and
rsaldmtlrrstrations. H gins y
kind, v analog,v) that rr finds
'ltgorisof differ h will us:
tr. This disussion prlvids his
s nd diffrns mong parts'
lnd I am()ngtraits of hrtr.
n uniftlrm and nrrrrniforl parts
Wntrnal nd intrrral parts,
'isin nin groups of ..gratkinds.''
our vrtrts)ar irds, fish,
;, and four.lggdgg-laying ani-
mollusks), soft-shlld
(tsta()tls
nd instdnimals (insts).Aris_
ation'' tl stalisl-rrlatiorrships of
..kinds''
gnralit,v, using tlr aov
i nirnls(ilLrling humrr ings)
ri tidinssis not his pimry goal;
systmatiorrllltions,arrd ths
onshipsi1mong:-rrrimaldiffrnti:l.
)s that plain a part's diffrntial
nands of an arrimal'sway of lif.
tl'sattmptto provid strh ausal
th propr mthldsfor iologial
ial planatiln(planations that
l and ffiintausal plnatiorr
it is produd)nd for orrditional
's ar plind ing slr<lwnto
rsth priority lf tlology.)This
lossthat arrimalsar unifid om.
litis,whih Aristotl rfs to as
'
in biologialontxts rfrs to th
, loomltion,and lgnition, whih
of multiftorial division, whry
n paralll in ordr to haratriz
: lsldinds mtlrod fo st.
hismtlrld.P II-IV attmpt oth
plantionsthat prlll th first
trg u.rks is G, whih povids
.orgnizdin H V_VII.
:sAristotl'sdisussion of th intr-
ristotle 425
! hv spokn' thrr, of th trnal pts of rh loodd animals-
how many thy r, of what sort and in what ways thy diffr from on
anothr' Nt w nd to disuss th arrangmentof th intrnal parts,
first off in loodd animls-for it is y som ing loodd and othrs
bloodlss that th grat kinds diffr from on anothr. . . .
Now all that ar four-lggdnd liv-ringhav an sophagus and
windpip, positiond in th sam Way as it is in humn ings;likwis
too in all of th four-lggd animals that lay ggs, and in th irds. But
thy [ah of th kinds nrentiondl diffr in t onformation of ths
parts. Put grrrally,all and only thos that, y taking in air, inhal and
hal hav a lung, windpip and soph:rgus;and th plmnt of th
sophgusand windpip is similar' r.ltth orgns r not; whil in th
as of th lurrg nithr is th organ sirnilar nor is it positiond simi-
laly. ' . .
Not ll th loodd arrimais hav a lr-lng;for ampl, fish do not' nor
do any othr animals that hav gills. (rpts from H II. 505b24_
506a12)
In his rviw of what has n aomplishd, h nots that th umber, hr-
ter and diffrentitiozs of th trl parts of loodd animals hav alrady
bndisussd. is now turning to te itrl parts of th sm groups. H
first lists th kinds that hav oth windpip and sophagus, and thn gnral-
izsto a otnsiv orrltion-ll tat inal and hal air hav ths trro
ogansand a lung' !hy did h not start hrr? Baus thr is no ..gratkind''
at that lvl; this rrlatiln rangs ovr ids' humans, four-lggd liv.
aringanimals, and fou-lggdgg-layingirnimals.\Whynot ..loodd''?B-
aus of th fish, whilr a loodd, ut hav rron of ths organs-this
ntwork of organs dos not oinid with th loodd animals ither.
This disussion is rmarkal in its omination of auray and gnral_
ity. In disussingth snaks hr h nots that ..th st of th intrnal parts
ar th sam [in snaks] as in th lizards, pt that owing to thir lngth
and narrowness th visirar long and narrow . . . for ampl th wrnd_
pipis vry long and th sophagusvn longr'' (508a13_19).Th taans
(whalsand dolphins), though mntiond in his opning list of loodd ani.
mals (50.530)and disrrssdxtnsivlywhn th tivities nd ways of iif
of animls ar irr fous in ooks V_VI, lrr ar ssumd undr his wid
gnraliz,ations without ommnt.
It is qually important tO noti wht is tzot said in H. There are o
usl epltils on offr; univrsal orrlations aound, Uito re|sons
ar givn for thrn. Aristotl rfrains trom laiming that ths orrlations
arnssaryand says nohing irout why tlrsanimals hv or do not hav
ths parts. Th lgug of tur, mttr, fort, nssit, us ad
ess|1c' so trl to Aristotle's pbilosoph of ture, is rfttll uoidd.
Now ontrst th disLlssionof th sarorlations in P I. :
Nt all animals hav nk, ut only thos with th partsfor th sk of
hih the nk is turll prsnt-ths ar tlr windpip and th part
426 Aristotl

known as th sophagus.NoW th larynx is prsnt b ntur flr the ske

al to know t
o/athing; for it is through this prt that animls draw in and pl ir
.his would ra
whn thy inhal and xhal. is h tose without lung hav no
of animls-on
nk, .g. th kind onsisting of th fish' Th slphagusis th part through
sulr s pirinting
whih nourishmnt prods to th gut; so tht animals without nks
mor th study <
manifstly do not hav an sophagus. But lr ls ot nessrv to hav th
hold thir aus
sophagusfor th sh of nutrition; for it onots nothing. And furthr' it
at th xaminat
ls possibl for th gut to plad right nt to th p<lsitionof th mouth'
thr is somthi
whil for th lung this is irpossible. For ter ds first to sonrthing
ommon lik a onduit, Whih thn divids in two and through whih th
on Frttary 22
air is sparatd into ssags-in this way th lung m bst aonrplish
ths wlrds in a r
inhalation and xhalation. So, thn, th organ onntd with rathing
month rlir. In
from neessiryhas lngth; thrfor it is ssryfor thr to an soph-
I h a d s nI h a d a l
agus twn th mouth and th stomh. Tlr sophagus is flshy, with a
mot notion Wht
sinuous lastiity-sinulous so tht it ma dilat whn food is ingstd,yt
my two gods, thou
flsh so tht ttis soft and yilding arrd is not danrgdwhn it is sapd
to old Aristotl.'. r
y th food going down. (P I. ,664a14_35; mphassaddd)
you for your kind
I hav highlightd th languag of natttr,nssity,possiility (nd impos- m u h p l a s u r .I a
siility), and tlology in this passgin ordr to ontrast it with t disus- ompltd th Trin
sion of th sam organi orrltions in our H passag. Hr th go:rl is m gat satisfati
planation-parts ar prsnt and hav th haratr thy do primarily . riod of grat suffr
aus of th onditional nssityimposd y th ognism's funtional -
quirmnts. On ss hr th Aristotl that so imprssd th grat Frnh B|LIoGRA
naturalist Gorgs Cuvir: Aristotl is nt only systmatiallydisussingth
nglislrtr:rns[tion(
adaptiv funtions of ah of ths organs; h is also showing th anrazing wy UnivrsityPrs
in whih th intrnal parts of animals onstitut an orgni sStm. r.ommnd:
Aristotl.|949. Pri
Prss.
oNLUsloN 1992.Aristo
How slf-onsious was Aristotl of th rvolutionary natur of wlrat h was [. D. -N4.Blrn'
20|. th
doing? ln Rspirtio, ft ritiillly rviwing th rrrsof prvious
larndorrrs
writrs, h rfltson th rason for thir failurs: ..Th nrost basi rasn
For furthrdisussi
that prvious invstigatorshav not disussdspiration proprly is a lak
G o t t h l f , . , a n dJ . (
of prin with th intrnal parts nd failur to grasp what nturr Billog1'. Cambr
maks thm all for; for Wr thy inquiing what spiration is prsnt in an- Kullmann, \/.'and S
imals for, and wr thy doing so whil insptingth parts suh zrsgills and Stinr.
lung, thy would hav disovrd th aus quikl,, (7124-29|. l,nno,J. (;. 2001.
Spulation on th purposs trf rgns without doing disstion and Sic.mbr
own, R. |992. ll
anatomy' or doing disstionswithout thinking about funtion, is domd to
hiag:Univ
fail. Propr biologial inquiry rquirs that on ask the funtional qustion Pllgin,P. l 986.
whil doing th mpirial work. Suh work an vry unplasant.Nr th Uit of th ri'
los of his philosophial introdr"rtion
to th study oi lif, P I, Aristotl d- lifrniaPrss
monishes us to kp our y on th priz: Finally, thris fs
G o t t h l f ,A . 1 9 9 9 .I -
vn in th study of nimals disagralto prption,th natu that
3-30.
raftd thm likwis provids traordinary plasursto thos who ar

pfsntb ture for the sk
animals draw in and xpl air
thos without a lung hav no
sophagusis th part through
o tht animals without nks
it is ot |Issrto hav th
)notsnothing. And furthr, it
(t to th position of th mouth,
r ndsfist to b somthing
lin two and through whih th
th lung m best aomplish
rgan onntd with rathing
:ssr|or thr to b an soph-
Th sophgusis flshy, with a
lilat whn food is ingstd, yt
-rotdmagdwhn it is srapd
1-35;mphassaddd)
rssity,pssiility (and impos-
jr to ontrast it with th disus-
ur H passag.Hr th goal is
haratrthy do pimarily -
y th oganism'sfuntional r-
at so imprssdth grat Frnh
only systmatialldisussing th
i s a l s s h o w i n gt h a m a z i n g w y
itutan organl sstem.
rolutionarynatur of what h was
rviwing th rrors of pvious
..h mlst asi rason
failurs:
ssdrspirationproprly is a lak
c a failur tO grasp what natur
p r s n ti n n -
; w h t r s p i r a t i o ni s
sptingth parts suh as gills and
quikly'' (47\24_29).
ns without doing disstion and
rking about funtion' is doomd to
at on sk th funtional qustion
k an vry unplasant. Nar th
rh study of lif, P I, Aristotl ad-
ll to prption'th natur that
linary plasursto thos who ar
ristltle 427
al to know thir auss and ar y natur philosophrs. Surly it
would b unrasonal,vn asurd, for us to njoy studing liknsss
of animals-ln grounds that w re at th sam tim studying th art,
suh as painting or sulptur, that mad thm-whil not prizing vn
mor th study of things onstitutd y natur' t last whn w n b-
hold thir auss.For this rason w should not hildishly disgustd
at th amination of th lss valual nimals. For in ll ntural things
thr is somthing mrvlous. (P I. 5, 645a7 =1'7)
on Frury 22, 1'882,driddn aid nar dath, harls Darwin rd
thswords in nw translation snt to him as a gift y William ogl a
..From quotations
month arlir.In a thank-you not to Ogl, Darwin wrot:
I had snI had a high ntion of Aristotl's mrits, ut I had not th most r-
mot notion what a wondfr.rlman h was. l,innaus and uvir hav ben
my two gods, though in vry diffrnt ways, ut thy wr mr shool-boys
..Thank
to old Aristotl.'' Ogl rplid just thr days for Darwin did:
.th parts of animals'. It gav m
you for your kind and ulogisti lttr r
muh plasur.I am glad also to hav addd a third prson to your gods and
ompltdth Tinity.'' As a studnt of oth of thsgrat naturalists,ir givs
m grat satisfation to know that Aristotl affordd Darwin, during a p.
riod of grat suffring)som momnts of plasur.
|BLIoGRAPHY
nglishtranslations an fundin Harvard
of all of Aristotl'sbilgialtratiss
UnivrsityPrss'sL lassilLirar.In addition,for translations I
rommnd:
Aristotl.1949.Prbr d Postriorltis..$7. D. Ross,d. Oford: larndon
Prss.
AimIiumI and De Grtionimlium
1992.Aristotl'sD l,rti|:ls
I. D. . Balm.trar.rs.Oford:(]lrndonPrss.
2001'. th Prtslf imls. Books l-IV. J. G. Lnno, trirns.oford:
larndonPrss.
Forfurthrdisussionof Aristotl,siologilworks and thirsignifin' s:
A., and J. G. Lnrro,ds.1987.Philosophillssusi ristotl's
Gtthlf,
Bioklg,mridg:mridgUnivrsityPrss.
.W.,
Kullmnn, Biologi.Stuttgart:Franz
and S. Follingr,eds.1'997.ristltlish
Stinr'
Lnno,J.G.2001. ristrltl'sPhilosophlf l|illog:Stttdisith rigis of Lf
Si.amridg:ambridgUnivrsityPrss.
own,R. 1'992.h Hutri Ltrsi omprtiuett m. P. R. Sloan,d.
hiago:Universityof hiaglrss.
P. 1986. ristotl'slssifitilof imls: tsiologd th lcptl
Pllgrin,
I.Jitlf tb Aristltliorps.Anthonv Prus'trans.Brkly:Univrsityf
liforniaPss.
disussionof Drwin'sthoughtson Aristotlin:
Finally'thris a fasinating
Gotthlf, ^. |999. Darwin on Aristotl.Jourllf th Histor of tsiolog32:
3_0. -1.G.r,.
428 Th rkstts rtil Tril

for th nt dade'
Th Arkansas ration Trial
Amrian sin-r
Th mdrn rationist movmnt dats from th puliation in 1 961 of The
|LloGRAPH
Genesis Flood, oauthod y ilial sholar John . lil/hitom and hy.
draulis nginring profssor Hnry M. Morris (Numrs 2006). This Gilky, L B. 1985.re
..young rth'' aount Minnapolis:Wins
of ration attratd supPort and grw in strngth
Gish, D. |973. ulluti
during th 1970s, thirnks to th poslttzingwork y formr iohmistDu- N u m r s 'R . L . 2 0 0 6 ' 7
an T. Gish, author of uoltio: Tbe Fossi/s Sy No/ (|97|' Sin th Desig.pnddr
Sops trir[,tlr lgal situation hd harrgd.No longr was it possil to x. Pnnok,R., nd N{.RL
lud volutiorrfrom sh<lols.Th prolm now Was to gt th Bil in. Nt- Qltestili th r,
urally, th rrionist supportrs wantd to gt thi idas taught in th Pronrtlrus.
Rus, N4.,d. \988,Bul
iology lasss of stat.supportd shools, so thy dvisd a variant-
retio/t,'oluti
ration sin-that supposdly mad no rfrnto th Bil and was .ihitomb, ., nd H
J.
asd on purly sientifividn,ut just so happned to support a young- d Its Sitln
ag arth' si days Of ration, with humns last, and somtim thraftra Pulishingompa
massiv worldwid flood.
In 1981 th stat of Arkansas passd a law mndating th baland trat.
Artifiial Lif pro
mnt in shool iology l:rsssof volutionary idas :rdrirtionsin'Im-
mdiatly, th Amrian ivil Lirtis Union (ALU) supportd a suit volution is tlr dfin
laiming tlrat th l:rw was unonstitutiorralirusit rahdth sparation whrvr it riry ot
of hurh nl st.lt.Th trial was hld in Littl Rok in Dm 1981, with ird pollinting a fl
ALU xprt witt-tsss inluding philosophr ihl Rus, palontologist uts of volr.rtion.Li
Stphn Jay Gould, and population gntiist Franiso J. Ayala. Th as and is bginning to 1
.W.alla
Was mad that ration sinis far from ing traditional hristianity, ut unravld thr
is rathr an idiosyllrati prvrsion of nintnth-nturyAmrian Prots- dtails. A rlativly n
tantism; that it is rlot gnuin sinut rligion drssdas sin;that th instans of voltrtio
mpirial fts nd thortial hypothssspak against it; and that it is n- \/hilmillions of li
tirly inppropriat tat iology lasssshould prsnt indiffrntlydiffrnt thy all tra ak to
positions simply usvrious mmrs of soit-vhav strongly held dif. only a singl t of li
frnt pt-lsitions. iology and our p
Aftr a tw<l-wktrial, Judg William R. Ovrton ruld firmly that th [w truly ompartiv i
was indd unonstitutiona1and should disardd.Th stt didd not rquir knowldg of
to appal th dision. In oming to his onlusion, Ovrton argud that si- tion. It is livdth
n follows rtin fixd ruls of mthodology-it appals to unrokn law, ut' unfortunatly, th
it opns itslf to omparison aginst th findings from th mpirial world In its ssn,th p
(falsifiation),ani it is always tntativin its onlusions-and that ration gneti variation and
sinfails on all of ths sors.Aftr th trial, sonrphilosophrs ojtd atristis of tlr popu
strnuously that rro ition of dmaration an proply sparat sin arth is th produt r
from rligiln. Altlrouglr tlris oasiond xitd disussion in pofssional howvr, th poss
irls, gnrirlthinking in th sineotntrlltnity'not to mntion tlr world Thr hirs n s
lf journalisrn :rnd puli opinion, sidd lvith ovrton and this smdtO may th produt
th nd of mttrs. irth to nw univrs
rationists thmslvsargud that th right way to pursu thir nds was vary among unlvrs
not through laws that ar oviously opn to attk' ut at th grassroots Darwinian fitnss,lt
lvl, prssuring shool oards and individul tahrs.This st th pattrn physis, whih govr

om th pLrliationin 1961 of b
.!7hitom
nolar John . and hy-
'. orris (Nurrs 2006). This
td support and grw in strngth
ng work y formr iohmist Du-
'ossi/s S1, No! (797). Sin th
d. No longr was it possil to -
1 now was t gt th Bil in. Nat-
l to gt thir idas taught in th
ls. s thy dvisd variant-
no rfrnto th Bil and was
it so happndto support a young-
rns last, nd somtim thraft a
lw nrandating th land trat-
rary idasnd ration sin. Im-
Union (ALU) supportd a suit
l a u s i t r a h dt h s p r a t i o n
-ittlRok in Dmr 1981, with
phr Mihl Rus, pillontologist
:tiistFraniso J. Ayala. Th as
l ingtraditional hristianity, ut
ntnth-nturyAmrian Prots-
rligiondrssds sin;that th
s spak against it; and that it is n-
rould psntindiffrntly diffrnt
.s of soityhav strongly hld dif-
(. ovton uld firmly that th law
disardd.Th stat didd not
trnlusion,ovrton rgud that si-
olgy-it appals to unbrokn law,
findings frm th rnpirial wold
r its onlusions-and that ation
l trial' som philosophrs ojtd
ttion n proprly sprat sin
xitd disussion in profssional
mmunity,l1ot to mntion th worid
vith ovrton and this smd to b
' r i g h t w a v t o p u s u t h i r n d s w a s
n to attak' ut at th grassrots
idual tahrs.his st th pttrn
rtifiil Lif Progrts nd uolution 429
for th nt dad, until th lntllignt Dsign disussion gn nd th
Amrian sin-rligion quarrl gaind nw lif.
BIBLIOGRAPHY
Gilk,v,L B. 1985. reti'listltl Tril: ulltfu d Glld t Littl Rok.
innapolis: \/instlnPrss.
Gish' D. 197. uolutil: Tb Fossils Sar, No/ San Digo: ratiln-Lif.
Numbrs, R' L. 2006.
-h
rtklists: From Sitifi rtioism to ltlligt
Dsign. Bandd d. amridg, A: Harvard Univrsity Pss.
Pnnok, R., and.. Rus, ds. 2008. But Is lt Sie? The Philosoohil
Qustio i t'rtion/uolutio ltrr,,rs\,. Updatd l. BLrfflo. NY:
Promhus.
Rus, M., d. 1988. Bt ]s lt Si? Tb biloslpbicl
Quastin i tb
rtk/uoltio otrouersy. Buffalo' NY: Promthus.
Whitom' J. ., nd H. . orris . 1961. The Gesis Fklod: h Biblit Rclrd
d lts Sitifi Implitilns. Philadlphia: Prsbytrian and Rformd
Pulishigo-rpny. -.R.
Artifiial Lif programs and volution
volution is th dfining proprty and rativ poss of lit, on arth and
whrvrit may our. Th htah running down its pry' th humming-
bird pollinting a flowr, and th drama lf th human mind ar all prod-
uts of volution' Lif hs ompltly rshpd th srrrfa of our plnt
and is bginning to pro th rst of our solar systm. Sin Drwin and
Wallaunrvld th mystry of volution' many sintistshav studid its
dtails.A rlativly nW approah to th study of volution is to rat nw
instansof volution' an approah that hs n alld Artifiial Lif.
!hil millions of living spisurrntly xist on arth' it is livd that
theyall tra ak to a ommon anstor' illions of yars go; thus thr is
only a singl tr of lif on rth. From this prsptiv,our nri sienof
biologyand our xpinwith volution is asd on a sampl siz of on. A
trulyompartiv iology nd tuly road prsptivon volution would
rquirknowldg of othr instans of lif and its gnrativ pross' volu-
tion. It is livd tht lif xists on many plants throughout th untvrs
ut,unfortunatly, thy ar out of our rah'
In its ssn,th pross of volution involvs slf.rpliating ntitis with
gntivarition nd diffrntialsurvival, whrh lads to hangsin th har-
tristisof th population of ntitis ovr th ours of gnrations. [,if on
rthis th produt of volution inhiting th mdium of aron hmistry;
howvr,th pross of volution an oprat in othr mdia as wll.
Thr has n som spulation among physiists that th univrs itself
may b th produt of volution. Som hav suggestdthat lak hols giv
birth to nw univss, and that th fundamntal onstants of physis may
varyamon]univrss.Univrssthat produ mor lak hols hav a high
Darwinian fitnss, lading to th volr'rtionof th fundamntal onstants of
physis,whih govrn tlr haratristis of univss.
40 Artifiil Lif Progrms d uoltil

Spulation asid, w hv disovrd that volution an inhait th yl an rpt svra

mdium of digital omputation. If w apt that volution is th dfining parasits'' that atually
proprty and tiv pross of lif, thn instansof digitl volution may Somtims, whn on(
also onsidrd instans of lif, alit dramatially irlin lif. If w had suh that all th ratu
th opportunity to obsrv lif on othr plnts, it likly would aron- iality in whih individ
asd lif and thus would hav tht muh in ommon with lif on arth. Ar- d u , u t t u r sl i v i
tifiial lif in th digital mdium shars only th volutionary pross itslf in this kind of digital wor
ommon with lif on arth' and so is mor alin than lif on lthrplants. mat to produ young
Th digitl omputational mdium is not a physial/hmialmdiium;it is gnti matrial int th
a logial/informational mdium. Thus ths nw instans of volution ar tion ontinud vn wl
not suit to th sam physial laws as organi volution (.g.'th laws of raturs had invntd
thmodynmis),arrd thrfor thy xist in wht amounts to anothr uni- produd ontaining m
..physial laws'' of th logi of th omputr. Thy
vrs' govrnd y th nt. It involvs a kind o
nvr ..s'' th atual matrial from whih th omputr is onstrutd;thy nti matrial from th
s only th logi nd ruls of th U and th oprting systm.Ths ruls hav did.
ar th only ..natural laws'' that govrn thir havior. Thus thy liv in a Whil th ologial
radially alin univrs. Inoulating volution into th digital mdium givs driving for for volu
us a roadr prsptivln what vllutionis and what it dos. on form of ffiiny
On of th most sussfulapproahs for rating digital volution is to du an offspring. Thi:
..digitalor-
writ slf-rpliatingomputr progrems, whih hav n alld gnom that dsrist
ganisms'' or simply ..raturs,''and run thm on a OmPutr with a Dar- mak a opy. Howvr
winian opratingsystm.A Drwinian opratingsystmmanagsa popultion volution of mor om1
of rpliating digital organisms in suh Way that whn nw raturs ar Th volution of mo
orn, oldr ons di to fr spa in th mmry inhaitd y th programs. grail of Artifiial Lif. t
Th Darwinian oprating systm also introdus mutation y flipping its plx and autiful lif :
(twnzro and on) in th od of th raturs.Th random mutations gnrat omplity th
aus gnti varition in th population' with th rsult that slmindividu- unralizd goal of Artil
als ar al to rprodu ttr thn othrs, wl.rihlds to natural pross opn-ndd volution,l
of Darwinian volution in n artifiial digital mdium. as ourd On arth.
Although this produs a dramatially alin instn of volution, it is Lif appard on ar
found to hav som striking parallls to organi volution on arth. Prhaps form of singl.lldol
th most signifiantfinding is that th digital organisms volv adaptationsto point in tim, lif mad
th psnof othr digitl organisms in thir nvironmnt. A digital ol- singl-lldforms lk
ogy mrgs,whih omsth main driving for of volution. forms with ompl ani
A omputr an sdd with a singl slf-rpliatingdigital organism, apl of oordinatr
whih will quikly rprodu and fill th mmory of th omputr with urrd so auptly that
population of raturs.Th raturs ar thn a vry prominnt fatur of xplosion of divrsity.'
th nvironmnt and om an important sour of sltivfors. Pr- Som hav put forth
sits ar on of th first things that typially vllv, and thy st off an srv for digital organ
ologial-volutionarydynmi that lads to an ongoit-tgsrisof volution. ould ontriut som t
ar innovations. prsrv' and digital or1
.sfhatvr
kind of ratur is most ommon omsa targt for ploita- putr to omputr, f
tion y othr raturs. Or if parasites lrn()mm()n' thy driv othr that if a large and om
raturs to volv dfnss.A typial snrio would gin with th volu- natur, thn a digital '
tion of parasits' followd y th volution of immunity y tlrir hosts. This or thr.
 Artifiil l,ife Pr>grmsd uoltion 41

rat volution an inhait th yl an rpat svral tims ut may progrss to th volution of ..hypr-
t tlrt volutilnis th dfining pasits''that atually ttak parasits,stalingthir nrgy.
t n st l f . l i g i r e l v o l u t i < l nm a y Somtims,whn on kind of rtur ompltly d<lminatsth mmory'
amatillyalin lif. If w had suh that ll th raturs ar losly rltd, thy will volv kind of so-
rts,it likly wor"rld rborr- iality in whih irrdividul ratr'trsliving in isolation ar not bl <lrpro-
ommon with lif on arth. Ar- du, ut ratrsliving in groups ar bl to rprodu. Th raturs in
h volr'ltitlry pross itslf in this kind of digital world ar typially sxul in th sns tat thy do not
in than lrf on othr plants. mat to produ young ut rprodu individully, opying only thir own
ph,vsial/lrmial mdiunl; it is gnti matrial into thir offspring. Howvr, it was disovd that volu_
n w i n s . t t r . 'si v o l u r i o n a r tion ontinud vn whn mutirtions wr turnd off. This was eausth
n i v o l u t i o n( . g . 't h l a w s o f raturshad inventd a kind of primitiv suality in whih offspring wr
r ',htamounts t() anoth uni_ produd ontaining mitus of gnti matrial from mor than l par_
h logi of th omputr. Thy nt. It involvs a kd of s with th dad, in whih offspring inlud th g-
1omputis onstutel;thy nti mtrial from thir parnt, mid with som gns from raturs that
1 oprtingsystrn.Ths ruls havdid.
ir havior.Thus thy liv in a !hil th ologial ovolutionary dynami dsrid abov is th main
' l r i n t o t h t l i g i r a ln i d i t r m g i v s driving for for evlution' thr is also prptual sltion for ffiiny.
s and rvhat it dos. on form of ffiiny involvs rduing th amount f tim it taks to pro-
r r a t i n 8t,l i g i t l v o l u t i o n i s t > du an offspring. This is usually aomplishd y rduing th siz of th
..digital or-
rih hv nalld 8nomthat dsris th lrganism, thry rduing th tim that it takS to
lm On a Omputr with a Dar- mak a opy. owvr' som optimizations hav n ahivd through th
ing systrrrmanags a populatiorr volutionof mor ompl omputr ld.
a that w nr"Iatlrrs ar Th volution of mr omple od is a tantalizing ampl of th holy
nory inhaitd th progams. grail of Atifiial Lif. volution transformd simpl moluls into th om-
ldttsmutation y flipping bits pl nd bautiful lif foms tht w 6nd on arth today. It is th aility to
. h antlommutations
r i u r s gnratmplity that is th sour of volution's powr. It rmains an
itlr th rsult thertsom individLr_ unralizdgoal of Artifiil Lif to produ an tifiiirlsystm that hiits
wlrih lds to a natural pross opn-nddvolution' lading to vr mr omple artifiial lif forms, suh
ll rnditrm. as ourrdon ath.
l i n i n s t a n t l f v o l u t i o n , i t i s Lif appard on arth rughly 3.5 illion yars ago' ut rmaind in th
3anivolLrtionon arth. Prhaps form of singl-lldorganisms until aout 600 million yars ago. At tht
l o g n i s rs v o l v a d a p t t i o n st o point in tim, lif mad n arupt transformarion from simpl mirosopi,
:hir nvironmnt.A digital ol. singl-lldforms laking nrvous systms to l;rrg and omplx multilld
:ng flrof voltrtitlrr. forms with ompl anatomis, physiollgis, ologis, and nrvous systms
s l f - r p l i a t i ndgi g i a l o r g a n i s m , apal of oordinating sophistiatd havior. This transfomation o-
mmory lf th omputr with a urrdso auptly tht vlutirnaryiologists rfr to it s th ..amrian
thn a vry promillnt fatur of plosionof diversity.''
t sour of sltiv fors. Para- Somhav put forth a vision of a digital ntllr' a kind of iodivrsity r.
;irlly volv, arrd tlry st off an svfor digital organisms, prlrps distriud aross th Intrnt. Popl
to an ongoing sris of volution- ould ontriut som of thir mmory and CPU yls to th digital natur
prsrv'and digital orgnisms ould liv in t spa' migrating from om.
ron ots targt for xploita- putrto omputr' fding on unusd Inmory and CPU yls. Th ida is
Omommon' thy driv othr thatif a larg and omple rgiol of yerspa ould st asid for digital
rrio would gin with th volu- natur'thn a digital amrian plosion of divrsity and omplity might
-rof immunity y thir hosts. This ourthr.
42 Artificil Life Progrms d uoltil

Humans hav n managing th volution of othr spisfor tns f thou- Although th vast in
sands of yars through th domstiation of plants and animals. It forms th out of rah, mor mo
.W
asis of th agriultur that undrpins our ivilizations. manag volution approah has n to f
through rding' th ppliation of artifiial sltion to aptiv populatins. prforming omputatio
Similar approahs hav n dvlopd for working with volution in th plx omputations. Thr
digital domain. It forms th asis of th filds of gnti lgorithms and gn' algorithms and has
ti programming. Howvr, aus digital volution has not yt pssd to produ what appa
through its vrsion of th Camrian xplosion, thr xists th possiility to Studying organi v
..managing'' digital volution. .!nd
us a radially diffrnt approah to ours ovr vast sals
not limit ourslvs to using volution to produ superior versiRs of.qisting proah to th study of
softwar appliations. Rathr, w should llow volution to find th n ap- srv th pross of v
pliations for us. To s this pross mor larly' onsidr how w manag modst omplity. Als.
appliations through organi volution. allowing us to know tht
Som of th appliations providd y organi volution ar ri' orn'
what, arrots' f attl' dairy attl' pigs, hikns, dogs, ats' guppis, ot- lLIoGRAPHY
ton, mahogany' toao, mink' shp, silk moths, yast, and pniillin mold. Adami, . 1998.Itrrldu
If w had nvr nountrd any on of ths organisms' W would nvr hav G o l d b r gD ' . . 1 9 8 9 .
.W
thought of thm ithr. hav mad thm into appliations ausw r- Lrnig. Rading,N
ognizd th potntial in som organism that was spontanously gnratd Koza, J. R. 1992.Ctt
s of Nturl S
within an osystm of organisms volving frly y natural sltion.
l - n g t o nC' . G ' . d . l 9 q 5 .
any diffrnt kinds of things our within volution. Brding rlts to Lnski,R. ., . Ofri,R.
..ttr,''
volution within th spis, produing nw and differnt, possily of ompl fturs.]
forms of isting spis. Howvr, volution is also apal of gnrating Lvy, S. 1'992'Artifiil Li
spis. vn mor signifiantly, volutin is apal of ausing an plosiv Books.
inras in th omplxity of rpliators, through many ordrs of magnitud R a y , T . s . 1 9 9 1 .A n p p r
of omplity. Th amrin xplosion may hav gnatda mplity in. J. D. Farmr,and S. F
Worksh'ptl r!ifii
ras of ight ordrs of magnitud in a span of 40 million yars.Harnssing .!sly.
ths normously mor rativ proprtis of volution rquirs a ompltly 1994a.volutin
diffrnt approah. 239_26.
.W |994. An volu
know how to apply artifiial sltion to onvrt polr-quality wild
r. r' .L "d tr ir .r .r a t r| ri rL". r 4r '. t. i, f/ ,r L. li 6 1
orn into high-yild orn. Howvr' w do not know how to rd alga into
d.,rtifiil Life:
orn. Thr ar two ass to this inaility: (1) if all w know is alga, w Smolin, L. 1997. Th Lif
ould not nvision orn; and (2) vn if w know all aout orn' W do not
know how to guid th volution of alga alng th rout to orn. our p-
rin with managing volution onsists of guiding volution of spis
through vriations on isting thms. It dos not onsist of managing th
gnration of th thms thmslvs. Ayala, FranisoJ.
An altrnativ is to lt natural sltion do most of th work of dirting Franiso J. Ayala was l
volution and produing omplx softwar.This softwawill ..wild,'' liv. guishd mmr of th
ing fr in th digital iodivrsity rsrv. In ordr to rap th rwards and knowldgd for his stu
rat usful applitions, w will nd to domstiat som of th wild digi- sin.H has bn D
tal organisms, muh as our anstors gan domstiatingth anstors of vrsity of alifornia at
dogs and orn thousands of yars ago. U n i v r s i yo f l i f o r n i
This vision of digital natur is urrntly unattainal, and may always r- Ayala's arr has ot
main so. Still, mny sarhrsin th fild of Artifiil Lif ar still working xpansion of molular
toward an opn-nddvolution in th digital mdium. has n a ladr in otl

tlrrspisfor tns of thor-r-
rnts and animals. It forms th
.$
zations. manag volution
,tionto ptiv popLrltions.
working with volution in th
f gntialgorithms and gn-
voltrtionhas not yt passd
, thrxists th possibility to
.W
rg'' digital volution. nd
: supriorvrsionsof xisting
l volution to find th nw ap-
rrly, onsidr how w manag
ani volution ar ri, orn,
. k n sd. o g s , r s .g u p p i s , o t -
thsoyast,nd pniillin mold.
rganisms,w would nvr hav
o appliationsausw r-
was spontanouslygnratd
ly v naturl sltion.
t volutiorr'Bding rlats to
..ttr,,'
nd diffrnt,possily
l i s . r l s o p a l o f g n r a t i n g
:apalof using an xplosiv
ugh many ordrs of magnitud
ravgnratda omplity in-
f 4 0 m i l l i o n a r s .H a n s s i n g
evolutionrquirs ompltly
r to onvrt poor-quality wild
t krrow how to rd alga into
1) if all w know is alg, w
now all about orn, w do not
ng th rout to orn. Our p-
t guiding volution of spis
s not onsist of mnaging th
l most of th work of dirting
'his ..wild,'' liv.
softwarwill
ordr to rap th rwards and
mstiatsom of th wild digi-
domstitirrgth nstors of
rattainal,and may always r-
f Artiliial [,if r still woking
l mdium.
l 4
Although th vst inrassin omplxity nvisiond for digital nture ar
out of rtrh'mo[ modst inrassin on,rplxityhav n ahivd. on
approah has n to fd numrs to digitl organisms nd rward thm for
prfrming omputations on thm; gratr rwards ar givn for mor om-
pl omputations. This pproalr has ld to the volution of quit ompl
algorithms ar-rdhas n usd as a dtnonstrtionof th aility of volutirl
..irrduily''ompl stuturs.
t produ wht appar to b
Studying organi volution an frustrating aus it is a proess that
ours ovr vast sals of tim and sp. Artifiial Lif is an xiting ap-
poah to th stlldy volution aus it provids an opportunity to o-
srv th pross of evo'lution in ation, gnrating novlty, ologis, and
modstonrplity.Also,.it roadns our prsptivon lif and volution y
allowing us to know th volutionary pross in a nonorgani mdium.
BIBLIoGRAPY
Adami, . 1998,ltrodutioto rtifiil Llle. Nw Yok: Springr-Vrlag'
Goldrg,D. . 1989. GtiIgorithmsi Serh,optimiztio, nd hine
Lerittg.Radirrg,A: Addison-Wsly.
Kza' J. R' 1992. GetiPrr-lgrmmig: th Progrlnmigof omputrsb
s of Nturl Seltit'l. amridg'A: tr4ITPrss.
Lngton,. G., d. |995, rtifiilLif:An t,eruie. mridg,A: IT Press.
l-nski,R' I].,. Ofri, R. T. Pnnok'nd . Adami. 2003.Th volutionaryorigin
of omplfaturs.Ntur 423: 139_144.
Lvy,S. 1992.rti|t:ilLif: Th Qust for Ne rtio.Nw York: Pnthon
Books.
Ray,T. S. 1991'An approahto th synthsisof lif.In C. G. Langton'. Taylor,
J. D. Farmr,and S. Rasrussn,ds.'rtifiil Life II: Procdingsof the
Workshlpo ArtifiilLife' vr>|' l1, 371_408.Rdwoodity, A: Addison-
Wsl.
1994a.volution' onrplity'ntropy'nd rtifiialrality.PhsiD 75:
2.9_26.
1994.An volutionrry approhto synrhtiioiogy:Zen and th art of
rating\if,rtifiil Lii 1, no. |l2: 179-209. Rprintdin . G. Langtn'
d.,ArtifiilLii: u,i,|79-209. mridg,MA: IT Prss,1995.
Smolin,L' 1997.TheI'ifof t orls.N,York: Oford Univrsityo'..,-'.*.
Ayala,FranisoJ. (b. 1934)
FranisoJ. Ayala was on in Madrid' Spain, on Marh 12, 1'94.A distin-
guishdmmr of th intrnationl sintifi ommunity, h has bn a.
krrowldgdfor hls studis in volurionary iologv and th philosophy of
sin.H has bn Donald Bren Profssor of Biologial Sins at th Uni-
vrsity of alifornia at Irvin sin 1987 nd Univrsity Profssor of th
Univrsityof alifornia sin 2003'
Ayala,s arr has oinidd with and' to a larg xtnt' has ld th ris and
pansion of molr.rlrvolution nd th philosophy of iology. Indd, h
hs n :r ladr in oth filds for many yars. Aftr his arly invstigtions of
434 l

th pross of spiation and th rol of gnti variation in vlurion, pr- th Amazon a

formd using traditional gnti mthods whil h was a graduat studnt, h ratory prlm
saw that th dvloping thniqus of molular volution md possil a and othr nvir
mor powrful approah to ths issus.Latr in his arr, Ayala usd mol- that natural s
ular mthods to luidat th volutionary origin and population strutur of phism. H dv
Trptosom' Plsmodillm, and othr human parasits.Th impat of Ayala's and frquny-
..Biology
ontriutions to th philosophy of iology startd in arnst with his I 1 ' 9 7 1 , a
..Tlologial th Univrsity t
as an Autonomous Sin'' \mrin Scintist, 1968) and his
planations in volutionary Biology,, (Philosoph of Sie, 1970). palontologist J
As a youth, Ayala was intrstd in sin and didd to study physis at th 1 970s tha
th Univrsity of adrid. Latr' as rsult of his philosophial and rligious lams; Antarti
inlinations, h immrsd himslf for fiv yars in thlogy at th Univrsity starfish' sa ur
of Salamana. tala nxt didd to om an volutionist, oming to,th th amount of
Unitd Statsin 1961ie am an Amrian itizn tn 1977. sours' ut on
Undr th advi of his formr Spanish tahrs, gntiists Frnando om adaptd t
Ga16n and Antonio de Zulueta, Ayala studid volution from a gnti pr- promptd nw
stiv with Thodosius Dozhansky, th Russian-orn Amrian gn- fossil rord.
tiist, at Columbia Univrsity in Nw York twn 7961 and 1964' During During th 1
his thr yars as a graduat studnt at olumbia h arrid out invstiga- ifornia t Irvin
tions aout th pross of spiation in Australian Drosophil and disov. as' whih is
rd rypti spis and intrmdiat stags of spiation. introdud nw H showd th
mthods for masuring population fitnss and dmonstratd that th at of thought at th t
volution orrlatd with th lvl of gnti variation. Ths studis wr l o n a l t h o r yo {
publishd in 1965-1966 in a sris of paps in Sic' Genetics' uoltio, tur of thsp
mricn Nturlist, ad Pifi Ists. man sourgsr(
In 1,965,as a postdotoral fllow' Ayala introdud ltrophortith. -N4orrntl
niqus for dtting prtin polymorphisms in Dobzhansky's loratoy at prum, th ag
Rokfllr Univrsity. H am an assistantprofssorthr in 1967.With that th world t
ths mor powrful thniqus, h now pproahd th arlir prolms of rnt' starting
masuring gnti variation (polymorphism) and how this impatd th rat that th spi
of voltion. H also initiatd a omprhnsiv invstigation of th gnti volvd in rsp
hangs onomitant to th spiation pross and th volution of rpro- In addition t
dutiv isolation. Invstigations omind laratory primnts with th of iologial pr
invstigation of natral ppulations of Drosophil illistonl and rlatd tintiv haat
spis of the Amrian tropis. pulishd th first omparative data ana- tions to th phiJ
lyzing gnti diffrntiation as a funtion of taonomi lvl in Drosophil of natural sl
spp.' as wll as in various othr animal groups. of living forms.
In ollaoration with Dozhansky, Ayala onfirmd thortial prdi- tiularly, volu
tions aout th distriution and adaptiv signifian of gnti variaility in larization of r
natural populations. on main ontriution to volutionary iology was his forfully argu
dmonstration that thr is muh mor gnti polymorphism in natural nd not in
populations than prviously thought. m lading partiipant in th Ayala has
ontrovrsy rgarding whthr molular polymorphisms Wr adaptivly maks sns
nutral or drivn y natural sltion.H argud forfully for th lading rl volution mak
of natural sltion asd on th distriution of tnsiv, ut gographially Franiso J. ,
fairly uniform' gnti polymrphisms in natural populations of Drosophil in wll as th autl

variationin volution, pr-
was a gradutstudnt,h
v<llutionmad possil a
is arr,Ayala usd mol-
and population strutur of
rsits.h impat of Ayala's
..Biology
in rnstwith his
..Tlologial
1968)and his
of Sice,1970).
d i d dt o s r u d p h y s i s a t
p h i l o s o p h i al n d r l i g i o u s
l t h o l o g y t t h U n i v s i t y
volutionist,orning to th
tizen in 1971.
hrs, gntiistsFrnand
lolution from a gnti pr-
ssian-ornAmrian gn-
nl96i nd l964. Duing
t i . rh a r r i d o u r i n v s t i g -
ian Drosophil ad disov-
iation.H introdud nw
monstratdthat th rate of
liatiorr'Ths studis wr
ii,G tis, L'olutio, tur of thsparilsits is prvailinglv lonal, vn though ths prvsiv hu-
ldud ltrophortith-
Dozhansky's laboratory at
,rofssorthr in 1967 'Wlt|t
hd th arlir prolms of
1 how this impatd th rat
' i n v s r i g t i o no f t h g n t i
nd th volution of rpro-
ator.vxprimnts with th
'phil illistoi and rlatd
firstomparativ dta ana-
onomi lvl in Drosophil
onfimd thortial prdi.
:anof gntivariaility in
volutionaryiology was his
i polymorphism in natural
La lading partiipant in th
rmorphisms wr adaptivly
fofully for th lading rol
tsiv'ut gographially
populations of Drosophil in
Al 45
th An-razonasin anl tlrroughout th Caibban islands. dsignd labo-
ratry xprimrrts t masuring tlr fftth amount of food, livirrg sp,
and lthernvironnrntaIfators hd on population fitnss.Rsults orrfirmd
that natural sltion Was th driving fato imparing gnti polymor-
phisnr. H dvlopd Iaoratory mthods for masuriIlg dnsity.dpndnt
and fquny-dpndntntural sltion.
In |97|, yIa and Dozlrnsky movl to th Dptmnt of Gntis at
tlr Univrsity of lifrnia at Davis. At UC Davis, Ayala ollaoratd with
palontologistJams \W. Valntin irr sris of invstigationspublislrd in
th 1970s that involvd divrs organisms, inluding tropial rid,n
lams; Antarti, tmprtl, and tropial oani shrimp; and dp-oan
starfish,sa urhins, and rahiopods. Tlr invstigationsdmonstratdthat
th amlunt l gnti vriarlrty 1pnddnot On th abundan of r-
sours'ut on nvironmntal staility, whih allows gnti variants to b-
om adaptd to diffrnt nvirnmtal fatursand rsours.Th rsults
promptd nw hypothssto plain th major tintions osrvd in th
fossil rord.
Duing th 1980s' rvorkirrgt U Dvis and latr at th Univrsity of al-
iforni t lrvin, Ay:rl studid TrpLl'Iosomruzi, th agnt of hgas dis-
as,whih is a svr disas ndnri in many South Amrian ountris.
H showd that it rprodus lonally, rathr than sually, as gnrally
thougl-rt at th tim. Ayale and ollaor;rtorihl Tiayrn formultd th
lonal tlrory of parasiti protozoa. hy proposd that th population stru-
m11sourgsrtain th apity for sxual rprodution.
or rntly, Ayala has ronstrutd th volution of Plsmodium fIi-
pru|' th gnt of malignant malaria' and rlatd spis.H has shown
that th world pansion of P. flipltzzthroughout th tropial rgions is
rl1t'starting from :r singl Afian propagul rrrund 5,000 yrs ago, and
tlrat th spisis grrtiallylargly uniform, pt for mutations rntly
volvdin rsponsto th human imt-nunsystm and to mdiins.
In addition to his ssayson iology as an autonomous sin,th onpt
of iologial progrss' and th id of tlology, Ayala has xplord th dis-
tintivharatistisof th sintifirrrthod.H has m:rdothr ontiu-
tions to th philosophy of sin,inluding xplorarions of how th pross
of r-raturalsltionprovids a ausl planation of tlr volution and dsign
of living forms. \/ithtnsivknowldg of th history of iology and, par-
tiuIirrly,volutionry thoris, h has ontributd tnsivly to th Popu-
larization of volutitlnry thory as th ntral or of biology. H has
forfullyargud thirt tlr tlror'l'of l,olution, as wll as sin in gnal,
ndnot in orrtrzrditionwith rligious lifs.
Ayala hs tndd Dozhansky's rnownd phras ..nothing in iology
maks sns pt in the light of volution'' with th phrase ..nothing in
volutionmaks snsxpt in th ligt of gntis.''
Frniso J. Ayala is th author of ro thn 900 spializd artils as
wll s th author or ditor of mlr thn 20 ooks. is a mmr f th
46 Al

National Aadmy of Sins, th Amrian Aadmy of Arts and Sins,

th Amrian Philosophial Soity, th alifornia Aadmy of Sins,th
Russian Aadmy of Siens,th ltalian Aadmia Nazional di Lini,
and aadmis of othr ountris' inluding xio, Sria, and Spain. H
has srvd as prsidnt of th Amrian Assoiation for th Advanmnt of
Sin, th Sigma i Th Sintifi Rsarh Soity, and th Soity for th
Study of volution. H has rivd numrous honors and awards, nd h
has n dsignatd Dotor Honoris us univrsitis in Argntina,
zh Rpuli, Gr, Italy, io, Spain, and Russia. In 2002 h r-
ivd th National dal of Sin, th highst sintifi award in th Batrial volution
Unitd Stats, from Prsidnt Gorg W. Bush.
Th Miro is so vry
|BLIoGRAPY Y o u a n n o m a k h i n l
-
B u r m n s n g u | np
Ayala, F. l. |965a. Drlsophildomii,a nw silingspisof teSerrt
grolp. Pc lsts7: 62-622. To shim throughtht
1965. volution f fitnssin primntalpopulationsof Drosophil
serrt'Sie150:903_905.
1965.Silingspisof t Drosophil serrtgroup.uolutio 1.9z Atually, Hilair Bllo, w
538-545. out of dat. iros w
t966a. Dynamisof populations.I. Fatorsontroilingpopulationgrowth looking at srapings from
and populationsiz in Drosophil srrt.Tb Amrin Nturlist 100: labld btri in 1828 L
33-344. rg' from th Grk wor
|966. volution of fitnss.I. Improvmntsin th produtivityand sizf
ogists disovrd th r
irradiatdpopulationsof Drlsophilserrtand Drosopbil birhii. Geetis
5 3 :8 8 3 - 8 9 5 . undrstand that divrsity
1968.Biologyas an autonomoussin.AmriSitist56:27-221. v o l u t i o n o f a t r i ai s o
1970.Tlologialxplanations in volutionaryiolog.Philosopbof xiting aras of study.
Sie37: 1-|5. B t i a r r l t i V l y:
206.Drtl,i d ltelligetDesig,innaplis,N: Fortrssrss.
maning that thy do not
207.Dri'sGift to Sieed Rligio..Washington' D: Josph
nry Prss. th gnti matrial is hou
.!. batrial ll. First, thr
H.y, J., . Fith, and F. J. Ayala, ds.205.Sstemtisd th origin of
.Washington,
Spis:n rst r's 100thAiursary. D: National ll hat primrily ari
AadmisPrss. omponnts. Sond, thr
Russll'R.J., !. R. Stogr'and F. J. Ayala'ds.1998'uolutiornd olulr primarily involvd with rr
Biolog: SietPrsptius o Diui titl. Vtianity Stat/Brkly,
not hav mor sophistiat
A: Vatin )srvatorynd th ntrfor Thologynd th Natural
Sins. -A.B. for powr gnratin) or
with ells that do hav nu
organlls usd for uild
Thy rprodu y simpl
nd ah daughtr1lis l
Lik all organisms, a
tion: lls with advantag
thm. Advantagous fat
First, th gns of at
vrsity an aumulat r:
of sual rprodution tl
gnti matrial n thir
t h h v a r i s ni n o n
Aadmy of Arts and Sins'
ornia Aadmy of Sins' th
:admia Nazional di Lini,
io, Sria, and Spain. H
ition for th Advanmnt of
Soity, and th Soity for th
,us honors and awards, and h
l y univrsitis in Argntina'
in, and Russia. In Z02 h r-
highst sintifi award in th
h.
l siling spisof the Srrt
al populationsof DroslphiI
ierrt groaP, uolutk |9:
,rs ontrolling ppulation growth
b mric Nturlist 1:.
:ntsin th produtivity nd siz of
and Drosophil birhii. Getis
meri Sitist5 6: 207 -22|.
.ionary iolog. Philosl1lh of
innapolis,MN: Fortrss Prss.
.lo. Washington, DC: Jsph
Sstmtisd th rigin of
.Washington,
,. D: National
' 1998. uoluti-lrd olulr primarily involvd with rgulating th ll's mtaoli prosss. Batria d
:tio. atican ity Stat/Brkly,
.Thology and th Natural
Batrial volution
Th Miro!-is so vry small
You anfot mak him out at all
But ma{lysanguinpoplhop
To shim thrughth mirosop.
Atually, Hilair Bllo, who wrot thsamusing lins, Was svralnturis
out of dat. iros wr first sn in 1674 Antoni van Luwnhok,
loking at srapings from his tth through simpl mirosop. Thy wr
laldbtri in 1828 y th Grman iologist hristian Gottfrid hrn-
..small stik.'' In susquntyars' biol-
brg,from th Grk Word maning
ogists disovrd th normous divrsity of atria. Th struggl to
undrstandthat divrsity, its volutionary history, and th ontinud rapid
volution of atria is on of volutionary iology's most hallnging and
iting aras of study.
Batria ar rlativly simpl, on-lld organisms' lald prokaryots'
maning that thy do not hav a spial ompartmnt' or nulus, in whih
th gntimatrial is housd. Thr ar two typs of gnti information in a
atrialll. First, th is a small, usually irular hromosom within th
ll that primarily arris th gns rsponsil fo uilding th ll and its
omponnts.Sond, thr ar a numr of gns floating in th ll that ar
not hav mor sophistiatdll parts (organlls)suh as mitohondria (usd
-4,8, for powr gnration)or hloroplasts (usd' .g.' y ukaryots-organisms
with lls that do hav nulei-for phtosynthsis). Thy do hv riosoms,
organllsusd for uilding protins frm th instrutions givn y RNA.
hy rprodu y simpl ll division; on ll divids into two daughtr lls
and ah daughtr ll is a lonal rprodution of its rnothr ll.
Lik all organisms' atria ar opn to volution through natural sl-
tio: lls with advantagous faturs prolifrat fastr than thos without
thm.Advantagous faturs aris in atrial lls through th avnus.
First,th gns of atria hang spontanously,or mutat, and lonal di-
vrsityan aumulat rapidly. Sond' som atria ar apal of a kind
of sxual rprodution through th hang, twn lls' of som of th
gntimatrial on thir hromsoms. This gn hang plas mutants
thathav arisn in on l1lin into othr ll lins. Third, atrial gns that
47
48 Btril ullut

af not on th main hromosom an oftn transfrrd dirtly from on onntrations of o1

ll to anothr. This pross' alld hlrizontalgn transfr,an failitatth of th most ommon
rapid sprad of a gn throughout a atrial ollny and hang th har- of th arth's istn
tristis of tlr olony quikly. Bus prokarylts a sirnplr than eukary. availal. As th ar
ots, th natural assumption is that th formr volvd for th lattr, and ris and sltion pr
that th lattr volvd from anst<lrsthat rsmldth formr. This assump- gan. Nt, oygn
tion sms tr born out y fossil rords, y th natur of ll funtion, and rgy. Th first vidn
y omparativ studis, prtiularly thos at th molulr lvl. Th arth is ould swith from o:
around 4.5 illion yars old and during its arly yars th surfa wag fa t othr (using oygn)
hot t sustin lif. Th first plausil tras of fossil lif dat ak to around lvls ontinud to ri
3.75 illion yars ago. At around .5 illion yars thr is solid vidnof 2". out2.25 i]
trial g!51gngg-yanoatria, othrwis known as lu-grn alga. os) appard.
Thy lustrd togthr in shts known as stromatolits and livd in th s ukaryots appa
long th Oasts.Shwing that hang is not a nssaryondition of lif on now livd' thank
arth, lu-grnalga of th sanr rognizal form still flourish today. Margulis, that uk:
Thanks partiularly to th 'uvork of th Amrian miroiologist arl among various prok.
!7os, it is thought that th last univrsal ommon anstor-th parnt of asord othr mir
ll living things past and prsnt-was aout 3..5 illion yars ago. !os, DNA on th nular
howvr, strsssthat it is probably Wrong to think in trms of just singl arhaa, it appars t
organism giving irth to all othrs.In fat, th tr of lif is rally a pitur of lls-ar dsndd
how lif volvd aftr rprodution nd inhritn through ll division today's wll-known
mrgd and am prdominant ovr rampant horizontal gn hang ing humans.) Th
and transfr. Bfor thn, it was likly that th fous of .thvolutionry pro. photosynthsis,also
ss was on th prolifration of individual gns nd small lustrs of gns. truly himras, whi
Th disrt linags rtrad thrlugh phylognti mthods wr not likly many diffrnt typs
in istnamong th arlistlls. It is ttr,and mor aurat' to think Th symioti rl
in trms of a group or population of anstors,all of whom would hav n omplx rlationshi
.Wos
important. nd his group rvld a mor aurat pitur of th tr Batri today ar ut
of lif than hithrto, and on of th most astounding findings is that th Many plants dpnd
prokaryots onsist of two vry diffrnt groups: th atria (or tru bat- thy an us it thm
ri or uatria)and arhaa (r, as thy hav n alld, Arhaatria). frtilizrs, and it is v
Thr ar signifiant diffrnstwn atri and rhaa. Most im- to modify plants gn
portant is that arhaa ar mor lik th ukaryotsin th way thy rad and tria, thy an thm
us th information of th DNA moluls.This apprs to th first major ria in thir stomah
split that w an rtra twn th atria and th omlon anstor of food. Batria in th
th arhaa and th ukaryots (s Figur 1 in th main ssay ..olular tin disassor stop
volution'' by Franiso J. Ayala in this volum). any arhaa ar apal us atria for thi
of or nd to liv in xtrm onditions (for mpl, hot or salty).This had usd in laning up t
ld to spulation that thy may rsmlsom of th arlistlif forms, s- Unfortunatly' it :
pially if' as many now suppos,th arlistlif was found in vry hot, dp- animals (spiallyh
sa vnts. Somwhr along th lin' thrfor,tria gan to los ths to history, tooth d
ailitis, or thy wr univrsally lost and thn rgaind. mouth: Strptloc
Th volution of atria saw nrajr hangs in th ways in whih nrgy plasant atrial di
an otaind and usd. In partiular' as an dtrmind from molu. cobterium i
lr studis as wll as rmains in th gologial rord, th first atria wr holra, ausd by
oligatorily anaroi, meaning that thy ould not surviv in signifiant terium tuberulosis

] transfrrddirtly from on
gntransfr,an failitat th
olony and hang th hara-
yotsar simplr than ukary-
.volvd flrth lattr, and
nldth formr. his assump-
th natur of ll funtion, and
t molular lvl. Th arth is
y yars th surfa was far too
tssii lif dat ak to around
Farsthr is solid vidn of
known as lu-grn alga.
omatolits and livd in th sa
L nssyondition of lif on
form still flourish today.
\mrian miroiologist arl
nmon anstor-th parnt of
.Wos'
3.5 illion yars ago.
think in trms of just a singl
trof lif is rally a pitur of
ritan through ll division
)ant horizontal gn exhang
l fous f th volutionary pro-
ls and small lustrs of gns.
nti mthods wr not likly
:r, and mor aurat, to think
, all of whom would hav n
or aurat pitur of th tr
stounding findings is that th
|ps:th atria (or tru at-
nalld, Arhaatria).
atria and arhaa. N4ost im-
:yotsin th way thy rad and
l i s a p p r st o r h f i r s t m j o
l and th ommon anstor of
..N{olular
in th main ssay
n). ny rhaa ar apabl
xampl, hot or salty). This had
n of th arlist lif forms, s-
if was found in vry hot' dp-
., atria gan to los ths
n rgaind.
s in th ways in whih nrgy
Lnb dtrmind from molu-
l rord, th first atri wr
uld not surviv in signifiant
Bteriluolutio 49
onntrtions of oygn. Thy found thir nergy y nonoygn mans; on
of th most ommon mans was y frmntation. Of ours, for th first half
of th arth's xistn' thr wr no signifiant onntrations of oygn
availal.As th arth sustaind hmial hangs, oygn lvls gan to
ris and sltion prSsurtoward oxygn-tolrant (arotolrant)organisms
gan. Nt, oygn-tolrant organisms gan to us oxygn to prod n-
rgy. Th first vid.oJ th faultativ aroi organisms-organisms that
ould swith frm on form of nrgy prodution (without oxygn) to an.
othr (usinglxygn)-an trad to aout 2.5 illion yars ago. Oygn
lvls ontiriud to ris until thy wr aolt 1o of today's prsnt lvl of
20%. Aout 2.25 illion yars ago' xlusivly oygn-usingorganisms (ar-
os)appard.
ukaryots appard shortly aftr th mrgn of oligat aros. It is
now livd, thanks to th rilliant hypothsis of Amrian iologist Lynn
Margulis, that ukaryots Wr formd through symioti intrations
among various prokaryots. Som miroorganisms, prsumaly arhaalik,
asrd othr miroorganisms, som of whih wr atria. \Whil th
DNA on th nular hromosoms of ukaryots indiat thir affinity with
arhaa' it appars that th mitohondria-th powr plants of ukaryoti
lls-ar dsndd from th sugroup protoatria. (sherihi oli is
today'swll-known protoatria; it livs in th guts of all mammals, inlud-
ing humans.) Th hloroplasts of plants, thos llular struturs that drive
photosynthsis,also appar to hav a symioti origin. Thus ukaryots ar
truly himras, whih ar th orgnisms arrying gnti instrutions from
many diffrnt typs of anstors.
Th symbioti rlationships that formd th ukaryots hav hos in th
omplx rlationships that modrn ukayots hav with modrn atria.
Batriatoday ar ubiquitous and othr organisms annot liv without thm.
any plants dpnd on atria to xtrat th nitrogn from th air so that
thy an us it thmslvs. Aiding this pross is th aim of many artifiial
frtilizrs,and it is wll known that tday griulturalists ar working hard
to modify plants gntially so that, thanks to gns transfrrd in from a-
tria, thy an thmslvs rat usal nitrogn. ammls us many at-
ria in thir stomahs and intstins, usually alld gut flora, to digst thir
food. Batria in ths plas also hav othr funtions, lik prvnting r-
tain disassor stopping various potntial allrgis.or gnrally,humans
us atria for thir own nds, as in frmntation. Tody, atria ar vn
usdin laning up oil spills.
Unfortunatly, it is oftn diffiult to liv with atria. Many disass of
animls(spiallyhumans) and of plants ar du to atria.Rvrting ak
to history,tooth day is du primarily to four kinds of atria found in th
mouth:Streptoous snguis, S' sobrinus, S' mitis, and S. mt.s.Othr un.
plasantatrial disasesin humans ar lprosy (Hansn's disas),ausd
cobterium lepre; whooping ough, ausd Bordtll prtussis;
holra,ausd Vibrio holor; and turulosis' ausd cob-
trium tubrlosis. Rntly, ppti ulrs wr found to ausd y
440 Br

Heliobctr plori. rtain intstinal disass, suh as Crohn's disas, a had n assumd to i
thought to th rsult of th ody's failur t rogniz its own atrial gut with fluid mittd .
symionts. his dirtor's pt dog
Batria ar known to rapid volvrs. Prft ampls of natural sl- ry with whih his n;
tion in ation, thy gntially put up arrirs against human prvntativ mryologial studi
masurs quikly. Antibiotis, spially pniillin, w th grat mdial t i k lgsgs h ih tt
disovris of th twntith ntury aus thy ntirly dstroyd.targtd 1 8 2 8 n d 1 8 3 7 .I n 1 t
batria without affting th human host. Unfortunatly' thanks to muta- drtook various kind
tion, high rats of rprodution, and horizontal gn transfr, atria d- Russian mpir. In t
vlop natural rsistans to antiiotis. For instan, almost as soon as though also a didd
.War
pniillin was introdud during !7orld II' it was found t'at Stphlo- Lik Gorgs uvi
coctts turswas dvloping gn-dirtd iohmial mthods of dfns. sparatd into fou
Th sam is tru of othr atria. Notoriously' vnral diss usd to urhins), th mollus
urd with just on shot of pniillin. Now it taks a oktail of drugs to and ras), nd th vr
omat it. It dos not hlp that humans do many things that nourgrsis- pitulation thory-th
tan dvlopmnt' suh as whn prostitutskp thmslvson ontinuous' through morphologi:
rlativly low doss of antiiotis. Batria sm to find this a hallng, find. ganisms lowr in th h
ing ways to surviv and thriv. imal mplifid fror
Th apaility of rapid volution in atria also prsnts a hallng for Urflrm of that parti
.W ..islrady at tJ
undrstanding thir arlist volutionary history. ar attmpting to r- srtd,
tra vnts that bgan at th dawn of lif, hanging rapidly and through man ftus, h hld, w.
mhanisms lik th horizontal transfr of gns with signaturs not asy to form of a gnralizd
tra. ]t4odrn advans in molular iology ar hanging our undrstand. and finally a partiul
ing, and th study of atrial volution is among th most hallnging and fronr th gnral to t}
iting topis in modrn volutionary iology. fo, nvr assumdtl
Dspit Bar's rj
|LIoGRAPHY
within a givn arh
Daon,l. Th icrobiI World. http://hlios.to.d.a.uk/t/miros/. arlir stagsof mr
Howland, J. 2000. The SurprisigArh.Nw York: oford UnivrsityPrss. attriutd to Louis A1
Univrsityof alifornia usum of alontolog'Itrodtilto tbe rh. tl mryos that h
http://www.ump.rkly.du/arhaa/arha.html/. -J.' d .R.
birds' or vry young
ths animls is quit
yos. But vn if th
Bar, Karl rnst von (1792_1876)
indiat anything; si
Karl rnst von Bar was prhaps th most minnt morphologist and mry- of irds, as wll s th
ologist of his day. H was orn in stonia to a Grman nole family. H at- mntal form'' (l828_
tndd th nw univrsity in Dorpat and ontinud mdial studis in Brlin Though h am
and Vinna. or intrstd in th thortial aspts of th sin, h pur- many Grmn morpl
sud postdotoral study at.!(iirzurg (1815), whr th rsidual influnof onrrtd him with t
Fridrih Shlling (-]"775_1854)ould still flt, transmittd thrugh th ratd y Shlling.Hr
gnial hands of Ignaz Dllingr (1770_184I). In 1817 h movd to Knigs- ampl, that th arh
rg to om assistant to Karl F-ridrih Burdah (1776-1847). Bar's guidd th ratur'
sor xprimntal approah to anatomy ld him to shd muh of his arlir ..t
mal,'' h dlard,
nthusiasm for a mor romnti and asthti intrprtation of natural ph. govrns its ntir dv
nomna. In 1826 h gan studying th fomation of th vrtrat mryo Bar ojtd to t
and initially onrnd himslf with th origin f th gg in mammals' whih rootd in Grman i

is'suh as rohn's disas,ar
o rognizits own trial gut
rftaInplsof natual sl-
r s a g a i n s th u m n p r v n t a t i v
niillin, w th grat mdial
thy ntirely dstroyd targtd
Unfortun:ltly, thnks to lnuta-
)ntal gn transfr, atria d-
rr instan, alrnost as soon as
: lI, it was found that Stplo-
iohmilmthods of dfns.
.rsly,vnraldisasr'rsdto b
l it taks a oktil of drugs to
r n yt h i n g st h t n o u r a g r s i s -
k pr h m s l v os n o n t i n u o u s '
m to find this a hallng' find-
:ria also prsntsa hallng for
s t o r y .W a a t t m p t i n gt o r -
:, harrrging rapidly and thr<lugh
;nswith signaturs t-lotasy to
3y ar hanging our undrstand-
among th most hallnging and
'g.
.d.a.uk/to/miros/.
,York: oxfod UnivrsityPrss.
, ltrodutioto th rh'
raa.html/. -.I .T. nd .R.
minnt mopholgist and mbry-
'o a Grman nl famil. H at-
ntinud rdial studis in Brlin
al asptsof th sin,h pur-
). whr th rsidual infln of
flt, transmittd through th
1 ) .I n 1 8 1 7 h m o v d t o K n i g s -
:h Burdah (1776-1847). Bar's
d him to shd rrruh of his arlir
:ti intrprtationof natural ph.
.mationof th vrtbrat mryo
;in of th gg in mmmals' whih
Ber 441
had rrassumd to idntifid with th Graafian follils of th ovirris or
with fluid mittd ,vtlr follils. Irrth sprir-rg of th nt yar h sarifid
his dirtor'spt dog nd loatd minut yllow ggs in its follils' a disov-
ry with ,|rilrhis.rrzrm would lrrrfotlr assoiatd.H ontirrud his
mryolial studis, pulishing th two parts f his mastrwork, Die n-
tiklgsgsllihtder hir (h dvlopmnralhistory of animals), in
1828 and l837. In 18.34h ptd a hair in St. Ptrsurg,whr h un-
dertllkvaious kinds of thropologial and thnographi studis for th
Russian rnpir' In his latr yars h am a modrat volutionist' al-
thor"rgh also a didd opponnt of Darwinirr thory.
l-ik Gorgsuvir (1769-1182), Bar hld that th anirnl kingdom ould
sparatdintl lur distiIrtarhtyps:th radiata (.g.,salrfish and s
urhins),th mollusa (.g.,lams and otopuss)'th artilata (.g.,insts
nd ras),and th vtr.lta(.g.'fislr nd humarr bings).H dnid ra-
pitulation thory-th ida tlrat th mryos of mor mpl nimals passd
through nrorphologial stagsorparal with thos of th adult forrns of or.
ganisms lorvr in th hirarhy of lif. H maintaind that th mryo of an n-
imal xmplifid from th ginning of its gstation only th arhtyp or
Urform of that partir"rlrorganisnl. ..Th mlrryo of th vrtrat,'' h as-
s r t d.,. i sa l r a d ya t t l r g i n n i n ga v r t r a t '('1 8 2 8 _ 18 7 , 1 : 2 2 0 ) . S o a h u -
man ftus, hld, w<>uld mov through stagsin whih it would tak on th
forr of a gnralizdvrtrat'a gnrirlizdmamral, a gnralizd primat,
and finally a partiula humn ing. Tlr frm of th growing ftus movd
fom th gnrl to th spifi. Th human mryo in its arly stags' thr_
for,nvr assumdth mtu form of arr invrtrator of a fish.
Dspit Br,s rition lf strit rapitulation, h did allow tht spis
within a givn arhtyp displayd virtually i]ntialstruturs during th
arlirstgsof mryognsis.In a famus passagthat Darwin mistaknly
ttriutdto Lois Agssiz (1s07-187)' Bar mntiond that h had two lit-
tl mryos that h hd flrgttn to lal: ..Thy might lizrds, small
birds' or vry young lrrmmIs.h fomation of th hds nd trunks in
thsanin.ralsis quit similar. Th trrnitisr not yt prsntin ths.m-
bryos.Brrtl,nif th wr in th fist stgsof dvlopmnt,th1.would not
indiatanything; sin th ft of lizards and mammals, th wings and ft
of irds, s l,l,lls th l.randsand tto[ mn dvlop forrr th sm funda-
m n t a lf o r m ' ' ( 1 8 2 8 _ 18 3 7 ' I : 2 2 1 ) .
Though h m wrv of th romnti Nturpbillsophi hiitd y
manv Grman morpl-rollgists, Bar rtaind rtain mtaphysial idas that
onntdlrir .itlrtlr taditilnlf transndntalidlism tht was inaugu-
ratdy Shlling.H smsnvr to hav aandond th onvition, for -
anrpl,tlrt th :rrhtypof th organism, as kind of traphysial ntity'
guidd th ratur's morphologial dvlopmnt. ..Th typ of vry ani-
ml,'' h dlard, ..tlth oms fixd in th rryo at th ginning and
g o v r n si t s n t i r d v l o p m n t ,(' 18 2 8 - 1 8 7 , 1 : 2 2 ) '
Bar ojtd to rlr transmuttionzrlthory th21tWas quikiy oming
rootdin Grmn iology. H undrstltldthat idas of rapitulation and of
442 Br

spis voltion gav sdutiv suor to on anothr) and h firmly op- flutuat in both p
posd thm oth: advan. N4orovr
swampd out. Bar
On gradually larnd to think of th diffrnt animal forms as dvlop'
plants appard sud
ing out of on anothr-and thn shortly to forgt that this mramor-
diatd, and tht in
phosis was only a mod of onption. Fortifid y th fat,tht in th
transitions within
oldst layrs of th arth no rmains from vrtrats Wr to found,
mammals-all arhi
naturalists livd thy ould prov that suh unfolding of th diffrnt
th prsnt tim wa
animal forms was historially groundd. Thy thn rlatd with om-
tntativly ut hop
plt sriousnssand in dtail how suh forms aros from on anothr. ,,for a tru undrst
Nothing was asir. A fish tht swam upon th iand wishd to go for a
i n t l l i g n ' ,( \ 8 7 6 '
walk, ut ould not us its fins. Th fins shrunk in radth from want of
volutionists and al
ris and grw in lngth. This wnt on through gnrations for a ou-
Bar.
pl of nturis. So it is no wondr that lut of fins ft hav finally
mrgd.(1828_1837, |: 200) |LIoGRAP1
Th ommunity of Grman zoologists livd along svral onptual fis- Bar' K. . von. 1828
surs. Som, lik th rdoutal Bar, rjtdth notion of rapitulation R|leio.2 vo|s
and th supportiv dotrin of spis dsnt. Gorg Hinrih Bronn \876. UbrI
(1800_1862)' Darwin's first Grman translator, thought that spis pro- Ntrissensh
grssivlyappard on th arth ovr vast priods of tim aording to a di- 1 1 8 8 6119 8 6
Oppnhimr, d
vin plan. H rognizd that mryoni dvlopmnt or a strong analogy Publiations.
to th morphologial dvlopmnt of spis;ut prhaps faring th ons. D a r w i n , . 1 8 5 9 .o
,l'96
quns of th rapitulational ida, h strssdth analogial haratr of oppnhimr,I.
th paralll and raffirmd th oundris of mryognsisthat Bar had amridg,:
Rihards, R. 1992.Th
stalishd. Othrs, lik th romanti Lornz kn (1779_1s51) and th
IdologilRo
asthtially drivn Gorg Gustav arus (1789_1869), unhsitatingly ad- Prss.
vand th thory of th rhtyp and its attndant notion of rpitula-
tion, whil th mrylogistsFridrih Tidmann (1781_1861) and Johann
Fridrih Mkl (1781*1833),undr th influnof Jan-BaptistLamark,
Bartholomw,
aptd th dotrin of spistransformation. Darwin adoptd rapitula-
Shmidt-Nils
tion thory dspit Br's striturs.H agrd that th mryo passd from
a gnralizd to a mor partiula morphologial stat. H simply hld that Although not mains
th phylognti anstor was of a gnral typ-for instan, that th vo- Knut Shmidt-Nils
lutionary prognitor of man was a gnralizdvrtratwhos form would tion. Bartholomw l
rapitulatd in th human mryo. ..Thus th mryo,'' Darwin o- pionrd th stud
srvd in the origin of Speies (1859, 38)' ..omsto lft as a sort of adaptations to sp
pitur, prsrvd by natur, f th anint and lss modifid ondition of thy dvlopd ask
ah animal.'' ampl' on of Shm
By th arly 1870s th volutinary thsis had swpt through urop' lav- watr and salt balar
ing only th most ralitrant in its wak. Bar undrstood tht th kind of among spis that
mpirial vidn Darwin itd, spially from iogography and palon- hmial proprtis
tology, pointd in only on dirtion, ut h would not ountnan Dar' varials. On of Ba
win's dvis of spis hang-han variation and natural sltion, rows living in salt n
whih ould not aount for th tlologial strutur of organisms. If Dar- a apaility not sha
winism wr orrt, intrmdiat forms should b produd y sltion, salt marshs. Th t
ut non wr found. han variation. whih th thor assums.should inspird, rval how

alothr' and h firrly op-
t nimal forms as dvlop-
forgt that this mtamor-
fid ,vth fat that in th
trtsw to found,
r unfo|dingof th diftrnt
3y thn rlatd with orrr_
ts ros from on anothr.
h land wishd to go for a
rk in radthfrom want f
lugh gnrationsfor a ou-
rt of fins ft hav finally
zrlongsvralonptl fis_
th notion of rapitulation
nt. Gorg Hinrih Bronn
'r' thought that spis po-
lds of tim aording to a di-
pmt o a strong anlogy
r.rtphapsfaring th ons_
c th analogial hartr of
:mbryognsisthat Bar had
okn (|779_1851) and th
i9_1869|, unhsitatingly ad-
ndant notion of rapitula-
r n n ( 1 7 8 1 - 1 8 6 1 )a n d J o h a n n
,rof
Jan_BaptistLamark,
. Darwin adoptd rapitula_
tht th mryo passd from
:al stat.H simply hld that
-._for instan, that th vo-
/rtratwhos form would
rs th mryo,'' Darwin o-
..oms lft as sort of
to
rd lss modifid ondition of
l swptthough urop, lav-
: undrstoodthat th kind of
m iogographyand palon-
would not ountnan Dar-
ation and naturl sltion,
ruturof orgnisn-rs.If Dar-
ld produd y s|tion,
r th thory assums' should
])rtoIomtad Sbmidt-NiIsn 44
flutuat in botlr positiv and ngativ dirtions' thwartirrg volutionary
advan. Morovr, sltd traits would, through random mating,
swampd out. Bar rathr suppsed that th grat lasss of animals and
plants appard suddnly in anint tims, as th gap-riddn fossil rord in-
diatd, and that intrtral,tlologial laws lrad to dtrmin volutionary
trrsitionswithin ah of th largr groups of arrimals-fish' rptils, irds'
mmmals-all..arhing toward partiLllr gorlsand diminishing it ptlwr as
th prsnt tim .wzlsapproahd. Br offrd his spifi hypothsis only
tntativly ut hopd that h had awknd in th radr rognition that
..for a tru undrstanding of natur' W annot dispns with a govrning
i n t l l i g n ' (' 1 8 7 6 , 2 : 4 7 | . R m m b r i n g h i s a r l y w o r k i n m r y o l o g y ,
volutionists and antivolutionists alik rvrd th rat Kl rnst von
Bar.
|LIoGRAPHY
Bar,K. . von. 1828-1837' aikelugsgshiht dr Thir:Btlbcthtugund
Re|leio.2 vols. Konigsrg:Borntrigr.
1876. UbrDris Lhr.Inis St,lif dem Gbitdr
Ntrtuissshfte,2 vls.' 2: 235_480. St. Ptrsurg:Shmitzdorff.
[1886]1986.Autobiogrphof Dr. Krl rst uo Br.2nd d.J'
Oppnhimr' d..;H' Shnidr' trans.Nlv York: SinHistory
Publiations.
Drwin,. 18.59. th rigin of Spis.Lorrdon:John urr1'.
opnhimr,I.|967. sssi th istlr1,o| mbrologd Billlg.
amridg,A: IT Prss.
-h
Rihards'R. 1992. ig lf uolutit:h orpbtlogilostrutild
lf Dri's Thlr.higo:Univrsit1'of hiago
IdellogilRrlstrutil
Prss. -R./.R.
Bartholm Gorge (. |91.9),and
Shmidt-Nilsn,Knut (\9I5 -2007 )
Although not mainstram volutionry iologists,Gorg Bartholomw nd
Knut Shmidt-Nilsn Wr normously influntial for th study of adapta-
tion. Bartholomw rrd Shmidt-Nilsn,with thir ollagusand studnts,
pionrd th study of physiologial and iohmial traits of animals as
adaptatins to spifi nvironmntal onditions. Th rsarh programs
thy dvlopdskd how animals funrilrrirr hallngirrgsitr:arions.For -
ampl,on of Shrrridt_Nilsn,s projts zrskdhow dsrt rrinrlsmintain
watr and salt alar-r.Ths typs of qustions ld to roadr omparisons
among spis that showd how wll diffrnsin physiolgial and bio-
hmial proprtis wr mathd to diffns in partiular nr,ironmntal
varials.on of Btholomw'sproits :rmindhow popultiorrsof spar-
rows living in salt nrarshswr al to obtain watr y drinking sawatr'
apility not shilrd in populations of t sanrspisthat do llot liv in
salt marshs.Th umultiv rsults of thir work, along with th work thy
inspird,rval how wll th funtional svstmsof diffrnt spis'and vn
444 Bts

diffrnt popultions of th sairrspis'lrav n moldd tO mt diffrnt F{ turnd ollttls

nvironmntal irumstns'prslltinga formidal as for th powr and lassi mpl, and
p r i s i t l no f a d a p a r i o n . ral sltionin atior
Brtholomw nd Shmidt-Nilsn wr Ontnrporaris;Barthlorw r- As ws omnlon
ivd his PhD in 1947 fom Gorg Clark arrd Shrnidt-Nilsnrivdhis manufaturir-rgity .
DPhil from Nol Laurat August rogh tn 1946' For ovr 40 yars thy di- duatd hoyist. J
rtd ritial studis in plrysiology that wr nimatd y ologil and pulishing from y
.Wall,
vlutionary prsptivsand th onvition tht nimls wr funtionally frd Russl
intgratd,maning on hd to undrstand th ndrenirnl in its nviron- trst as wll in th
mnt. Both sintistsstudid a wid rang of prolms in a divrsity of ani- rad widly on spi
mls in many nvironmnts. Barthlomw's st-known studis rvolvd naturl law. Thir rt
around tmpr.aturrgulation and nrgy mtrolism'topis that applid to work in Brazll.
survival in xtrm limats, hirnation, an] th nrg'tisof 1lying in I n 1 8 4 8 , B a t sa n d
irds. Shmidt-Nilsn is st rmmrdfor his aminations of watr and to sll oti sPim
slt :rlirnin ]srtnd rrrrinaninrals s rvll s his latr work n th llltingpaid tlri
saling of mtrolirats and th nrgtiosts of loomltion in trrstrial addrss th natur at
vrtrats.ah is wll known among gnrati()nsof studntsfor their stud- spnt th nxt nin y
is of urious or harismti prolms, suh as Bzrrtholomw'ssrutiny of th his spimnsnd n
ad:rptationsto diving in nlarin igunas and Shmidt-Nilsn'sstudy of th xprt on svral gr.
aml's adaptation to dsrt onditions. Bartholomw and Shmidt-Nilsn old and talntdx.
oth wr rnownd lassroom tahrs,mntors' and writrs; Bartholomw Bats'sfildwork s
has ovr 1,000 irrtlltualdstrdants(studntszlnd studnts of studnts) its ttrrtionto th r(
and Shmidt-Nilsn'sttooks ar among tlr most popular nd appaling that Bats and Wall
ollg txts vr writtn. ah inflund iologists far yond his irnmdi- origin of spis.Up
at disiplin' atld ah ws influntial in uilding two of th st zoology d- n irr sris <rft:
partmnts in t Unitd Stats,Brtolomrv t ULA and Shmidt-Nilsn ologial obsrvti
t Duk Univsit. and ]]atstnan
Th intlltul lgay <lfBartholornw and Shmidt.Nilsn an found origi of Speis.
in murnvaras, from th str"rdyof ilhmialadaptation at th molular distriution of ins
ivl to th ornparisolr lf physiologia| prprtis at th popultion nd dn of vOlutiona
spis lvl. Th mhanisti approah to adaptation thet ah promotd winism in natur, wl
rprsnts on Ilollg svral mth<rdsusd to r-rndrstandthis ntr.rlon- stunrring support b
Pt in Darwinirr voltltion' p
for suh a strikir"rg
Bts Ws a rsp
BIBLIOGRAPHY tis, ut his arr
Dwsln,!. R. 200.5.GorgA. BrthlIor'nv's tl itltglrtiv
ontribLttions ltnd vd th old guard.
onlparativiology.Ilttgrtiud omprtiu Bblog1,4 5 : 219_2. British usr'rmlf ]
Shnlidt-Nilsn,K. 199[l.Tll ml's Nls:mlirsof rit>usSitist. origirrsand lak lf s
Washington'D: IslandPrss. -].T,
am asSistantsr
sponsil frth or1
to pr()dua ltlng st
Bats, Hnry Waltr (L825_892) for othr olltors
Britislr naturalist nry Waltr Bats is rditd with th disovry of prot- Alorrg with mirni
tiv mimir. Today, th snario irr whih an tlrganisrr-l rsmlsn unr- Th Nturlist t tl
latd' unpalatalspisis alld Btsian rimiy. Although not th first t immdiat suss'
noti shard apparansrf spis,Bats rinvntd th sujt in 1862. s i g h t s ,a n d s k i l l t l l
 Bts 445

nmoldd to mt diffrrrt H turnd olltos' odditis into i1 powrful as study. Mimir1'eam :l

:midalas for th powr and lssiampl, and on of th most popular ttook iIlustrations,of natu-
ral sltionin ation.
)Irtporris; Barthol omW r- As w:rs ommon in Vitorian ngland, tl,ryoung Bts' a lrk in th
nd Shmidt-Nilsnrivd his nranuftuingity lf Listr, pursud natural lristory as a largly slf-
1946.blrov 40 yrs thy di- duatd hyist. was arr aritious it-lstolltr and taonomist,
:r nimatd y ologil and pulishing fronr a yollng ag in natural history iournals. Wit his frind Al-
that animals wr funtionally frd RLrssl!7alla, h turnd to pofssional ollting. Thy sard n in_
h r r t i r a n i m . r li r l i t s n v i r o n - trst as wll in tl1 thortil qustions ing raisd in th 1840s. Both
f prlmsin a divrsity of ani- rad rvidly on spisnd wr dawn to arly disussinsof volution and
's st-known studis volvd natural law. Thir rading trf trvlrs' ollnts of th topis ld thm to
ta<llism, topis that applid to wrk in Brazil.
nd th nrgtis of flying in In 1848, Bts arld \Wallagan an ploration of th Amazon, arrngig
lr his xamirrations watr and of to sll xoti spimnsto olltors through an agnt in London. Although
rs wll as his lirt work on th ollting pLridthir way' thy hopd tl e al to tls thir osrvationsto
:osts of loomotiot-l ir-rtrrstril addsstlr nat and oigin of spis.Goir-rgsparatwys in l850, Bats
ationsof studrrtsfor thir stud. spnt th nxt trin yars long tlr Arnzll-l.H irmwll known through
s Brtholonr''s srutiny of th his spimnsalrd nts n hvior and distriution.Knor,vrras a taxonomi
c Shrnidt-Nilsn'sstudy of th xrt on svrl groups of insts,Bats lso rnd rdiility y ing
rthol<lmwnd Shmidt-Nilsn old nd talntd xplorr.
ntors, and r,ritrs;Bartholomr, Bats'sfildwork stands out for its rl intrstin distibutionl dat nd
;dnts and studnts of studnts) its attntion to th rlations of spistl ah othr. orrspondrrvals
.\/alla
that Bats and disussd t lngth how sr'rhfats might addrss th
; th rost popular and appaling
liologistsfar yond his immdi- origin of spis.Uporr his rturn to nglarrd in 1859, Bats usd his pri-
ilding two of th st zooiogy d- n in a sris of taonomi plrprs and spis nots, whih inludd his
W t UCLA and Shmidt-Nilsn ologial obsvations.hrls Darwirr agud along lins familiar to Bats,
and Bats amrr immdit' nthusiastiDalvinian aftr rading tb
nd Shmidt_Nilsnan found rigilt of Spis.Amtlng a sris of paprs on vriatitln and gographial
Lialdaptation at th mllular distribution of insts' Bts dvlopd th s lf mimti uttrfliss vi-
)roprtisat th population and dn of volutionary hng' prlviddth dtails of an ampl of Dar-
l daptation that h promotd winism in natur, whr th strugglfor istnld to dptatioIl.It was a
d to undrstandthis ntral on- stunning support aus no prvious natlrrlisti xplanation hd isd
for suh a strikir-rgglhtromnon'
Bats was a rsptd naturalist arrd mmr of tl-rrnajor sintifi soi-
tis' ut his arr path shows that Darwinians did not always or asily in-
vde th old guard. applid ut faild to sur a tlrtrrialpost at th
ontiutionsto intgrativnd
,.l|L|tiL' 4 5 : Z|9 _2-30.
B iol>g British sum of Nrural History. H ws hamprd y is working-lass
1moirsof 'urils Sitist. origins and lk of sial onntions' s rvll s y sintifipolitis. H -
-1.r. am ssistantsrtar}'ofth R<lyalGogrphial Sity,whr h was r-
sponsil for th organization of svrl m;rior pditilns.Bats ontinud
to produ a long srisof taonlmi paprs. H also did ontrattralwork
{or othr olltors,spializingin svraltrtlpial inst groups.
litd with th disovry of prot- Along witlr mimiry thory, Bats am famous through his nrrativ,
t an organism rsmls an unr- he |Jturlist o tbe Riur mzons (786), a lssi still in print today. An
nrimiry. Although not t first to immdiat suss'it was highly praislfor its obsvations' thortiai in-
s rinvntdth suit tn 1862' sights,nd skillful portrayaI tlf t trlpis.
446 rgso

ILloGRAPHY stimulting. His first

Bats' H. \. 1863' Tb Nturlist t th Riur mzllls. London: Jhn urray. ws opnly inflund l
lodd, . [1863] 1892. moir of th author. In H. W. Bates,Th Nturlist o th lishd his massiv ovrv
Riur mllzs, vii-lxxi. Rprint d. London:.|lrn Murray. -'l ludd tlrat vitalism is
oon, |1.P. 1976. Hr Wltr tsls RS, 182.\-1892:plorr, Sitistd world as progrssingup
Drtuii. I'istr:ListshirN[Lrsums.Art (]llris.and Rods Sr-
this supposition.
vl.
Tday, fw would op'
O'ara, J. . 1995. Hnry Waltr Bats-His lif and tlntriutinst iolog.
A r h i u st ' [ N l t t r I H i s l , ' r 2 2 l .| g 5 _ 2 | 9 -w..K' volutionists who rpud
..ordr for fr,'' srt
mhanisti dtrminis
larly tlr motivation
Brgson,Hnri (1859_94|)
Riurtting th Srd'
Hnri Brgson ws a Frnh philosophr who tught for many yars at th
BlLIoGRAPHY
ollg d Fran. H had grat influn in his ountry and in Britain (his
mothr was nglish)'|n \927 h was awardd th Nol Priz for litratur. Brgson,H. 1907.I-'uolu
Born of Jwish prnts' h drw los to athliisnr ut rfusdto pudiat Huxly, l. S. \912. Th l
UnivrsityPrss.
his hitag. did at the ginnirrgof th Vihy rgim from old trght
| 9 4 2 . u o I t t t i ' ' n
whil waitillg for an idntity :rr1' Katlffrarn,S. 2008. Rei,ll
ntrs th story of volutiln aus of hts l-'ulltio rtrie, first Rligio. Nw York: ]
pulishd in |907 (nglish translation, rtiu tloltio, \911)' Brgson, Rus,M. 20,Drtuit
who was influndy th writings of rrt Spnr,always saw volution N4A:Harvard Univr
as an upward progrssto humnkind, ut h did not 21gf with th Darwin-
ian laim tht a blind pross lik rrurturalsltionuld do th jo. Aprt
from anything ls, h ould not s how sltiorrould aout fo th ft
Biogography
that somtimstwo quit sprtlins smdto t:rkth sam volutionary Biogography is t spa
d i t i o n s ,a q u i r i n gt h s m i n n o v t i o n s . trstd in th ology ;
This ld Brgson to introdu his famous, or notorious' notion of a vital glo; th palontologr
spirit, what h alld the l uitl (ls uittu in tlr plural)' This is il for, isms s rvaldthrrug
rthr lik a lif spirit of th kind ndorsd y Aristotl, that suposdly btwn tlr two br'l
guids and drivs th volutional.y pross. Although Brgson pprohd u t i o l - l so f o r g a n i s m si l r
th prolm as ir philosophr' h found support fror som sintists,notaly Th is a parado a
th Grman mryologist Hans Drish, who as a rsult of his primnts ontology. ,For th layp
on dvloping organisms was likwis inlind to a vitalist approah, alling to th point that many ;
lris vrsion of th for t|teetel|ll'. thing thzrt mttrs. For
Vitlism Was i1nathmato most i<lllgists, spiall :rt th beginning of tuth of volution, iog
th twntith ntury whn mn like Jaqus Lob wr ndavoingto put his ws rtainly th
th sujt on a matrialisti and naturalisti sis. Thos attratd tO Brg- rr of natural sltion
son's vitl fors inludd a }'rnh athli prist and palontlogist, ing of his turn to volut
Fathr Pirr Tilhard d hardin. His sin-rligionsynthsisof th vo- diffrnt organisms su(
ltrtionary pitur, sing upward progrssto th Godhd, th omg Point, ia. Thn, famously, h
lrrly owd mtrh to Brgson. Arrtlrr inflund y Brgson was th nountrd olr th Gal
Amrian popultion gntiistSwall \iright,although proaly in is as diftnt' H ould not
th dirt influrl of Spnr'swritings Was mor signifiant. A third n- th mainland and thn t
.Dhn
thusiast Was th nglish iollgist Julian Huly, grandson of Thomas Drwin am t
Hnry Huxly nd oldr rothr of novlist Aldus Huxly. Sarhing for (
Speis 1859)' palont
a non_God-sd rr-raningfor lif' Jlian Huly for.rndBrgson's idas rord, tlr irpparnta

'|1zo|1
s. l-ondon:John urry.
H. \V. Bats' The Ntur|ist ll th
lon: John urray.
i_I 8'92:pllrr, Silttist d
;, Art Gallris,and Rords Sr-
nd ontributionsto iology.
_.w..K.
lo tught fr many yars at th
Lhis ountry and in Britairr (his
.d th Ntll Priz for litratur.
roliismbut rfusdto rpudiat
/ihv rgim frorn a old ught
i is L'uoltirl r,:ttrie,first
'tiuelloltil, 1971 Brgson'
)'
:t Spnr'always saw volution
. did not grwith th Darwin-
:ltionould dl th jo. Apart
:tion orrld oul1t for th ft
d to tak th sam vllutinary
. or notoriotrs'notion of vitl
lx irr th plLlral).This is a for,
d y Aristtltl' tlrat sr"rpposdly
Although Brgson pproahd
ct frol sol silltists, nltly
| o a 5 ; l s u l to f h i s p r i m n t s
d to a vitlist pprolrh' lling
s, spiallyat th ginning of
s Lo wr ndavoring to put
asis.Ths attrtd to Brg-
oli prist and plorrtologist,
r-rligionsyntsis of th vo.
th Godhd, th Orrrga Point,
nflund y Brgson Was th
rt, althoughproaly in his s
as mo signifint. A third n.
Huly, grandson of Thomas
t Aidous uxly. Shing for
uIy found Brgson's idas
Bilgogrpb1, 447
stirnulating. His first ook, Th ldiuidul i te itl igdlm (1912),
was opnly inflund y Brgson's thinking. B 1942, whn Huxly pu-
lishd his massiv ovrviw, uolutiott: Th odr 31,thsis,h sadly orr.
ludd that vitalism is no asis for sin' ut h still saw th volutionary
world s progrssingup to lrumankind, tying to giv naulisti bking to
this supposition.
Today, fw would o.lly susi to an l UitIl'vt many suspt that
volutionistswho rpudi.1tstrit Darwinism in favor of ltntivs,suh as
..ordr
for fr,'' srtly]harbor philosophial yarnings against lind nd
mhanisti dtrrninisnr, th sal dislik that nrtivtd Brgsoll. lris is
larly th motivation of th mrrvmnt's ladr' Stuart Kauff-ran, in his
Rinutig th e Sred.
lBLloGRAPHY
Brgson,11.1907. L'uolutiortri.Paris:Alan.
Huly,J. s. L912.h ldiuiduli thiml igdlm.amridg:ambidg
Univrsitrss.
1942.uoltil:h odr Sthsis.London:Alln and Unlvin.
Kauf{man,S. 2008. Riutigt Srd: N Vituof Si,Rs,d
Rligil.Nw Yrk: Basi Bloks.
Rus,M. 203,Dri ttdDsig:Does uolutilue Purpos?mridg,
MA: arvrd Univrsity Prss. -.R.
Biogography
Biogographyis t sp as palontology is to tim. Th iogographris in-
trstdin tlr llogynd volution lf orgilnisms s distributd arourdth
glo;th palontologistis intrstdin th ology and volution of organ-
isms as rvaled through th fossil rord. Of ours, thr is muh ovrlap
twnth two ausoth ar intrstdin how th past influnsdistri_
u t i o n so f o r g a n i s m si n t h p r s n t .
Thr is a parado out th rsptivstatussof iogogphyand pal-
ontology. For th layprson thinking aout volution, fossils om to mind'
to th point that n-ranypopl thirrk tht th fossil rord is vitually th only
thing that mattrs' For th volutionary iolgist sking onvition of th
truth of volution, iogographyis simply outstanding.
This ws rtinly th as for harls Darwin and irlso for th odisov_
rrof natural sltion'Alfrd Russl \/alla.In his autoiograph, in talk-
ing of his turn to volr-rtion, Drwin highlightd th w in whih sirnilar ur
diffrntorganisrssuddalr othr as h journyd down South Amr-
ia. Thn, filmously' lr spok of th organisms (tortoissand irds) tlrat lr
nountrdon th Ga16pagosArhiplago irr th Pifi: similar ut slightly
diffrnt.H ould not s how this ould unlss foundrs had om fronr
th mainland and thn volvd as thy sprad Out aross t islanlgroup.
lfhn Darwin m to prsnt lris thory of vllution in tb rigi of
Spcis(1859), plontologl. was somWht prolmati-gaps in th fossil
rord,th apparnt asnof lif forms for th amrian, and so forth.
148 Biogogrph

But iogograph rvas a triumph. Patiulariy important for Darwin was tlr liAll]t|D
Way that th inhaitants of islands ar so simila to th inhaitants of thir
losst ontinnts and not to th inhitants of lthr ontinnts farthr i'ALAYSIA
away. ..Th most striking and imprtnt fat for us in rgard tl th inhai-
sttGAPoRl
tnts of isl:rrrds,is thir ffinity to th<lsof th nsrrrrinlnd,rvitlrtrt-
ing atually th sanrspis.Nunlrous instansould givrrof this fat.
I will giv only tln, that of th Glapagls Arhiplago, situatd undr th
quator' twn500 nd 600 mils from th shosof South Amria. Hr
almost vry produt of th lnd and Watr rs h unmistakalstamp of
t h A m r i n o n t i n n t ' '( D a r w i n | 8 5 9 , 4 1 6 \ .
.s7all J
was likrvisvry snsitivto th signifinof iogography.His
first papr hinting t volution (in 1855) flusdon th ways in whih or- t 0 cs
ganisms sm to pttn thmslvsround th glo. H found that nwly Wi
arrivd organisms st-nalwys to onnet with livirrg organisms los to
thm...vr spis lras om into xistnoinidr-rtoth in spa arrd
t i m r v i t h a l o s l y a l l i d s p i s ' '( . W a l l a 1 8 5 5 [ 1 8 7 1 l , 2 5 ) . I n T h c e o -
t
N
i t i d i f l s

0 00Mi|es
FF
grphil Distributillt of ils (1'876)l/allaturnd to issus in io. 0 300Kilometers
gograplry,lolking oth at th spial issusthat suround islands as wll as 100B
1100
road qtrstionsof distiution. \Vll krrown is th sl-lldVallil's lin
(sfigrrr), brirk twntwo vr diffrrrtsts of organisr-ns, Asian and Walla,slin. Whn h w
lWalla rea|ized that thrt
Australasian, running hrough th Mlay Arhiplago, twn Borneo nd
Asia and Australasia.He <
Sulawsi, and twnBli (wst)and Lomok (ast).
in Th ogrphil Dist
A majlr qury for ths nintnth-nturyvolutionistsWas iltly how
a voagerwith Frdinnd
orgnisrs gt disps].Darwin spnt onsidrabl tinr running pri-
mnts trying to shorv how things lik sds anl slrils ould rvitlrstandslt
watr nd gt arrid y jtsam or y irds. H disussd his rvork tn-
sivly in te rigi. A liffrntapproeh was takn y Darwin's los frind, Aftr th synthti th
o t a n i s tJ o s p h D l t n H o o k r ( 1 8 5 ) . H a r g u d t h a t p l a n t d i s r r i u t i o n s tory of iogography w.
wr rtr xplaind r, on_istingland ridgs tlrt hav now disap- 1960s. Plat ttoni th
pard thnks to th ising sa lvls.This was prt of a thOry of Iimat, ontinntal drift. It now
proposd y goIogist harls l-ylt (1830-183), whih supposd that al. that had onrnd th n
though ontinntsannot mov sidwys' thy ar alrvayshanging shap - of lost ontinnts ut of
aus th arth's surf is forvr risirrgor flling. groups away from ah .
Btwn Darwin's tirn and th frlrrr-ringof tlr synthti thor in tlr th sam tim gology
1930s' iogographytook ak sat to palontology as most ffort was di- pothses.A lassixam
rtd toward th tring of arth'shistory. !7hn sltionthory am ak of Lstrosurs. This hr
into fashion' most attntion was peid to rirovolutionary usal issus' million yars go' is fat, r
prtiularl,vth ways irr r'',hihgns s;lrad and hrrgin popularirrs.Yt hav bn a world trav
thr ws som rvivd intrsti iogograplr,v. Partiularl implrtantWas found in th sam fossil
th influn of systratistand ornithologist rnst yr. In his lassi Sys- (Southast)Asia, and An
tmtis d th origill of Specis (\942) h disussdin dtail issus to do fat that, mor than 200
..rings thy wr part of Panga
with distriution' most notaly highlighting th irrrportantlf of
ras.'' Hr, on hs a rirrg of adjar-rt nd intrdingppulations of th gologial hypoths
..Th I
spis' lrut whr tlr nd points ar rprodutivly isolatd. Nlayr argud in th min ssay
that this is dfinitiv vidnof th formation of spisin a grdual rathr Th dvlopmnt of
thn a suddn pross. idas about how to r
 Biogeogrph 449

ortant for Darwin ws th TliAltAl'ltl PlLlPPlNs

to th inhaitants of thir
I othr ontinnts farthr II,IALAYSIA t'jlAlYslA
us in rgard to th inhai- stGAP0
rst minland, without - )Uquao
ould givn of this fat.
plago,situatd und th
Sumatra
e,
rso{ Soutlr Amria. Hr
.hunmistaklstamp of 3

e,
anof iogography. His .
100
s
l on tlr ways in whih o-
Job. found that nwly Wal|aeLine
living organisms los to
inidnt oth in spa and
N
t i**iin*i:**l

' [ 1 8 7 1 ] ,2 5 ) . I n T b G o .
turnd to issus in io-
surround islands as wll as
th so-alld \7alla's lin
ts o{ organisms, Asian and
lago, twn Borno and
sast).
olutionistswas atly how
r l t i m r u n n i n g
snails ould withstand salt
disussdhis work xtn-
ln y Darwin's lcs frind,
udthtpln distriuions
idgs that hav now disap-
r part of a thory of limat'
3), whih supposd that l-
:ahvayshanging shap -
tq
tlr synthti thory in th
tology as most ffort was di-
n sltionthory am ak
o v t l l u t i o n a 1 1a u s a l i s s u s .
J hangin populations. Yt
,. Partiularly important was
nst Mayr. In his lassi Sys-
sussdin dtail issus to do
.h importan of
trrdingpopulations of a
:tivlyistllatd. ayr argud
of spisin a gradual rathr
0 300Miles
FF
0 300Kilometers AU$RAL|A
,120U
1000
11 0
130! 1400
!alla's
lin. Whn h was ollting spimnsin th Far ast' Alfrd Russl
!allarealizedthat thr was a lin of sparation twnt organisms of
Asia and Australasi.H disussdthis and othr iogographialphnomna
tnTh Gogrphil Distributio lf nimls (1876). Th lin was first notd y
a voyagrwith Frdinand agllan in 1521.
p i -
Aftr th synthti thory was wll stalishd, th major vnt in th his-
tory of iogography was th oming of plat ttoni thory in th arly
1960s.Plat ttoni thory offrd gophysial xplanations for th ida of
ontinntaldrift. It now Was possil to addrss proprly th kinds of issus
thathad onrnd th nintnth-nturyvolutionists. It was not a qustion
of lost ontinnts ut of ontinnts that mov sidways, arrying on-joind
groupsaway from ah othr. Biogography rivd a powrful tool and at
th sam tim gology rivd strong onfirmation for its audaious hy-
pothss.A lassi xampl of th two-way pross is givn y th distriution
of Lstrosrs.This herivorous, mammal-lik reptil, found mor than 200
million yars ago' is fat, short, and squat. It is rtainly not a ast that wold
hav n a world travlr' rgularly swimming aross oans. Today, it is
foundin th sam fossil dposits (Lowr Triassi) on th ontinnts of Afria,
(Southast) Asia, and Antartia. This would a miral if it wr not for th
fatthat, more than 200 millin yars ago' all thr ontinnts tohd whn
.,rings of thywr part of Pangaa. Th fossils support th gologial hypothsis, and
thgologialhypothsis maks snsof th fossil distriutions (sFigur 27
in th main ssay ..Th History of volutionary Thought,' y A4ihal Rus).
Th dvlopmnt of plat ttoni thory oinidd with th advnt of
idsaout how to ronstfut spis phylognis using a st of xpliit
4s0 Biogogrphl

ruls, that is, idas aout ladistis. Th fusion lf ths onpts ld to th A mjor iogograpl
shool of viarian iogography, artiulatd in th 1981 volume Sstem- al illgist Rort
tics d Biogogrph: ldistis d Virie, ditd y Garth Nlson d w r d ( ) . W i l s o n 1| 9 t
and Nomn Platnik. Viariarr iogographrs, following Vnzulan lands in tlr s or oas(
Lon roizat, rrrphasizdth ir-nportanof sahingflr omnron iogo- iving and spis l
graphi pattrns mong unrlatd groups of organisms and dignlsingth funtion of suh fro
rol of gollgial vnts in rating thos pattns. For xampl, a lassi r h a r t l f t h i s | a n di
viarian analysis would idntify th ommon pattrns of rang limits and oIonizitln nd in
distriutions rf gnti variatitlrrmong s<-ruthstrn U.S. fish spis,pat- urv' a mathmatial
trns tied to oundaris t major rivrs' and diagns th formation of rivrs fourrd in lt lo:rti<ln inr
and th istn of old stuaris nd sa hnrrlss paramount vnts in SPis-lluvshlltl
braking on-ontiguous distiutions and ating prsnt-day pattrns. offrd th first quantiti
In ontrast, so-alld disprsal iogographrs argu that th disprsal of T h o l o g i |t h
spis aross road rangs and hangs in dispr:salrats aross major fa- usd th thory to rir
turs lik mountains or rivrs ar mor imprtant for prsnt-daypattrns. t h d i s t i u t i o no f s p
Th prlpr nswr is surly a omination of both viarin and dispr- a t i n 8a r . r g n l n r l
szrl.Drift and tlr susqr.rnt irrrirsar vry imptrrtant.Howvr, no on a u s l m h n i s m st
truly an say that ontinnt:rldrift is th only ky ftor in organi disprsals. of additionl ioggr
assiv disprsals hav ourrd twn Nortlr and South Amia as th rangS also tnd to hav
land links hav n stalishdtwn thm: th ..Grat Amrian Intr- Thr hav nsv
hang'' (rshall et a\. \982 s Figur 26 in t-hmain ssay ..Th History t h o y t l f M A t h u rl
of volutionary Thought'' y Nlilral Rus in this volum). This is th ra- S p k < l s k( i| 9 7 8 , 1 9 7 9 ,
son marsupils, suh as Opossllnls,xist in Notlr Ar-rrriaand r<ldntsp- argr.ringthat t rtin1
par in South Amria. (oviously, th golgil hngs hav tin part mpty r-rihsthat ar t
a funtion of drift; it is diffiult to kp th two fators apart.) rht mth th prdi
..Palontology
It is not orrly thanks to gology tht inrasing intrst is ing takn in and th
ausal fators in iogography.arly n, population gntiistsralizd that Th thlr of th ttl
gographial distriutions ar sigrrifiantin thi str-rdis. A lassi s was s t i k i n g x t r t s i o nI.n t l
th distriution irr Afria of sikl-ll nmia, a dissin whih th vitir and, on irlr islnd on
usually dis for rahing his or hr fifth birtdy. Why dos it tlur in i n i r lp o p u l t i O n x p e
rtain pats of th ontinnt trt not in othrs? In on dirtion, it was pos- d l i n l l n d t i n t i o nr
sil to pin th disas down tr th possssion of a rtain kind of gn- d i t i o n s . l t i s , i n f te,
homozygots for that gn show th diss-and it was shown that th gn prosss rvith volutio
.Why
auss th ollps f rd lood lls. But why th distriution? in th tions ol1stntlyo
Gold oast ut not in South Afria? Th answr am austh gnsin y Rort Riklsnd
singl doss (htrozygots)onfr a natural immunity to malari, and th this pt-tssin Wst lnt
mosquito is major pst in thos and only thos aras with th gn.In othr ogy and volution oul<
words, th distriution is linkd to sltionstting up arrirs against inf- T h s o n n t i O n s t
tion (Allison 1'954a,954). standing iogographi
\/ithth advnt of molular nrthods,population gntiistswr 1to t h t a n i m l I s i n r a i r
tnd si iogographil prinipls to amin larg-salpatrrns of pttn is vidnt in
gn distriutions in spa. Fron thspattl-ts, iologists hav n al to plnations not in sir
draw infrns aout th likly loations in whih a spis or group of h t I n t l r f | t i v | yl
spisoriginatd and th history of disprsalinto its prsntrang. Ths r- humidity, growing sas
onstrutions of history, alld phylogography, rprsntth modrn vr- Biogography ontin
sion of Darwirr's and !alla'spiorrringplratilr-rs. o n p u i l lu n d r p i n n

ion of tlrsonpts ld to th
d in th 1981 volume Sstm-
,i, dited y Garth Nlson
raphrs' fllowing Vnzulan
:sarhingfor ommon iogo-
I organisnrsand diagnosirrg th
)attrns.For xampl, a lassi
lon pattrnsof rang limits and
l t h s t r nU . S . f i s h s p i s .p a t -
diagnosth fomation of ivs
h n n l s s p r m o u n t v n t si n
[ rating prsnt-dy pattrns.
1rs argu tht th disprsal of
rlisprslrts ross majo fa-
)rtnt for prsnt_daypilttrns.
n of oth r,iarianand dispr-
:ry important. owevr, no on
.ky fator in organi disprsals.
rlorth nd South Amria s th
..Grat Amrin Intr-
Lm:th
..Th History
i in th main ssay
in this volum).This is th ra-
North Amria and rodnts ap-
o g i l h a n g sl l a v b n i n p t mptv ihs that ir thn oloniz.dy organisnls at rts arrd with rsults
:wo ftorsapart.)
:asingintrstis ing takn in
lpulation gntiistsralizd that
thir studis.A lassi as was
ti, a disirsin whih th vitim
r l r t h d a y .W h y d o s i t o u r i n
rs?In on dirtion, it was pos-
'sion of a rtain kind of gn-
-and it was slrown that th gn
vhy th listriution?\/hy in th
n s w r m a u s t h g n s i n
al immunity to malaria, and th
hosaraswith th gn. In othr
l sttingup irrisagainst inf.
opulationgntiistswr abl to
amin larg-salpattrns of
rns,biologistshav n bl to
i n w h i h s p i so r g r o u p o f
al into its prsntrang. Ths r-
:aphy,rpsntth modrn vr-
:olorations.
Bi-lgogrpb, 4'\ 1
A rrrjor iogogaplriaillogial tlrory ppard in th 1960s. Thorti-
al iologist Robrt aArthur nd inst spiirlis(and futur soioiologist)
dward o. !7ilson (1967) agud that spisnmrs on islands-whthr is-
lnds in th sa or Oassin th dsrt-rh n quilirium vn spisr-
riving and spis lving (prhirps through tintion). This alan is ;-r
fur-rtiln
of sulr ftors as tlr distn lf tlr islnd fror tlr rr-rainiandnd of
th ra of th islnd itslf. In aldition, aArthur and ./ilson dsrid how
olonization and tintion ould ornin to gnratth so-alld spis-ara
urv, a mathmatial rlationship tht dsris how th numr of spis
foud ir-ra loation insswith tlr ara (suakilomtrs) of tl.latlotion.
Spis.raurvhd n t]sribdin tlr,p:rst,ut MaArthur and $ilson
offrdth first qtrntitativplnation for th shap f thos urvs.
Th ologial thory inspid a rnaissanin iogography. Biologists
usd th thory trl rintrprt arrl unify smingly disparate pattrns, fron-l
th distriution of spi'son isoltd mountintops to th ollsqunsof
rtinga fragnrntdhaitat fronr ontinuus forst.Th ttmpt to tst th
ausal mhanisnrsthat ould rate spis.aurvs ld to th disovry
of additional ilgographipattrns,suh s tlr fat that spiswith lrgr
rngslso tnd to lrv highr population dnsitis.
Th hav rrsvlinrportantvolutilnilytIlsions<lfth ologil
.Wilson's
thory of aArthur and \7ilson. studnt, palontologist J. John
Spkoski(1978, 1979,1984), applid th thory to hanging divrsity in tim,
arguingthat at rtin priods' suh as th girlningof the amrin, W s
that 1athth prditions lf tlr iogograplrithory (s th main ssay
..Palontology
and th History of Lif'' y ihal Bnton in this volum).
Th thory of th taxon yl, originally dsrid y !7ilson, is a partiularly
strikingtnsion.In th taxon ,vl,spisdisprsthrough an arhiplago
and' tll-tah island on whih rh1-prsist, undrgo an volutionirryyl of
initial populatill xpansion, daptation tl island onditilrrs,and vntual
dlinand tintion undr th prssurof nw llonistsand hanging on-
ditions.It is, in fft,a lif yl of island iota. This thory linkd ologial
prossswith voltrtionaryons nd offrd an xplanation for why xtin.
tilnsa onstrrtlyorring,llrit n a vry lorrgtimsal.Gnti studis
b R<lrtRiklfs and ldridg Brmingharrr(2007) rvldth signaturof
.Wilson's
this pross in sst Indian irds and vrifid insight into how ol.
ogy and volution ould onntdthrough igography.
Tlrsonntions ar also ing mad y ltaildinvstigirtionsof long-
str-rdingiogogrphipattrns.For xar-npl,Brgmarrlr'srul (1847) st:rts
tht animals inrsin siz with ithr inr:rsinglatitud <rrlvation.This
pattrn is vidnt in wrm.loodd and old-loodd animals and has its
plnations nrt in simpl thml rlationslrips (.g., largr animals hold
hat mor fftivly)ut in th otnpl rltilnshipsamong tmpratur,
humidity,growirrg sason' and nrgtiffiinis.
I}iogographyontinus to fllurish,as it did in Darwin's day. It offrs th
onptual undrpinning for applid prolms lik th dsign of natur
452 Birds

rsrvs and ontinus tO play k rol in tlr onfirmatoy support tlf vo- h immdiat an
lutionary thory. that rsmld anima
holsts' Fathrs,whi
lLIoGRAPl.|Y
atually volvd rvitlr
Allison, A. . 1954a. Prottilt-t th sik|.]ltrait gainst sutrtinralriaI ovring in anir-nalssu
inftion. B rit i s h di l J rl r l 1,: 2.L)
.
loet pinnr sh
1954. Th distriutilrof th sikl-lltrait in l]st Afri:r arrd lswhr
heoptr. Thir pr
and its apprnt rlationship to th ini]nf subtrtianmli.
rnstios o| t Rrll,l ,\oit tli rtlpil cdil Hgict, 48: |2. would hav svd a
Avis, J. . 2000. Phlogeogrpl:y.mr.idg,MA: rvrd Univrsity Prss' t h i r i n s u l t o r yP r O p
Brgmnn, . 1847. Uer di Vrhiiltnissdr wdrmokonomi dr Thir zu ihrr rytially olord w
Gross. Gttigr Stdi.]. no. 1: 59-5_708. tion, or amouflag;s'
Darwin, . l8.59. () th Origi of Spal-..Londo: .)lhnlvlurv.
nough vidn of ol
Hookr, J. D. 185.]. Itrlduttt ss tl tb F|or f.Ntu Zelnd. l,ondot-t:
Lovll Rv. f a t h r so n t h f i n g r s
Lomolino, . V., B. R. Riddl, ltnd J. H. Blwn. 26.Biogeogrp.3rd d. gsts thi1tth fthrs
Sundrland, lr{A: Sinur Assoiats. posttl prsvdin s
Lyll, . 1830_] 83. Priipl,s tf Golog1,:Big- ttmpt to pli th I f t h p r s n o f I
Former hgs i th rth's Surf b Rfrc to ss Nola i
flight volv? rlttl
prtio.3 vols. l-ondon: Jhn urry.
i s t h 6 r s r a n i m l i r rt l
aAthrrr, R. .. r-r] . O. Wilsln. 1967. he h<t\'tf Isld Biogogrph1'.
Printon, N.}:Printon UrivrsitPrss. f o r m w i n g p l o
.!,
arshll, L. G., S. D. J. J. Spkoski Jr., nd D. . Rup. 1982. ammlin sit flight strok and u
vlution and tlr grat Amian intrhng, Silt2l5: 1.5\_157. faturs is indirt but
Mayr, . 1'942' Sstttisd th rigi tlf Spis, Nr,v York: (]olLrmi S i n t h l 8 8 0 s , w
Univrsity Prss.
hav n priodiall;
Nlson, G., and N. Platnik. 1981. Sstmtltisd Biogogrph: ldistis d
Viri.Nlv York: olLrmbi Univrsit Prss. and usd inipint win
Riklfs' R. ., and . Brminglrn. 2007, Th ussf vllutionrvrditins in Th sond is tht ir
arhiplagos:Pssritrirds in th l-ssrAntil|s. mrill Nturlist |69l. nt willgs to gin inr
285-297. p d r ( ) r s .V a r i o u s m t
Spkoski, J.J., J.. 1978' kinti modl of Phrrrzlitaxonomi divrsity.
nithr lrypothsisis tt
I. Analysis f nrirrinordrs. Plrlbi<ll<lg1,
4:22_251'.
|979. ^ kinti modl oi hrrrzoitaxot-tlrni divrsit. lI. arly
storsould lim r
Plozlifarilisand muItipl quiliria. Plo b i tllog 5 : 222-25 2. d o s n o t g U a a n t f l l
1984. A kinti modl of Phanrozi taxonomi divrsity.IIl. Post- trstrial ipds and 1
Plozoi familis and mss tintions. Plobilllgl,10: 246_2-67. liftoff?
.!(alla'
A. R. 18.5.5[l871]. Or-rtlr law lvhilr lras rgultdth introdution of nw Rnt rsahhas
spis. Als nd g:in of NtrI Histor 16: 184-196. Rprintd in
lution of th flight strt
ontributkls tll th Tlr of Nturl Sltio. Lndln: millan, 1_25.
|876. Th GogrphitlDistributitlll lf imls.2 vols. Llndon: movmnt ol, th for
Mrnillan. Deio,bus and VI
1905. 1' l-if: Rlrd lf uets nd o1liltions. Londn; hapman nd i n h r i t d :t h o n s
Hll. -I'. d .R.
o n s h d l n o d i f i dj t
thos prdatory dinos
Wr gly lollgtd
Birds
lik ird wings. Whn
Sin originating from small arnivorous dinosaurs in th Lat Jurassi whippd forward and
(aout 150 million yars go)' irds hv raditd into sm 9,600 spis gatd fthrs,dlivr
around th glo. Thir olors' songs' havios,and ologial adaptations th most frqurrtlyus
ar ndlssly olplx. What sorts of fators hav n ky in thir volu- Of urs, flight s
tionary radition? pod Di()hus \a
 Birds 15

1onfirmatorysupport ot vo- Th immdiat anstorsof irds wr small' prdatory, ipdal dinosaurs

that rsmld animals suh as Vloirptor, irorptr>r, a'd Surorit-
holestes.Fathrs,whih W usually think of as a diagnosti fatur of irds,
atually volvd within rrrivorous dinosaurs, first as a short filamntous
trait aginstsutrtiann-rlarial ovring in animals suh as Siosuroptr, and thn in inrsingly mor
laorat pinnat shaps as in BipiosL|rus' diptr, and Protr-
raitirr ast Afria nd lswhr
hopter. Thir,rimary funtion was oviously not for flight' ut thy
of sutrtianrnalari.
| diI Hgie 48 31'2,
would hav srvd a narly automati thrmrgulatory rol y virtu of
.Whthr
[: Hrvrd Univrsity Prss. thir insulatory proprty. thy wr rightly' haratristially, or
rmokonomi dr Thir zu ihrr ryptially olord would hav dtrmind rols in display, spis rogni-
tion, r amouflag; som olor anding appars in arly fathrs, ut not
on: Johrrurry. nough vidn of olor rmains to indiat othr rols. Th prsn of long
t of Ntu ZId. Lond<ln:
fathrson th fingrs of at last on oviraptorid thropod, udiptr' sug-
206. tsiggrh,3rd d. gststhat th fthrs ould hav n sprad ovr th ggs whn nsting-a
postur prsrvd in svral fossil xampls.
Atttpt tl pli th If th prsn of fathrs did not automatially onfr flight, how did
'frtl uss Nltu i
flight volv? rheopterx, univrsally rognizd as th first known ird,
is th first animal in th thropod-ird lin to hav fathrs larg nough to
Thor f lsld Biogogrph.
form a wing apal of sustaining th animl in th air, assuming th rqui-
d D. . Rup. 1982.Mammalin sit flight strok and undrlying mtaolism Wr prsnt.vidn of ths
e . S i 2 1 5 : 1 3 5 1 _ 1 3 5 7 . f a t u r si s i n d i r t u t s u g g s t i v .
,isN . w Yrk: llumlli Sin th 1880s, two ompting hypothss for th origin of ird flight
hav n priodially dbatd. On is that ird anstors limd trs
d Bilgogrh:Idistis d
and usd inipint wings fist to parahut, thn to glid, and finally to flap.
Prss.
r u s s f v l u t i n r yr a d i t i n s i n Th sond is that ird anstors ran along th ground and flappd inipi-
\ntills.mri NturIist |69l nt wings to gain inrasd hight, ithr in jumps aftr pry or away from
prdators. Various modifiations of ths idas hav n proposd, but
r o z i t a n t l m i d i v r s i t . nithr hypothsis is tstal.Thr is no onvining vidn that ird an-
:22_25]'. storsould lim trs, ut most small animls an; howvr, arorality
onomi divrsity.II. arly
ilelbitlo g 5 : 222_2 5 2.
dos not guarant flight ility. It now appars that ird anstors W
onoi divrsity.IlI. Post- trrstrialipds and good runnrs' ut wr thy good nough to ahiv
obkllog 10 246-267 ' liftoff?
as rgulatdth intrdution of nw Rnt rsarh has shiftd th fous from ts vrsus ground to th vo-
istor16: |84_|96. Rprintd in lution of th flight strok itslf. This strok had its origin in th prdatory
,tio. London: amillan, 1_25.
.imIs' movmnt of th forlims of small thropod rlativs f irds' suh as
2 vol'. Londn:
Deinochus and Velocirptor. These dinosaurs shard a fatur that ids
pikls, London: hapmn and inhritd:th ons of th wrist that onnt with th palm (mtaarpal)
-J.T. nd .R. ons had modifid joints that swivld narly 180. from sid to sid. In
thosprdatory dinosaurs, this joint prmittd th hands and fingrs,whih
Wrgratly longatd,to foldd los to th ody whn not in us, muh
.Whn
lik ird wings. ndd to dlivr a prdtory strok, thy ould
dinosaurs in th Lat Jurassi whippd forward and downward. It is narly this xat strok, with lon-
radiatdinto som 9,600 spies gatd fathrs, dliverd at a slightly diffrnt angl, that onstituts on of
|viors,and ologial adaptations th most frquntly usd flight stroks of irds.
)rs hv n ky in thir volu- of ours, a flight strok is uslssto an animal of th siz of th thro-
pod Deionhus (aot 25 kg), ut th first irds Wr muh smallr;
154 Birds

rhtlptrl spil-nnsran.from pig,ontO l.ow siz' Nw wok on th Gauthir,J. A.' and L. F. G

mirostrutur of on tissusin dinosaurs and arly irds shorvsthat dinosaurs uolutiotlof Birds.N
as a group grw quikly' at rats omparal to th()sof m<lstmammls and Padin,K., and L. . hia
Biologil Ruies(z
irds' Howvr' th bon tissttso[ th first irds, suh lls ofuittstlris' . h o r i g i n.
1 9 9 8 T
rord rlatir.lyslowr growth during just thos stgs(juvrliland suadult) 3847.
whn typial dinosaurs grw most quikly. As a rsult, irds fftivly bam Prum, R. O., and A. ' Bru
mini:rturizddr.rlts. At tlris smallr siz,th rvuld lrvhd gratrratio of SintifiAmric,
wirrg surfa ara to ody mass, and this improvd wing loding would hav
n arodynamilly:rdvantagous.
Buffon, Gorgs-L
Did th first irds lrilv th rrrtabolisrrrnssarv for sustaind flight?
..Thr is no haptr of
Agin, dirf msurmntis impossil, ut th growth dvnamis of arly
irds and thir dinosaurin rlativs hav n ronstrutdfrom on tis. mor ontrovrtd than
sus.Th higlr rats of strstindgrowth rfltdin th tissusmaks it diffi- i o y 1 9 5 9 , 8 4 ) . L o v 1 o yi s
rrlt to draw any orrlusiorrothr thirn that dinosaurs, and thi avian phr playd as pomin
dsndants,had rlativly high mtaoli rats.Th prsnof fthrsis a Gorgs.Louis Llr d
furtlrr indiarion. amiguous in thir pin
Part of tlr shift to flight involvd a rorganizationof th llomotor mld- wrot at a tim whn th
uls of ipdal dinosatrrs.In ths dinosaurs,th arms Wr not usd i loo- highly omplx-was n
motion; th lgs proplld and sr.rpportdth animl' nd th tail assistdin volutionist' in othrs h
ioomotion in an ir-rrportant way. Th lng musular tail, lor lik a roo- qustions nd to dist
dil tail than lik tails of irds today' attirhdto th spins and prosssof fon an volutionist? (2)
th tail vrtra,and insrtdtoward th tlp of th thigh on nar tlr hip ida of organi volutlo
sokt. lts rol in pr"rllingak th lg, :rnht>ring tlr strok :rgainstth stabi- nd to valuatdin t
lizing tail, would hav n sustantial.In th dinlsaurslosst to irds, th Buffon's work touhd
tail am rdud arrd stiffnd,and this musl flund a mor important th Paris Aadmy of Si
rol irr ontolling movnlnts of th til in running nd in fliglrt.Th for- intndnt (had) of th .
lims, of ours' atnwings; instad of hving :r two-modul loornotory oldst sintifi instituti
systm in whih th lgs and tail workd togthr, irds volvd a thr- th ollg d Fan. ([
mldr"rlsystl-tlin whilr tlr wingd forlirr-rs,th lgs, and tlr tail elr pr. rrgnizd and rnam
forrns sparat funtilns in th air and on th ground. Phylognti trnds Jadin du Roi, Buffon
show that in th arl volution of ids, faturs of th flight mhanism 1741 he translatd N
(suh as th lula anl th ftrsior-rf orrs)volv{and rfirrd funtions (|736\. H lso produ
more rapidlv thn did ftursof th hindlims (stlhs prhing laws) and mography. From 1749 t
othr haratrsrlatd tl fding and loomotion. lishd in the 44 volum
Judging frorrrth disrriutiorrof ftursin irds and othr dinosaurs, th [Natral history, gnra
first irds proly had largr luths than th on to four ggs of today's laoration with svrall
irds. Thy wr likly prlialrathr than altriial' and thy would hav ton and Gu6nau d
uilt nsts on th grollnd rathr than in rrs;sol prnt:rlar is likly. dals in dtail with subj
Somtim during t rtous, fo th divsifiation of th living applid mathmatis,zo
groups of irds, growth rats apprntly inrasd,so that adult stagswr volution (to us th m<
rahd mr rapidly. Birds tldi.ryrah ftlll siz ar intrvals ranging from s n s i n t h i g h t n h
s v nd a y s ( s p a r o w )t o n a r l , v y a r ( o s t r i h ) . a oftn losly assoia
history of natur, a on(
BIBLIOGRAPHY in th ours of Buffon
L., and . Rlrv.1997'|listktitilt:Dioscluruoltirlnd
Dir-rgus, Although gology w;
.W..
th rigitt of tsirds.Nw York: . F-rmn. tur' it was not golog

to row siz. Nw wok on th
arlybirdsshows that dinosaurs
to th<lsof nrost nrmmals nd
ir.ds.suh s tl|'uiusornis.
sstags(juvniland suadult)
r rsult,irds fftivly am
lould lrvhd a grtr ratio of
rovd wing loadirrg would hav
nssaryfor sustaind flight?
: th growth dyrramis of arly
)n rOllstrutdfrom or-rtis-
:tdin th tissusnrks it diffi.
at dinsurs , nd hi vian
s.h Prsnof fathrs is
iztion of th loomotory mod-
th rrrswr not usd ir.rloo-
animl, nd th tail assistd in
tusulartail' mor lik a roo-
c ttl th spinsand prosssof
l of th thigh on nr th hip
ing th Strokagainst th stai-
dinosurslosstto birds, th
atrslfound a more important
.unningand in flight. Th for-
'ving a two-modul loomotory
)gthr' irds volvd thr-
;, th lgs,rrdtlr til ah pr-
lr grund. Phyllgntitrnds
atllsof th flight mhanism
v o I v d n d r f i n d f u n t i o n s
s (suhas prhing lws) and
otion.
L irds nd othr dinosaurs' th
th on to four ggs of today's
altriial,and thy would hav
)s;som parntal ar is likly.
l divrsifiation of th living
asd,stl that dult stags wr
l siz at intrvals r:rnging fom
).
'itil
: D iosurF'tlltttil
d Although gology Was ntral to Buffon,s notion of th history of na-
Buffo 455
Gauthir, J. A., and L. F. Gall' ds.2001. Nl Perspetiues'lth rigin nd rl
uolutil of Birds. Nw Havn, l.: Yl Univrsity Prss.
Padin, K., and L. . hiappe. 1'998a.Th lrigin and arly volution of birds.
Biologil Ruies (amridg) 73: 142'
|998. Th origin of irds and thir fltgt. Sitifi mri, Fruary:
3847.
Prum' R. o.,4nd A. tl. Brush.2003. Whih am first, th fathr or th ird?
ScintifiAmri,arh: 84-9. -K.P.
Llrd (1707_t788)
Buffon,Gorgs-Louis
..Thr is no
haptr of th thor of organi volution mor onfusd or
mo ontrovrtd thn that whih rlats to th position of Buffon'' (Lov-
io 1959,84). Lovjoy is orrt. No ightnth-nturysintistor philoso-
phr playd as prominnt a rol in th gnsis of volutionary idas as
Gorgs-Louis Llr d Buffon, ut then again, no othrs wr quit so
amiguous in thir opinion of volution. Thr is a rason for this: Buffon
wrot at a tim whn th vry ida of organi volution-whih, of ours, is
highly omplx-was not iarly dfind. In rtin rspts' Buffon was an
volutionist, in othrs h was not. From a historial prsptiv' two diffrnt
qustionsnd to b distinguishdaout Buffon and volution: (1) Was Buf-
.shat
fon an volutionist? (2) rol did Buffon play in th mrgn of th
ida of organi volution? All histoial data aout Buffon and volution
nd to valuatdin th light of ths two qustions.
Buffon's work touhd an inrprssivang of donrains.|n 174 h ntrd
th Paris Aadmy of Sinsas a mathmatiian.|n |739 h bam supr-
intndnt (had) of th Jardin du Roi (Th King's Gardn), on of th two
oldst sintifi instittions ratd y th Frnh monarhy; th othr was
th Collg d Fran. (During th l;rnh Rvolution' th Jardin du Roi was
rorganizdand rnmd th useum of Naturl istory.) As had of th
Jardin du Roi, Buffon dvotd mor and mor tim to natral history. In
174| he translatd Nwton's postlrumously pulishd thod oi Fluios
|176). H also produd svral original mmoirs on proaility and d.
mography. From ] 749 to th nd of his lif, his omplt works wr pu-
lishd in te 44 volums of his Hlsroire turelle, grI t prtiIir
[Natual history, gnral and partiularl, a sristhat h partly wrot in ol-
laoration with svral brilliant sintists of his tim, in partiular Daun-
ton and Gunau d Montliard (s figur). This giganti sum of work
dals in dtail with sujts as diffrnt as osmogny' minralogy, gology,
applid mathmatis, zoo|og, and anthropology. Buffon's viws on organi
volution (to us th modrn word, whih nnot found with its prsnt
snsin th ightnth-ntury writings) ar disprsd in many plas. Thy
ar oftn losly assoiatd with rfltions on th mor gnral thm of th
history of natur, a onpt that am to play an inrasingly important rol
in th ours of Buffon's work.
tur, it Was not gology that first ld him to th notion of a grrral and
4.\6 Buffon

tion of spis, in th
std in th history of
rspt. First, Buffon's
dsnt, with rsml
This notion of spi
trms of tmporal on
Buffon pliitly adm
b modifid y th at
For instn, h prop
thir ounterparts in ,
should onsidrd
Daunon's anattlm
L'As (175) that th
rats' and h said th
ommon anstry.
Ths idas had a m
on his thr points' h
an indfinit transmut.
to a rtain point. Bu
quadrupds (200) to m
nlargd spis.But hl
spis wr rintrpr
largd spisould n
osrvd at th right l.
nnt' as Natur hsl
sidrd th hypothsis
( l 7 5 3 ) ' w h r h p o v i
world as dsnding fr
of hand for two rason
anothr spisand (2)
mannt lin of distintr
this point. ost of th
fat that h progrssiv
only what w would
to th lvl of a ..famil1
Pitursof a rhinorosand an lphant,... *'r", s Nturl istor.The tion etwn spis an
-;
many forms of animls that w apprnt y th ightnthntury fasinatd |n Lpoqus d I t
th savantsand ld thm to spultaut th rasonsfor th divrsity. tory of th living worl
proposd that th art
omt' had progrssiv
alls mad of diffrnt
gradual volution (in th sns of an irrvrsil) history of natur (Rogr 100'000 yars had
1988). In Tbelr of the rth (1749), Buffon ws mainly intrestd in th from an initial stat of
a t u a l a u s s 'w h i h , y t h i r O n s t a n ta t i o n ' a o u n t f r a n u m r o f g - anothr stimation: 3 r
ologial fats (.g., th trnal ation of watr, whih aounts flr major puts forward an asolt
aspts of gographi rlif). It was only aftr h gan to laorat his no- formation of sdimnt

Buffon'sI'Jturl i stor. T he
th ightnthntury fasinatd
asonsfor th divesity.
:rsibl)history of natur (Rogr
fon was mainly intrstd in th
L o n ' o u n tf r n u m
lat, whih aounts for major
t r h s a nt o l r a t h i s n o -
Buffon 457
t i o n o f s p i s 'i n t h r i d - 1 7 5 0 s ,t h i r t B u f f o n g a n t o s r i o u s l y i n t r -
s t di n t h h i s t o r y o f n a t u r . T h r o f h i s i d s p l r y da m i o r r o l i r r t h i s
rspt.First' Butfon's onpt of spis was foundd on intrfrtility and
dsnt'with rsmlaning suordinatd to th ontinuity of linags.
This notion of spis suggstd a gnral vision of th unity of lif in
trms of tmporal ontinuity rathr th:rn in trms of trnal typs. Sond,
B u f f o n x p l i i t l y a d m i t t d t h t o t d o m s t i a t da n d w i l d s p e i e s o u l d
m o d i f i d y t h b a t i o n o f s v r l a u s s( . g . ,l r u r r r na t i o n o r l i m a t ) .
For instan, h p'roposd that th nimal spis of th old World had
thir ounterprts in Amria: th lattr dsndd from th formr, ut
should onsidrd today as distint spis.Third, on th asis of
Daunton's anaton]ial work on nimal skltons, Buffon argud in
L ' s e ( | 7 5 3 ) t l r a t t h r W a s a n r . r n d r l y i n gr - r r r i t 1 , otfy p o f r h v r t _
r2rts'and h said that this uniry of ryp suggstd th hypothsis of a
ommon anstry.
Tlrsidas had a massivimpat on ightnth-nturynatural historians.
on his thr points, howvr, Buffon did not rally adopt th hypothsis of
n indfinit transmutation of spis. Spies uld modifid' ut only
to rtain poirrt. Buffon proposd to rdu tlr numr ot spis of
qr"r:rdrupds (200) to mrly 32, intrprting th urrnt familis in trms of
nlirrgdspis.But h did not Want to go any furthr. ost of th urrnt
spieswr rintrprtdin trms of varitis' ut t limits of th nw n-
largd spisould not ovrstppd.Providd tat th naturl spisis
obsrvdat tlr riglrt lvl, thy ilr .,pduringniris,as anint, as pma_
nnt' as Natur hrslf'' (Scod Vue' 1765). Similrly, whil Butfon on.
sidrdth hypothsisof transmutation in his famous mmoir on th donky
(175),whr h providd an inrdily suggstivviw of th whol animal
wrld as desendingfom a ommon anstor' h rjtdthis hypthsisout
of hnd for two rasolls:(1)no true spisis known to hirv ppdfom
anothrspis:rnd (2) th infrtilitv lf hyrids provids dfinit nd pr-
manntlin of distirrtiontwnspis.Buffon nvr hangd his mind on
thls point. ost of th allgd Ontraditionsof his thinking rsult from th
fat that h progrssivlydoptd llotion of th spisthat inludd not
only what w would all today a ..spis''ut also vy possil taon r'rp
to th lvl of a ..ferrnily.'' But Br'rfforralways nraintaind an soltrtdistin-
tion twnspisnd varitis.
InEpoqus d l tur (7779), Buffon intgratd his idas aout th his-
tory of th living world into a gnral vision of th history of natur. H
proposd that th arth' aftr hving n gnrtd y a ollision with a
ot, hd progrssivly eom oolr' on th asis of xpimnts on
allsmad of diffrnt matrials (spiallyiron), lr alulatd tht at last
100,000 yars had n nssary frlr th arth to ool to its prsnt stat
from an initia1 stat of fusion. (In anothr manusript, h somtims givs
anothrstimation: 3 million yars; s Rog l988.) In poqus, h also
r o f g -
puts forward an asolrrtgologil hronology on tl] asis of th stady
formation of sdirnntary roks. p<tqusgav a massiv irpuls tl th
458 Buffo

no dsntwith modi
a lly pr-evolutionist
a aa
aa . . .. Buffon's volutioni
aa a :ltion of spis witl
aa
a This is prisly w
rjtd,so many hist
tionist in th modrn
insight into Buffon's r
.tf.' a
aa a ta aa'
. .. BIBLIOGRAPH
a a
a
Br'rffon,G.-L. (Llrd
vols.Paris:Imprim
mntindin this
a
.... prtiuIir.|
a
Gyon,J., d' 1'992.Bu1
.. .. t. .... ... ...... .....3 .. .
.. d l mrt d Bffo
a G. anguilhm.Pr
I Lvjoy,A. o' 1959.Bu
I
HA|N nd W. L. Strauss J;
1749 1755 1761 17661767 1780 1782 I 789 Johnsopkins Pr
Rg'J. 1988.Intrldu
harls Darwin was far from th first to us th pitrrrof th tr of lif as ditionsdu trsL
rprsntation r mtphorfor th histoyf lif on erth.On rlirusr was Shnlitt.S. 2007.Bttff
Buffon. This diagm pl<ltsth frqunieswith whih he usd th tr mta-
phor and othr metaPhors'maps' and hins.Not that th us of th tr mta-
phor in fat dlind with tim' possily a funtion of th fat that Buffon am Burgss Shal
undr rligious prssurfor his toying with evlutionary spulations'and, as
Th fossil rOrd larg
on always ken t sty on th side of th authoritis,h modifid (or at least
onaled)his thinking aordingly. (Adaptd from G. Barsanti, ..Buffon et Alnrost invarialy, ll
l'imagd la natur,''in Gyon |992,290.I shlls) as wll as tral
ganisms imprintd on
S u h v i d n sa r s t i
idn also has its dis
volutionary viw of natur. This mmoir also put forward a hypothsis prsrval hard prt
aout th origin of th diffrnt animal and plant spis.Lif had appard fossils ar undiagnst
as a onsqun of th spontanous assoiation of organi moluls, tial tras an bui
whih thmslvs had rsultd from natural hmial rations whil th All, howvr, is not
trrstrialglo was ooling. Diffrnt assoiationsof diffrnt organi mol- wirrdow into th dist
uls had gnrtd th diffrnt living spis. From th onst' ths pisolslf fossiIizati
spis Wr as ompl as thy a now. As a funtion of th physial ir- parllld irrsightsint
umstans (i.',limat), spisapprd sussivly'ut thy did not d_ th potntial to rsolv
snd from ah othr. All wr spontanously gnratd. Buffo also tirrv dposits yilding
argud that th various spis migratd from on pla to anothr as a irnportant is th Burg
funtion of limati hang, and tht thy am tint whn th irum- notaly thos from so
stanswr no longr favoral. Thr ar svral r
This final spulation is haratristi of Buffon. On th one hand, his on- of intrst.In partiu
ptions aout th antiquity of th arth, its gadul hang,th influnof rian xplosiln,th st
physial onditins upon th history of lif, and xtintion look gnuinly of tlr arrirrr[s. Th x
volutionist.But his onptual shm for th arupt origin of sPis,With s k l t a l f o s s i l s n d t h

'1780 1787
hpiturof th tr of life s a
lif on arth.On earlir usr was
ith whih h usd th tr mta-
. Not that th us of th tr mta-
ntion of th fat that Buffon am
volutionaryspulations,and, as
thoritis,h modifid (or at last
..Buffon t
d from G. Barsanti,
ir lso put forward a hypothsis
d plant spis.Lif had appard
Lssoiationof organi moluls,
ural hmial rations whil th
oiatins of diffrnt organi mol-
g spis.From th onst' ths
As a funtion of th physial ir-
d sussivly'ut thy did not d-
tanously gnratd. Buffon also
l from on pla to nothr as a
l am xtint whn th irum-
I Buffon. on th on hand, his on-
its gradual hang, th influn of
[if, and xtintion look gnuinly
lr th arupt origin of spis,with
Burgss Shl 459
no dsntwith modifiation and no room for inrasing omplity, is typi-
ally pr-volunist.
Buffo volutionism was a limitd volutionism. H aptdth modifi.
ation of spiswithin narrow limits, th limits of what h alld spis.
This is prisly what hrls Darwin nd nintnth-nturyvolutionists
rjtd,so many historians lgitimatlyssrtthat Buffon was not an volu.
tionist in th modrn snsof th trm. (Sth figure ln pag 458 for mor
insightinto Buffon's vaillations aout vllutin.)
BIBLIOGRAPHY
Buffon,G.-L. ([,lrde).1749-1804.Histir turll,grlt prtilir.44
vols.Paris:ImprimriRoyl;Plassn.INot:All writingsy Buffon
mntiondin this ntry wr pulishdas part of Histoir turll,grlt
prtiulir.l
Gayon,.|.,d.|992. Buffo 1988,tesdu ctllloqitrtiol du bietir
d l mort d l3uffo,Pris-otbrd-Dijo.rfy . yr, postfay
G. nguilhm.Pis:Vrin.
Lovjoy,A. o. 19.'9.Buffonnd th prolmof spis. In B. Glass,o. Tmkin'
and \W.L. StraussJr., ds.,Forrurs
of Dri: 1745-18.'9'Baltimor:
.l
1789 JohnsHokins Prss,84_] .
Rogr,J. 1988.Introdution. In Buffo:Ls poqusd l tur,i_lii.Pris:
ditions du Musum national d'istoir naturll.
Shmitt'S. 2007' I]uf|o-(ut,rs. Paris: Gllimrd. -1.G.
BurgssShal
Th fossil rord largly onsists of th rul and srapings of anint |if.
Aln-rostinvarialy, all that w find r roust pis of sklton (ons and
shlls)as wll as traks, trails, and othr signs of ativity th on living or-
ganisms imprintd on th sdimnt nd ar now prsrvd as tra fossils.
Suh vidnsar still ntral pillrs in doumnting volution, ut suh v-
idnalso has its disadvantags.F.irst,many organisms, vn animals, lk
prsrvalhard parts. For ampl, onsidr th slug. Sond' many tra
fossilsar undiagnosti as tl th original typ of animal, and fftivlyidn-
tial tras an uilt y unrlatd animals.
All, howvr, is not lost. This is auson oasion ntur throws opn a
window into th distant past. Thls is possil on aount of ptional
pisodsof fossilizatiln.Th rsultantsoft.odid fossils not oniy giv us un.
parallldinsights into th original divrsity of anint lif' but thy also hav
th potntial to rsolv fundamntal prolms in volution' of th sdimn-
tary dposits yilding xptionally prsrvd animals, argualy th most
importantis th BurgssShl (British olumia, anada) and its ongnrs'
notalythos from sluthwsthin and North Grnland.
Thr ar svral rasons why Burgss Shal_typfaunas ar suh a fous
of intrst.In partiular, thy throw rmarkal light on th sl-alldam-
brianplosion' th sminglyarupt apprnand volutionary adiation
of th animals. Th plosion is slf-vidntfrom th dramati apparanof
skltlfossils arrd th divrsifiati<lnof tr fossils, yt it is now apparnt
460 Burgss Sbl

that if w hd to rly on thos sours of infomurtionalon, ur un.{rstand-

ing of this rmarkale vt would gatly impovrishd.This is us
th soft-odid rihs of th Burgss Shl-ty1.lfaunas rval tlr traordi-
nary divrsity of amrian marin lif and in doing so pos qustions of ma-
jor volutionary intrst.\/ht.if anything, triggrd this plosion of lif?
D w nd to onsidr novl mhanismsof vtllr.rtion? lWhat was th rol of
ology, spially prdation? All ths qstionsar ons of mjor urrnt
dat.
Th typ mplar of suh funas, th Burgss Shal itslf, is from a loal-
ity nar Fild' British olumia, ut numrous Othr loalitis are knwn in
th wstrn Unitd Stats as wll as astrn lolitis in Pnnsylvnia and
Vrmont. Futhr afild, ky lolitis inlud Parv Land, Gnland (Sirius
sstfun)' southrn lrin (hngjiarrg,Kili), and South Australi (Kan-
garoo Islnd)' Dspit th rng of ags (approximatly 530 to 510 million
yars) and also nvironmntal sttingsthat varid sustantially in trms of
watr dpth and proaly oygn ontnt' th Burgss Shl-typ faunas
show onsidl homognitv. Thus thy are dlminatd y zrrthropds,
altlrough th wll.known tiloits form n irrsignifiant fation. Also
o m m o n r s p o n g sa n d v a r i o u s W o r m s ( s p i a l l yt h p r i i r p u l a n s ) ,w h i l
Th animal Hlluigl
rarr groups inlud the hodats and vn th dliat sa-goosrris
(tnophors).
Thr ar svalrasons why Burgss Shal_tl,pfaunas ar of partiular
volutionary inrportn. First, tlry ar wll known fo housing :r sris of
supposdly izrr anirals, inluding th aptly namd Hllcigni (sfig- ut th rmirrdr h
urs). In fat, th history of tlr study of this p:rrtiularanimal mplifisa ooks.
radial shift in sintifi thinking. Th fundamntal prolm hs n that Th quisit prs
not only do som of ths fossils look vry strang' but fitting thm into any traordinary insigh
shm of volr.rtionappard prolmati. Th diffiultis' howvr' Wr tors transpir to b r
largly a rfletior-r of inomplt information and somtimsa lak of img- of thir ativitisorr
ination. Rsolution of ths polms has dpndd on svral lins of in- mks. So too it s
quiry. In partiular' molulr iology has tratlsformd th undrstanding tlr arupt apparan
.Whn
of many aspts of animal volution. nw phylognis ar omind pr:ottiln.A partiu
with rintrprtations and frsh disovris frm th Burgss Shal_typ inirl sklton form]
funas, it orrrsapprnt thilt ths supplsdly izarr rrrrin ani- fliility.
mals ar sinrply staging posts in th volution of major groups' usually r- Th story of th B
frrd to as pyla. Thse fossils show us, thrfo, how oly plans wr dd, muh rminst
assmld and, just as importnt, th funtionirl and llogia1ontts of disovris, notaly
ths ky stags in th mrgnof nimals. It is qually important to ap- finds wr th lrl
priat, howvr, that th asi prosss of volution involvd in th nrrgd. So too, th
arnrian plosiorr appar to no diffrnt fror thos Ori1tiI1g eny- Cilrrrrin plosion
whr ls. Again and again th study of vllution appars to psnt un. triggr, may t th
ridgal gulfs twn wildly disparat forms. A mmonly posd ] n' or altrnativlyt
qustion is: ..ow on arth col|d tbt hav volvd? \/hatstps ould possi- look to multipllur
bly onnt ths organisms?'' Yt' rpatdlyth flssilrord r,alsa s- hav volvd from s
ris of intrmdiar,vstags that utifully display th transition of form. this trnsition r sp(
Almost invarily tsmissing links displ unptdan]rrovlfatus, triguing dira
 BrgssSbl 461
)rmationalln,our undrstand-
ly impovrishd.his is us
:ypfaunas rvl th traordi-
l doing so os qustions of ma-
triggrdthis xplosion of lif?
.s7hat
:volution? was th rol of
stionsar ons of maior urrnt

rgssShalitslf, is from a loal.

ous othr loalitis ar known in
n loalitisin Pennsylvania and
J Pay Larrd, Gnland (Sirius
Kaili), arrdSor"rth Australia (Kn-
proimtly530 t 510 nrillion
: varid sustantiallyin trms of
, th Bugss Shal_typ faunas
y r dominatd thropods'
n insignifiant fration. Also
. s p i a l l tyh p r i a p u l a n s ) ,w h i l
h nim1lluigispdrs,an inhitntof th
/n th dliat sa-goosrris BuressShl.

.ral_tvp faunas ar of partirrlar

li known for housing a sris {
lptly namd Hlluigi (s fig- ut th rmirrdr is tht .ratur is ustrally
a ttr tahr than tt-
Lispartiulaanirnal rnplifis a books.
ldarnntalprolm has n that h xquisit prsrvation o th Burgss Shal_typ
faunas also rvals
s t r a n g h, u t f i r i gt h m i n t o a n y traordinarvinsights into th ology of arly niml
ommunitis. i)rd-
. Th diffiultis' howvr, w tors transpir to b mor signifiantthirn on thought,
with dirt vidn
ion nd somtimsa lak of imag- of thir ativitislming from gut ontnts' f,oio-looking jaws,
and it
; dpnddon svral lins of in- mrks.So too it sems tht on of th hllmrks
of tt' "mi,n -piosion,
rs transforrtdtlr undrstanding th abrupt apparanof hrd shlly parts' is lrgly
a rsult of th nd for
nW phylognis ar omind prottion.A partiularly striking aturof som
hard parts is that th oig-
:is from th Burgess Shal_typ inal skltonformd a sort of naturl hain mail that
omind stnethwith
upposdly izarr arnrian ani. flxiility.
r t i o no f m a j o r g r o u p s , u s u a l l y r - Th stoy of th Burgss Shal_typfuns is y
no mans omlt. In-
, therfor'lrow ody plans w dd,muh rmins to olplishd.First, tlrris
stdy stram of nw
tionl and ologil Ontts of disovris,rrotaly frot hina. Prhaps th
most i..'p,.ta.,t of rnt
als. It is qually important to ap. finds wr th rlist-fislr, th group frlm
whih h.,,.'".', ultimatly
ss of volution involvd in th mrgd.So too, ths faunas rinfor our sns
of th ragnitud of th
f n tf r o m t h o s < t p r a t i n ga n y - mrian plsion' It is far from lr whthr w
nlto ,..k a uniqu
volution ppars to prsnt un. trlggr'may at th gnomi lvl or praps in
trms tlf tmosphri oxy-
at frms. A ommonly posd gn'or altrnativlyto dismiss a singl plntion
.sht as sintifillynaiv and
: voir,d? stps ould possi- look to multipl usations.Finally, what of th
ntdnts?Arrirals must
dly th fossil rord rvals s- havvlvd frll slmtlringls, ut tlr natur
and gologil timing of
Lv display th trnsition of form. thistrarrsitionar spulativ.Th amrian plosion
is prddy th in-
ay unxptdand novl faturs, triguingdiaaran assmlgs,ut thir ()nntlon
to th story of animal
462 Brgss Shl

voltion in th anrr
Shal_type faun:rsnray
thy rmin a ftil fil

ILloGRAP|-tY
Budd' G. .' and S. Jns
iltrianphy[a.8ill
onway orris, S. l998.
of Aimls.amrid
Llotr,..G., R. J. Aldridg
204. Th mbri
iml Lif. for1
Valnrin'J. W. 2004.
Prss.
Zhuravlv,A. Y., nd R. t
Rditio, Nw Ylk

Bush,Guy (b. 192

Guy Bush Was dut
th Univrsity of Tas (
Lansing. At arvard, B
rnst ayr, had of th
is allopiltri' that is, dt
lngd this viw liv
Spcis, that it is possil
graphi isolation) to tl
nor'rghto hav som ktl
vids y spializingon
[ p . H h a s p u s u dt h i s
Bush and his tlw
pomonell, known as th
worm. on onfind t
sprad widly to ultiva
ing hits of th flis rrr
Ronstruting lluigni sprs. Tlr uppr drwing shor,vsth first ttrrrpt
Bus hs n l to sl
at ronstruting fl' sprs.lt is lpitd as walking rln its spins. Th lowr
rpodutivly isolatd
drawing shows visd ronstrlltiOn, with th spills n()W upprmost'
prhaps ing usd for dfns. orr-rparthis with dward Drinkr op's thi rsptivhosts.
ronstrution of th plsilsaur, lsmosurus plturus (s figur on p] It turns out that odrlr
487). !hras op got his animal k ttl front' lnr,t,yMrris gt his th sptivodors of t
upsid down. This zr[[gos to shlw tht ronstruting th ast r1uis on hlst th is rpid r
ilnginatiln and thrly s mulr lrs llrut fossi] fin]s(s lso th Ollstrutd of this host and ntlt of
d i n o s u r s s h o w n i n F i g u r 1 8 o i t h m a i n s s i r y . . T h H i s t o r y f v t l l r r t i t l n r y availl and othr flis
Thought'' y ilrl Rus in this vllum). Again' pttion nd imginaritln
mthods and findings ol
triumph ovr fat. Popl thought o th dinlsurs s rutish and umbrsom,
..s la gns that ar involr
so thy rinford tlris pjudi witlr thir modls anl xprssions lik
outd:rtd s a dinOsur.'' Howvr' tl-rdinlsaus thrivl ior L.50milliln vrs, and wlrl-tspiation h
rrrdthir anstol.sr still flvic Out vrv'uvh. Te work of Bush a
 Busb 46

volution in th amrian is still largly unrsolvd. In onlusion, Burgss

Shal_typfaunas may a ky omponnt in undrstanding volution' ut
thy rmain frtil fild for all sintists, and spially th young.

|BLloGRAPHY
Budd,G. ., and S. Jnsn.2000.A ritialrappraisal of th fssilrordof th
ilarrianp|a.Bkllogil Ruitus75: 25-296.
onwy Morris, s. 1998. Th ruibl lf rtio:Th BrgssShlnd th Ris
of imls. amridg:Can.rbridg UnivrsityPrss.
Hu, '-G., R. J. ldridg,J. Brgstrm, D. J. Sivtr,D. J. Sivtr,and ..H. Fng.
2004, Th mbri Fossilsf bgjig,hi: Th Floeriglf rl
Aiml Lif. Oford: Blakwil.
Valntin,J. !. 2004. o th rigi of Phla.hiago:Univrsityof hiago
Prss.
Zhuravlv,A. Y., and R. Riding,ds.2001. Th llogof th mbri
Rditio.Nw York: olumiUnivrsitvPrss. -s...

Bush,Guy (b. 19291

Guy Bush Was duatd at Harvard Univrsity and spnt his working lif at
th University of Tas (Austin) nd thn at ihigan Stat Univrsity at ast
Lansing. At Harvard, Bush's intrsts wr sparkd y th strong laims of
rnst ayr, hd of th Musum of omparattve Zoo|og, that all spiation
is allpatri, that is' dmanding gographi isolation to ou. Bush hal-
lngdthis viw living, as did harls Darwin in his oz th rigi of
Speies,that it is possil for sympatri spiation (spiationwithout go-
graphi isolation) to our. Following Darwirr, Bush thought that it ould
noughto hav som kind of ologial isoltin, whr prhps a spisdi.
vidsy spializingon diffrntfoodstuffsvn though thr is physial ovr-
lap. H has pursud this inquiry, with onsidralsuss,for ovr 4 ears.
Bush and his owokrs and studnts hav fousd on Rhgoletis
pomoll,known as th appl maggot fly ut somtimsalld th rilroad
worm. On onfind to th hawthon ush' in the past 100 yars o so it has
spradwidly to ultivatd appl trs introdud from urop. By amin-
ing haits of th flis in th wild' as well s through ontolld xprimnts,
drawing shows th first attmpt Bush has n l to show that th flis ar reaking into sparatspis-
lkingon its spins.Th lowr rpodutivlyisolatd groups-simply y dvloping adaptations to utiliz
hspinsnow upprmOst'
thir rsptiv lrosts.
with dward Drinkr op's
It turns out that rdor is major fator t play. Th flis a rsponsivto
s pltrus(sfigur on ag
nt'onway Morris got his th rsptivodors of th fruits f thir hosts. On a population sttlson
strutingth past rquirs on host thr is rapid sltion prfting th aility to rogniz th fruits
l 6nds (salslth ronstrutd of this host and not of othrs.his an our vn whn thr ar othr hosts
r y ' . T h H i s t o r yo f v t l l u t i o n l r y availaland othr flis utilizing thm. Rntly, Bush hs n utilizing th
r i n . p t t i t ,ann d i m g , i n e t i l l n mthodsand findings of molular iology' oth to trk down th partiu-
jaursas rutishand umbrsom,
.,as lar gnstht ar involvd and to us this knowldg to map prisly how
ldlsand prssionslik
and whn spiation has ourrd.
urs thrivdfor 150 million yars,
Th work of Bush and his ssoiatsis an xllnt ampl of a ph-
rvhr.
464 Bush

nomnon that ritis of natural sltion oftn dmand for Darwinian

thory an takn as provn' namly, an osrvd group atually brak-
ing into two diffrnt spis. It would ni if th work of Bush ndd
this partiular ritiism forvr; howvr, sin so oftn th ritis ar mo-
tivatd mor y trasintifi prjudis of a rligious natrrr rathr than
y th fats' on douts that this will th final word on th topi.

BILIoGRAPHY
Darwin, . 1859. ()n th, rigi of Spies.London:John urray.
Howard, D. J., nd S. . Brlhr,ds. 1998.dlssForms:Spisd
Spitio.Nw York; Oford UnivrsityPrss. ain, Arthur Jams
Mayr' . 1942.Sstemticsd th rigi of Spcis,Nw York: olumbia
UnivrsityPrss. -^4.R. A. J. ain (h always us
of th group of British .
Ford's oIogil gni
.!Var
tr sorld II. Thy f
as a major ( t h yw o u l d
duatd at Oford f
sltion ws rgardd'n
gists, lik Linar Profs
on volution, and whn
drift wre muh favord
ti) rasons for distrustr
lin of thinking riv
| 9 4 6 o f h s p i r i r u l i st
Profssor of Zoology (;
Togthr with fllow
light of thory ing dv
ampl of Ford, ain st t
of ntur. Thir most
emorlis, widly found
trysid round oxford. l
ing and with plain shlls
takn to a pardigma
fturs Wr thought fal
ain nd Shppard sh
th akgounds in whi
xampl, is diffntfron
from th grass of a fild 1
a major lmnt in th dj
to th akgrounds th l
portant tool in making t
nvils (stons)on whih
ould ollt nd ount t
d i f f n tl l i r i s( i n
ml Speis d hir ]
sujt.
ln dmand for Darwinian
bsrvdgroup atually rak-
i if th work of Bush ndd
Lso oftn th ritis r mo-
a rligious natur rathr than
final word on th topi.

n:JhnMurry.
llessForms:Specisd
ain, Arthur Jams (92|_|999)
ies.Nw York: olumia
-^4.R. A. J. ain (h always usd only initials for profssionalpuliations) Was on
of th group of British volutionary iologists who wr mmrs of . B.
Ford's ologial gntisgroup at Oford Univrsity in th arly dadsaf-
tr NorldWar II. Thy firmly stalishdth signifianof ntural sltion
as a major (thy would hav said th major| aus of volutionry hang.
duatd at Oford for th wr, ain had ntrd a world whr natural
sltionwas rgardd with suspiion, if not outright hostility. Most iolo-
gists' lik Linar Prlfssorf Zoology . S. Goodrih, did not ltur at all
on voluti<ln,and whn th topi Was mntiond altrnativssuh as gnti
drift wr muh favord. Not a fw hd fairly ovrt ligious (at last vitalis-
ti) rasons for distustinga purly naturalisti approah to origins, and this
lin of thinking rivd almost offiial support with th appointmnt in
\946 of th spiritualist Alistr Hardy as sussorto Goldrih as th Linar
Profssorof Zollogy (ain and Provin 1991; Rus 1996).
Togthr with fllow Oonin iologist Philip Shppard, wrking in th
light of thory ing dvelpdy Ronald A. Fishr and th xprimntal -
mpl of Ford, ain st out to show that natural sltionis a powrful for
of natur. Thir most lratdand lasting work was on th snat| p
morlis,widly found in th woods, madlws,and hdgrowsof th oun-
trysid around oford. Th snail oms in many diffrnt ftrrms,with and-
ing and with plain shlls, and with pink nd rown and othr olors. It ws
takn to a paradigmti ampl of gnti drift; suh simpl and vrying
fatursWr thought far too insignifiantto hav any tru adaptiv valu.
ain and Shppard showd that in fat th shll ollrs ar tightly tid to
th akgrounds in whih th snails a found. A hw.lodforst floo, for
ampl, is diffrntfrom a dappld dith sida hdg,and again diffnt
frm th grass of a fild grazd y ttl. Thy disovrdthat th snails ar
a major lmnt in th dits of irds' thrushs spially' and that aording
to th akgrounds th snails ar apturd and atn diffrntially. An in-r.
portant tool in mking ths dtrminationswas th fat that thrushs us
anvils (stons)on whih to smash th snail shlls, and hn th rsarhrs
ould ollt and ount th rlativ numrs of forms takn y th irds in
diffrnt loalitis (ain 1954; in nd Shppd 1950, 1954). ain's zl-
ml Speis d Thir uolution (1954) m a standard work on th
suit.
165
466 i

Lving Oxford, ain vntually urrrr profssor at Livrpool Univrsitv

arsn, Hamptn
(whr Shppard had rnovd),a major entr of Britlsh volr.rtionarystudis,
havily susidizd y th Nuffild Foundation in an attmpt to show th sig- Th rsarh of amp
nifin of volutionary studis flr mdial issus,suh as th natural vari- most important studis
bility of lood type and tlr onsquntsusptiilitto varior"rsailmnts.I}y lutionary iologv of th
this tim ain had lost intrst in frontlin rnpiial rsarlr,although h hs nitd tnsiv
did ontinu to work with ollaorators and studnts (in and urry multidisiplinilry rsa
796a, 196)' Inrasingly,h turnd to th history and philsophy of his pattrns of th giant p
disiplin. H wrot loquntly and informativly or-rsuh issus as th n- arson dvlopd a po
tur of spis and th prolms f Linnan tonomy' pirtiularly aout formulatiorr of nw
its volutionar undrpinnings (ain |959a,1959,1962). Although his ma- spis' rson's F.oun
jor rativ work am bfor th major onptual vnts of th 1970s (s- in bringing togthr a tl
pilly th orr-ringof ladism), in did muh to rais th importan of work togthr in unravt
taxonomy ;rs a sin in its own right and on Worthy of thortial study' oigin of nw organism
(ain had spnt a yar arly in th 19.50sworking in Amria with ornithol- systmatis and taxono
ogist and systmatistrnst ayr. In itslf this shows how by mid-ntury gology, iohmistry,
Bitish and Arrian volutionistsWr snsitivto tlr mrits of intlltul prosss within on
oss-frtilization,and spifially in ain's as this was irportant for his Hawaiian Arhiplago.
own thinking about taxonomy.) aras, ithr [.lhis ow
Susquntyars hav shown, prhps ptdly,that th work of in othrs. H has movd
and Shpprd rrds rvision in light of other sltion-ontrolldfators proahs aros, and h
(Jonset a|' 1977). But thir work still stands s landmark in th history of lgus with nw and a
n"rpirialstudis in Darwinian volutionary thory. flxiility of his intll
mrltidisiplinary ppr
B|LIoGRAPHY
goup of insts.
rin,A. J. 1954.iml Spisd Thiruoltirl.L,lndon: utl-rinson. Hmpto arson w
1'959a.Ddutivrrdindutivmthodsin plst.l,innaan tonomy. sylvania. H otaind ]
Proedigsof th Li Soitof Lodrl 17():185-217. dgr (PhD, 1943) fro
1959.Th plst-Linrraan dvlmntlf ttlomy.Prl,!igs of th
assistant profssor of l
LiSlit of Lotltlll170 234-244.
1 ' 9 6 2 . T v o l u t i no f t a o n o m i p r i n i p l sI.n G . . A i n s w o r t ha n d souri, in 1943. arson;
P. . A. Sath,ds.,irobil lssifitio, 1_|3. amidg: ambridg p r o f s s o r .I n | 9 7 1 ' , e
UnivrsityPrss. th Univrsity of Hawa
A. J., and J. D. Curry. l963a. Ara fftsin p{lctl,
air-r, PhilosophiI rson pulislrdn
rnstiltts oi th R-l1'I Soitl'B 38l 269_299.
iology and pulishdl
196.Ara fftsi pon th Lrkhill ArtillryRangs,Slisur
P|ain'loull lf th LineSoietof l-odo Zooklg 4.5:1-1 5. itl, d Humn Lif (I\
in, A. J., and.!7.B. Provin.t991. Gnsnd login histry.In R. J. Brry, as on of th rnost inflr
ed.,Gsi oklg: h rd Smptsium lf th British llogilSoit. svrl dds. In 1!
Oford: Blkwll. :rwddrsln th L
in, A. J.' ad P. -N4. Shpprd.1950.S|tion in th;lolymorphiland snil
ual sins'Partiula
pmorlis(L.|. rdit4z 27 5-294.
ial Drosophlla. !hil
1954.Ntulsltiln in p.Gtis39: 89_1-l6.
S., B. Fl. Lith, and P. Rawlings. 1977.Polymrplrism in 1l: t th:rn four dls,hrs
Jns'J.
problmwith too ransolutions?ltul Ruitt, of ologLldSstmtis v a r s < l rhi s s t u d i
8:109-14.1. on mdiinal hs us
Rtts,\4. 1996.od to : Th tlcptof Progrssi uolutiotlrBiolog. lishd a papr on th r
mridg, MA: arvrd Univrsity Prss.
lt frtiliztiorrin a s1

profssorat Livrpool Univrsity
:r of British vrlutionarystudis'
on in n attlpt to show th sig-
l issus,sr.lhas th naturl vri-
ptiilityto various ailmnts. By
: mpirial rsarh,although h
and studrrts(ain and urry
:hhistory nd philosophy of his
rativlon suh issus as th na.
an taxonomy' partiular[y aout
, 1 ' 9 5 9 'l 9 6 2 ) . A l t h o u g h h i s m a -
)nptualvntsof th 1970s (s-
muh to ris th importan of
l on worth'- of thortial study.
,,orkingin Anria with ornithol.
I this shows horv y mid-ntury
:rsitivtl tlr mrits of intlltual
's
as this was important for his
ptdl,that th work of ain
oth sltion-ontrolldfators
rdsas a lndrnark in th history of
y tlror,v.
]utio.London:uthinson.
ds in plst-Linaan taxonomy.
d o | 7 : 1 8 5 _ 2 1 7 .
tt lf taonlt.lrr,. Prodigs
+.
nipls.Irr G. . Ainswortl-r and
rio, 1_13. amridg: amrilg
:s in p. Philosophicl
59-299.
-arkhill Artillr Rangs, Salisury
. , , n d t , nZ t t ' h t g r 4' . s : l - | 5 .
'd ologyi histry. In R. J. Brry,
nl of th British oklgiI Slit.
ln in th polymorphi land snail
94.
ltis39 89_116.
7.Polymophismi p:
l Rt,itutli log d Sstemtis
f Progrssi uoltir Billlg.
rso 467
arson,Hampton Lawrn (|9t4_2004)
T rsarh of rpton Lawrn Cirrstltr has n itd as on of the
most import:rrrtstdis of twntitlr-nturyiologv. His wok on th vo-
lutionary iologv of th inst family Drsophilid of th awaiian Islands
has n itd tnsivlyand is onsidrd on of th suprm ampls of
multidisiplinary rsarh. Basd on dtaild mintion of th anding
pattrns of th giant pllytn hronrosoms in th Hawaiian Drosophil,
arson dvlopd a plrvrfulmthOd of ppulation studis that ld to th
tlrrnulation of nw rnhnism ftlr plaining th formtion of nw
spis,arson's Foundr-Flush Thlry of Spition. H was instrumntl
in ringing togtlrra tam of prts in all asptsof iologial sinsto
work togthr in unravling ..that mystry of mystris'' (Darwin 1859)' th
origin of nw orgalrisms on this arth' arson and his ollagus hav usd
systmatisand taonlmy, gntis' ology, havior, plrysiology, otan,
gologv,iohmisty, and m<llulariologv to undrsttrdtlr spiation
prossswitl-rin on of th mlst rnrarkl grotlps of orgnisms in th
Hwiian Arhiplago. H has mad mjor ontriutions in ah of ths
aras,ithr y his own work or y ringing to light th rlvant work of
othrs.H has mrvd from on a tO anothr as th nd for nw ap-
proahs aros, and h gnrously nouragd th patiipation of ol-
laguswith nw nd appropriat skills. Th radth of his r'rriosityand th
flibility of his irrtIlt,in e sns' rprsnt almost an pothosis of th
multidisiplinary approah to invstigating th voltltionary history of this
group of insts.
Hampton arson was orn on Novmr 5,1914, in Phildlphia, Pnn-
sylvania. otaind oth an undrgduat(AB' 196) and postgradut
dgr(PhD' 1943) fom th Univrsit.vof Pnnsylvani.H was hird as an
of th
assistantprofsso of iology at washington Univsity in St. Louis, is-
souri,in 1943. :rrson rmaind on th faulty as assoizltptlfssorand full
profssor.|n I97|, h movd to Hlvaii to om prrfssorof gntis.-t
thUnivrsity of Hawaii's John A. Br'rrnsShool of Mdiin.
rson publishd narly 300 sintifi artils in th fild of volutionary
iology and pulishd a dfinitiv ook in th fild f gntis ntitld Hered-
ity,dHumn l-ii ]96). His paprs r ited widly' arld is rognizd
as on trf th rrrost irrfluirtil figurs irr volutionary iology during th past
svraldads. In 1985' th Plrilllphia Adrrrv of Natual Sirrs
awarddarson th Lidy Mdal for lris outstandingontributions to th nat-
ural sins,partiularly his work on th volutilnary illogy of th Hawai-
ian Drosolhila. Whil Carson's wlrk r>nth awaiin insts spannd mor
than four dds,lris ontriutions ttl tl-rnatural sinsgn narly 30
vrsflrhis stLldisirr th Hwiian Islands.In 1935, h ptllishda papr
on mdiinirlhrlls usd y th Lirrado skimls.Thn, in 1940, h pub-
lishda papr Ol-lth rd rossill, and in 1945 h ublishd a papr on th
-.R.
latfrtiliztionin a spisof snak. His intrstsin iology had oviously
468 stl

n strongly irrflr.rnd y an intrst in naturlll history that gn as a of gntis.Lik mny

young oy ut at-t-t fully dvl<lpdas a vrungollg studnt t-r-rnlr of ing to prolnrs of h
th Dlawar Vlly Ornitholt>giallu. ginr-ringof th twntith
Although son rtird frlm offiial dutis at th Univrsity of Hawaii's h was also th first to u
Dpartmnt of Gntis in 1985, h was still vry ativ in rsarhand on- th volutionist's wrkh
tinud to mak signifiant ontriutions to vlllttionry iolog. H is won th Nol Priz for
onsidrd to hv n on of tlr Urrivsityof Hawaii's most illnt si- astl's most famotls
ntists. was a mastr t:rhrwhtt, in th ours of his distinguishd- nd whit oat olos (
rr' gav of l.rimslfunslfishly t>his ollagusand to his malrv studnts. primnts that showd
Modsty is a virtu unparalleld in this grat sintist.On' whn h was populatin' from mix
'W'hat
ing intrviwd y th nws mdia during a syrnposium hld in honor of Phillips 7914). w:
his rtirmnt,h ommntd tht..it's vry humbling. Sinis a slialn- tion was finishd, th r
d v o r .T h r i s r r o l d r .I d s o m f r l m v r y w l r i l r l i r t i o n . . . . T l r point hivd' In othr
most gratifying part of all is to s how it [th awaiian Drosoplrila Projtl laimd and what rrran
has afftd th livs lf so many kamaina [Harvaiin word for loals] stu- prtnannt ffts on th
dnts'' (Hoolulu Str Bullti, .|un 12, 1985). Indd' arson's Hwaiian work provd that on :
Drosophila Projt has n an irportant trining gound for mor rhn 400 th starting point.
undrgraduat,gr:r1uat' nd postgrduatstudnts irr all spts of volu- Throughout th twn
tionay iology. Also, mor thatr 70 snior sitltistsfrlm all ovr th world slrowing that sltion
hav partiipatd in the projt primarily ausof arson,s ladrship in r h o u g h s t l . sw o r k i
th fild of voluti<rnaryiology nd his aility tl ring togthr th st W a ) . o m o d r n v o l u t
minds in th fild. H has n fthr figur wh<lhas n al tl motivat prirnnt intnding to s
and hallng th rr-rinds of youngr gnrationsttl r-trrlok th srtslf vo. first inrrst was in th
lutionary iolog. H rotivdotllrs to think out horv knlwldgan sllposs. astl wirs tr
hlp psrv th natural hritag of ou plnt. hs quatitis mai this gns' ar not nti an
grat sintisttlr huml prson that h was. up irs rprodution O
tim. It was only aftr }
|BLIoGRAPHY
th ntury that stl
Arrdrson, W. w., . Y. Kanshir,nlL. V. Giddings.l989. Hmpttlnf,:r,n fronr vritions us
rson:lntviwstrwarditlltul history.lll W. W. Adrsorl,K. Y. transmittd unhangd
Kanshiro.ar-rdL. V. Giddings, eds.,(}tis,Spitit' d th Fttdr
Ir-rth ours of his
Priipl,-28. Nw York: Oford UnivrsityPrss.
rson,11,L'. 196.Hrditd Hum Llle. Nw Yrk: olumiaUnivrsity mous and important ol
Prss. ( s P o v i n 1 9 8 6 ) . I n z
1 9 7 0 . h r o r o s o m t r a st lorf < l r i g i r . sr o
p f i e sS. i c1 6 8 : 1 4 l 4 _ 1 4 ] 8 . h foundrs of modr
1986.Su.rlsltionIrdsi:lti<lrr. In S. Krlin lrnd' Nv,ds., rsarh work was on t
uoltitl"tr Prrlsss d Thlt, 391_409.Ollrr-rd, FL: Aal-ri Prss.
tl.stthat wnt ilk t(
Darwin, . 1859. tb rigi o|-Spis. Lndn:Johr-r urry.
Provin,W. B. 1989. l.our"rdr fftsr.rdgntirvllutionsin mirlvolutionnd
|BLIoGRAPHY
spiation: A historilprstir.' In \{/.!(/.Andrson,K. Y. Kanshiro,and
L. V. Giddings' ds.,()tis,Spitio,d tha FodrPriiplc,4_76. stl,W. .' nd J. . I
Nw Ylrk: Oxfrd Univrsity rss. -.Y.. st f th fftit'
edli Cros-s,s \
Pror,in,W. B. 1986.Sap
astl, William F,. (1867-1962) f hiaglPrss.
Rus.A4. 1996.od tr
!illiam . astl, a profssorat Harvard Univrsity nd latr at th BttssyIn- irmridg,MA: H
stitlltion (an agitrlrtrrl muh tl th ilrlv yars
arm of Harvrd), ontrir'rt]

latural history that bgan as a
oung ollgstudnt mmr of
s at th Univrsity of Hawaii's
vry ativin rsarh and on-
o volutionary biology. H is
y o f H r w a i i . sm o s t m i n n ts i -
) oursof his distinguishd a-
gus nd to his many studnts.
rt sintist.on, whn h was
' a symposium hld in honor of
humling.Sinis a slial n-
vry whih dirtion. . . . Th
Hwaiin Droslphila Projt]
[Hawaiian word for loals] stu-
8 5 ) .I n d d , a r s o n ' s H a w a i i a n
ining gror.rndfor mor than 400
studntsin all spts of volu'
intistsfrlm all ovr th wold
laustlf arson's ladrship in
lility t ring togthr th st
. 'w h o h s n l t o m o t i v t
lns to unlok th srtsof vo-
rink out how knowldg an
:1nt.Ths qualitis mad this
dings. 1989. Hmpton Lawrn
.W.
y. In W. Andrson, K. Y.
s, Spitkltt'd tb Fldr
y Prss.
] wY t l r k : | u m l ' i U n i v r s i t y
f spis.Si 168: 1'414_141'8'
S. Karlin nd . Nvo, ds.,
). orland, FL: Aadmi Prss.
n :J o h n u r r a y .
rvolutionsin mirovolution nd
Andrson,K. Y. Kanshiro, and
d th FtludrPriipl, 4_76.
-.Y..
vrsityand latr at th Bussy In-
ltriutdmuh to th arly yars
stl 469
of gntis.Lik nrany of his gnration, h gan as an mryologist, swith-
ing to prolms of hrdity whn Mndl's work was rdisoverd at th -
ginning of th twntieth ntury. astl workd mainly n mammals, although
h was also th first to us th fruit fly Drosophil mlogstr,whih bam
th volutionist's workhors in th twntith ntury. Thomas unt Morgan
Won th Nol Prize for diphring th gnti natur of this organism.
astl's most famolls sris of primnts Wr on rats of varying lak
and whit oat olors (s figur on pg 47). |1 prformd sltion x-
primnts tht showd how on ould hang th ovrall rang of hus in a
population, from mixd to virtually ntirly whit or lak (astl and
Phillips |91'4)..Whatwas signifiantaout this work ws tht on th sl-
tion was finishd, th ts' oat olors ontinud to hovr round th final
point ahivd. In othr words, astl showd what harls Darwin had
laimd and what many ritis had dnid: seltion an hav lasting if not
prmanntfftson th ovrall natur of a population of organisms.astl's
work provd that on sltionis rlad, vrythingdlsnot rvrt ak to
th starting point.
Throughout th twntith ntury thr wr numrous othr primnts
showing that sltion an hv lasting ffts. Paradoially, howvr, al-
though Castl's work is rightly haild as a major xprimntal stp on th
Way to modrn volutionary thinking, h himslf did not ntr into th -
primnt intnding to slrow th worth of sltion. Rathr, as a gntiist' his
first intrstwas in th ntur of th hrital fators that Mndlism pr-
supposs.astl was trying to show that ths fators, what ar now alld
gns'ar not ntir and unhangalin th ours of gnration' ut split
up as prodution ours and romin in nw variations in th ours of
tim. It was only aftr Morgan's work was ompletd in th sond dad of
th ntury that astl ondd that h had n wrong and that (apart
from variations ausd y spontanous hngs' or mutations) gns ar
tansmittdunhang.
In th ours of his long ar, astl had many studnts' th most fa-
mous and important of whom Was gntiist and volutionist Swall \/right
(sProvin 1986).In anothr paradox, Wright is proprly known as on of
th foundrs of modrn volutionary thought, yt th ulk of his mpiril
rsarhwork was on th pattrnsof hrdity as shown y guina pigs' an in-
trstthat wnt k to his studntshioundr astl.
|LloGRAPHY
.W..
astl, ., and J. . Phillips. |L)14.Pibld Rtsd Sltio: primtl
stof th fftiusslf Sltiod of th Tlrof GmetPurit i
.Washington, .lfashington.
dlirosses. D: arngiInstitutionof
rovin,!. B. 1986'StullWrightd ullutior Biolog.hiago:Univrsity
of hiagoPrss.
Rus,. 1996. od to : h optlf Progrssi uolutilrBillog.
amridg, A: Harvrd Univrsity Prss -.R.
470 stle

fifrq
The usesof I

f,
-.2 - !

T 'r
iq
(i
J.B.s.Haldan's
is th apston txt
haild primarily for
hitts of populati
\/
( 1 9 3 1 ) ,a n d a l d n
I a synthsis of lassir
hmistry rvith popr
disussion is onfin
ial nratrial drawn
Arystwyth, ntitl
Haldan's prolr
n of natural sl
shas th prolrna
answr) intrwovn
part follows Fishr i
dt1 to low sltion
dan's support lf Fis
rrid th Inathml
hanging nvlrnm
and in part as a rlt
ovr, and important
ltion, r<lgnizig
,'.* populations may oI
Ths disussins
di of uss. Y|a|
uss of within- an
trs and a road ss
nal hrptr.
.Working
throug t
ry of Mndlism in
and btwn spi
pral 1oi, diffn
mosoms' or o{ wh
Gtics d th Ol
gu tht th sam s
Willianrastl,sdmonstrtionoftfftsofsltitrnforoato1or.op within spisou
r o L u : | e l a t o l o r . . . t . u , " d f o r l a s s i f i a t i o n . i l r l t r tht
l z : sastl's
l t i owork
n-forlight
olr. Th pradx roadns th ndt
o|or.Btlttrlt,ulz'.,.t....l. r",...t tat th units f
was aimd at disp.ov,nj .'"r.r *ntis(v showing Th onlusion
.."^.::pJ:u.c th ffts of natrtral
hrdity lnd in ah gnrtigl), thought sin Darw
a s t l a n d P h i l l i p s l 9 1 4 , P l t 1 . )
'.-i.t'om th sarn tim, Hal
dominant rativ {
rativ aus.hr
volution, inludin5
that mirrds ar sur

+4
\
! (1931),and aldan himslf. It is lss widly, yt just as dsrvdly,haild as
I
. sltionfor oat olor. Top
iddl rop: sltionfor light
' paradois that astl'swork
y shwingthat th units of
ng th fftsof natural
l.)
Th aussof volutin 471
Tb uses of uolutio (I. B. S. Haldan)
J. B. s. Haldan's (7892*1964) The ses of ullutio' pulishd tn 1932,
is th apston tt in th origins of thortial ppulation gntis. uss is
haild primarily for its important, ritial disussion of th work of th ar-
hitts of population gntis' R. A. Fishr (1930 [195s]), Swall \/right
a synthsisof lassil gntis,hromosomal mhanis' ytology, and io-
hmistry with population gntis.This is ironi austh mathmatial
disussionis onfind to an appndi addd to original disussionsof iolog-
.W.als,
ial matrial drawn from Haldan's 191 ltursat th Univrsity of
Abrystwyth, ntitld ..A
Rxamination of Darwinism.''
Haldan's prolmati may posd as, ..What is th natur and signifi-
an of natural sltionin Mndlian popultions?'' In this sns,Haldan
sharsth prolmati of his ontmporaris Fishr and \/right.And in his
answr, intrwovn with supporting mpirial vidn, Haldan in larg
part follows F.ishrin th viw that umulativ volutionary hang is mostly
du to low sltion prssursting on mutations of small fft. But Hal-
dan'ssupport of Fishr is not in th form of an unritial summary. aldan
arrid th mathmatial ploration of sltion in finit population sizs,
hanging nvironmnts, and multipl dimnsions furthr thn Fishr had,
and in part as a ritil onsidration of !right,s work on th topi. Mor-
ov, and implrtantly,Haldan emphasizd th problm of th units of s.
ltion, rognizing that sltiv fors ating on gamts' organrsms' Or
populations may om into onflit.
Ths disussions ar arrid orrt in th middl haptrs and th appn-
di of ss. Haldan mds thm in rih disussion of th gnti
auss of within- and twn-spisdiffrnsdisussd in arlir hap-
ts and a road ssssmntof th sin of volution as whol in th fi-
nal haptr.
\Vorking through mpirial vidn amassd from th tim of th rdisov-
ry of Mndlism in th 1900s, Haldn dmonstrats tht diffrns within
and twn spis ar du to diffrns twn individual gns at om-
paral loi' diffrns in numr of individul hromosoms or sts of hro-
mosoms' or of whol lrromosoms. Antiipating Thodosius Dozhansky's
Getics d th origi of Speies (197), Haldan uss this vidn to ar-
gu that th sam sorts of prosss that aount for th gnti diffrns
within spis aount for suh diffrns twn sis. Furthr, Haldan
roadns th ndlian ontxt of volution.
Th onlusion of uss mphasizs th progrss mad in volutionary
thought sin Darwin and th powf of volutionary planations of lif. At
th sam tim. Hldan is lar that whil h ss natural sltion as th
dominant rativ volutionary us, h dos not lim that it is th only
rativ aus. Througout ses Haldan highlights th multipl auss of
volution, inluding dvlopmntalonstraints ln volution. Haldan insists
that minds ar surely produts of volution;thy also ar in all liklihood not
472 ulliSforz

poduts of naturl sltion.Th road disussion in Haldn's onluding G e n e s \ 1 9 9 4 )f r

haptr is to md th sinof volution in distintly matrialist philos- through in th r
ophy of sin. ronstrution
.sfright
Haldan, similar to Fishr and for him, aomplishd on- whih thy spr
iliation of natural sltion and Mndlian hrdity. aldan forgd ahad graphi distriu
of his prdssors y furthr synthsizing a wid swath of iology with p<lp- riginal populati
ulation gntis. The uses of uolutio is part and parl of th origins of volutionary hil
thortial population gntis, forming what is ommonly alld th mod. In he Neolith
rn synthti thory f volution. Howvr, it is grundraking in its (1984|' h had l
g r a n d rs y n r h t i v i s i o n . to th sprad o
sal with sp
|BLIoGRAPHY
guag diffrnti
Dozhansky,. 1937. Gentisd the origi of Spies.Nw York: olumia guag lssifi
UnivrsityPrss. distriution of
Fishr'R. A. 1930 [19.'8l.The GetilTborof NtrlSeltio.2ndd.Nw
sals.
York: Dovr Pulitions.
aldan,J. B. s. 1932.Th usesof uolutio.I,ondon:I-ongmans. Givn th brr
.sright'
s. 191.volutionin ndlianpopulations.Getis16:97-759' grat dal of W
-R..s. that might hlp
rnsmissil l
avalli.Sforz
ntly his laor
Cavalli-Sf rza, Luigi Lua (t. |922)
tion usd to d
Luigi Lua Cavalli-Sforz an onsidrda modrn rinarnati<lnof R- Th study of th
naissan sholar. is major ontriution to volutionary ilogy mrgd ially fftiv il
from his knowldg of n unusually wid rang of filds that h uitivatd at Cavalli-Sforz
th profssional lvl with a sinr passion for undrstnding th human sarh. H is tI
ondition. olumns. Th t
Traind as a mdial dotor in Italy in th 1940s, h rifly pratid md- with W. Bodm
iin. H thn turnd his ommitmnt to rsarh.Attratd y th statistial fild in 1981.
asptsof gntis,h pursud his intstundr th guidan of R. A. Fishr
at amridg Univrsity. Almost at th sam tim (in th arly 1950s), h B|LloGF
was involvd in pionring wok in atrial sx, a highly innovativ topi. avalli.Sforza,L.
H soon fousd on humans. H ontriutd lasting mthodologial ad- Gtisf I
vans suh as th still-usd valli-Sforza_dwardsgnti distan,an s- avalli-Sforza,L.
sntial tool for phylognti and phylogogaphi studis (nothr fild h San Frnis
Cavlli-Sforz,l,.
pionrd using a diffrnt nam, th ..gographyof gns'').
ullutio,P
H rativly dvlopd ways to us th rihnss of historial rords in valli-Sforz,L.
Italy (.g.,parish ooks and onsanguinity dispnsations)in human popula- of Hum G
tion gntis.H was among th first volutionry sintiststo liv that avalli.Sforza,l-.
drift was important in th volution of human popultilns.In 204, o. GtiDrif'
sguiit, lbredig' d Geeti Drift i ltl organically prsntd th
rsults of his dds.longrsarhffort.
|n 1971h lft Italy to om profssorof gntisat Stanford Univrsity
hamrs, t
in alifornia, whr h is urrntly ativ mritus prfssor. Rort ham
avalli.Sforza was a pionr in th us of modrn human gnsfor ron- Britain. Togtht
struting th ntural history of man. The Histor d Glgrph of Humn abjt povrty

ussionin ldan's onltrding
in a distitlymatrialist philos-
for him' omplishd a ron_
hrdity.Haldan forgd ahad
rvidswath of iology with pop.
part and parl f th origins o{
lat is ommonly alld th mod-
. r . i t i s g o u n d r
f Spis.Nw York: olunrbi
f NturISltitllt.2ndd. Nw
. l,ondon: Lngmans.
tions. Gtis |6: 97_1'59.
)\
] a modrn rinarlrationof a R-
to voltionaryiollgy mgd
ang of filds that h ultivatd at
otr for undstandingth human
1940s,h rifly pratid md-
s;rrh'Attatd y th statistial
ndrth guidanof R. A. Fishr
rm tim (in th arly 1950s), h
. i l s .a h i g h l y i n n o v a t i v t o p i .
utd lasting mthodologial ad-
'_dwardsgnti distan, an s-
grphi studis (irnothr fild h
gaphyof gns',).
rihnssof historial rords in
. dispnsations)in lrurnan popula.
lutionary sintiststo liv that
rrranpopulations. In 24, o-
i ltl' organiirlly prsrrtdth
of gntisat StarrfordUnivrsit
mritusprofssor.
f modrn human gns for ron-
Llistlrd Geograph of Humn
hmbers 47
Gs (1994) fittingly slll.Ilsth radth of lris ndvor. Haild s brak.
through in th urrdrstanding of human volution, it offrs th first full-sal
ronstrution of whr humn populations originatd nd th paths y
whih hy sprd throughout th world. By mapping th worldwid go-
graphi distriution of gnsfor ovr 110 traits in ovr 1'800 primaily ao-
riginal populations' migations we hartd and lok was dvisd to dat
voiutionry history. ssntial information was ntriutd by arhology.
In Th Nolithic Trsitio d th Geetis of Popltios i urlpe
k i n gi n i t s (1984)' h had addrssdth qustion of wthr ultural diffusion was du
to th spad of idas or of popl. Th issu was invstigatd at a world
sal wih spial attntion to th link twn gnti divrsity and lan-
guagdiffrntiation.Although linguists hav not rirhdonsnsuson lan-
guag lassifiation at gloal sal' an intriguing ovrlap btwn th
distriution of gns and languags Was found at diffrnt gographi
sls.
Givn th brdth of his rsarh intrsts, it is not surprising to find a
gratdal of wok olltd in a ook ddiatdto th ploration of modls
-R..s. that miglrt hlp us undrst;rnd how ultur:rl traits trr trnsmitted (ulturl
Trsmissiond uolutil' oallthord with . w. Fldman' 19s1).
avalli.Sforz has njoyd a rmarkaly rativ sintifi longvity. R-
ntly his laboratly dvlopd nw thniqus for disoving DNA varia-
tion usd to dfin nw gnti markrs of th Y and othr hromosoms.
Th study of ths uniprntally tansmittd marks trnd out to esp-
ially fftivirr luidting popr:lation origins.
Cavalli-Sforza has natural gift for ommuniating his passion for r-
sarh. H is th author of mny gnral audin ooks and nWspapr
olurns. Th ttook The Getis of Humt Popultios (oauthord
with !. Bodmr, 1971| was awardd a prize as th most quotd tt in th
f i l di n 1 9 8 1.
ILIoGRAPHY
L. L.' and A. Ammrman.,l,984.h Neolithicrsitiond tb
availi.S{oza,
Gtiso|-Popultiotlsi rop. Printon,NJ: ritonUnivrsityPss.
.W.
avalli-Sforza,L. l-., nd Bodmr. 1971.b Gtisof Hum Popultios.
SanFrniso:W. H. Frmn;inola,NY: DovrPulitions.
avalli-Sforza,
L. L.' and . W. FIdmn.198l . ulturITrnsmissil d
uolutkl,Printon'NJ: PrintonUnivrsityPrss.
valli-Sforza,L. l-', P. nozzi, and.Piazza. 1994. he Histor nd Gogrph
of 17t,ttGs,Prineton,NJ: PrintonUnivrsityPrss.
avalli-Sforz,
L. L., A. oroni, and G. Zet.24,osguinit, Ibreeding,
d
GticDrift i [tl.Printon,NJ: PrintonUnivrsityPrss. -P.'
Rort(Is02_1s7I)
hamrs,
Rort hambrs Was ol-l of tlr most sussful usinssmn in Vitorian
Britain. Togthr with his rothr, William, h had pulld himslf up from
abjt povrty to found a thriving pulishing businss. Thy wr major
474 hmbrs

innovators in th us of stam prsss' and thy wr at th for of th pro- lLIoGRAPY

dution of popular litratur and magazins. A wkly puliation, ham-
Chamrs' R. l844. Vs/l
ers,slournal of Litertr, Si d Arts, of whih Rort and William
|845. pIli,'s
wr oditors and to whih Rort ontributd many smll pis,Wasvry Darwin, . 1859. th ()r
sussful. Rus' . 7996. od to
Rort hamrs was always rady with his pn. H wrot ooks and amridg, A: Harva
artils as wll, mainly of a popular and (with an y to sals) informativ, Sod, J. A. 2000. vi!ri
d Srt thorship tl
gossipy natur. Prim ampls inludd guids to walks around din.
hiago: Univrsitv of (
urgh, listing plas whr snsational murdrs and othr ill dds had o-
Tnnyson' A. 1850. In m.
urrd in past years-prft matrial for a family fad with th nd to utb orittiu t Bk
fill th long hours of yt anothr Sottish Sunda. As wr many othr Norton.
half-duatd mmrs of soity, hamrs was also vry intrstdin
sin, partiularly gology. In th arly 1840s, this ld him to writ what
was to om on of th snsational works of th ra, Vstigsof th Nt- Chan and voluti
url Histor of retio (1.844),togthr with a squ1titled pltios T h o n p o f h n i s l
(1845).
T h s a s s o i t i o n sr a n g
Apparntly, hamrs had st out to writ a ook on phrnology (th si- tan to playing a ntra
n of rain umps), anothr popular psudosintifitopi of th day' ut molular iology. Thr
h got sidtrakd into writing a ook on volution. (SFigur 3 in th main lutionary thory ar disr
ssay ..Th istory of volutionary Thought'' y Mihal Rus for dtails of D a r w i n 1 8 5 9 , 1 3 1 ) , h a r
th way that hamrs onivd of th volutionary pross.) Givn th and marovolution. Wit
ontnt of Vstiges' it was rtainly not intndd as a all to athism, ut disussd. Howvr, tra
mor an appal to th lawlik natur of th world. (Although hamrs was ar yond th sp of t
not kn on rgular hurhmn, h was always a livr in a disti way.) In It is imPortant to not
ommon with many othr volutionists' his ook ws a hymn f prais to dp, mtaphysial sns
progrss. As w hav improvmnt in th sial world' so also do w hav it of han is most oftn rl
in iology' and onvrsly as w hav improvmnt in th ilogial world, suh as in quantum ml
so also do w hav it in th soial. fair oin is preditd st
Baus of his sinss and soial position, hamrs wnt to grat Th rsult has a ausal t
lngths to onal his authorship. Indd, it was not rvld offiially until oin flip ar not random
1884, aftr his dath. It was proaly as wll that h hid his nam' aus disussions of han r
Vstigs was highly ontrovrsial. any in th sintifi ommunity wr x- mhanisms.
trmly ritial, lading harls Darwin, just thn formulating his volution. Natural sltion is thr
ary thory' to kp quit aout his own ativitis. owvr, thr wr many lution. It was, in part, f
who rspondd positiv| to Vestiges, most notaly pot Alfrd Lord Tn- from rival thoris and l
..In moriam,'' a triut to th
nyson. was omplting his long pom I y i n g o n h n m U t t i o
mmory of his dar frind Arthur Hallam. Dprssd by what h saw as th adaptiv omplity of t
mpty nihilism of Lyellian uniformitarianism-no hang, no progrss-h spis hibit han v
drw omfort from th optimism of Vstiges' with its support of progrss. dltrious. Barrs tlf s
Prhaps it was that th dad-too.young Hallam Was a suprior ing orn thus thy rprodu anr
for his tim. (S th alphatial ntry on Vstigs for dtails.) ovr tim, th sussf
Vestiges mad th ida of volution wll knwn in th mid.Vitorian r gumnt tht hn vaI
and pavd th way for Darwin and his o th rigi of Speis. By th tim strutur as omplx as
the origi appard, sintists and laypopl wr hustd from arguing, o f D a r w i n i a n v o l u t i o n
th shok of th ida of volution WS ovr' and a mor aland aount of
t 1 s 1 9 ] ) a r t i u l a t dt h i s
hang ould graspd and aptd y many. world that was a sul

:IyWr at th for of th pro-
. A wkly puliation' ham.
\William
;, of whih Rort and
td many small pis,was vry
r his pn. H wrot ooks and
l t h n y t o s a l s )i n f o r m a t i v '
guids to walks around din-
drs and othr ill dds had o-
r family fd with th nd to
Sunda. As wr many othr
rs Was also vry intrstd in
840s,this ld him to writ what
s of th era, Vestigs of th Nt-
lith a squl tit|edpltios
' a ook on phrnlogy (th si-
dosintifitopi of th day, ut
olution.(SFigur 3 in th main
t'' y ihal Rus for dtails of
:volutionaypross.) Givn th
'tndd as a all to athism, ut
world. (Although hamrs was
ays a livrin a disti way.) In
.s ook was a hymn of prais to
ial world, so also do w hav it
ovmntin th iologial world,
sition' hambrs wnt to grat
it was not rvald offiially until
,,llthat h hid his nam' aus
th sintifiommunity W x-
rstthn frmulating his volution-
ivitis.Howvr' thr Wr many
st notaly pot Alfrd Lord Tn-
..In Mmoriam,'' a triut to th
Dprssdy what h saw as th
ism_no hang, no progrss-h
lges,with its support of progrss.
{allam was a suprior ing orn
on Vstigesfor dtails.)
ll known in th mid-Vitorin
t the origi of Specis. By th tim
l p l w r h a u s t df r o m a r g u i n g .
.r' and a mor aland unt of
mny.
h d L,uoltttio 475
BIBLIOGRAPHY
hamrs, R. l 844' Vstigs rlf the Nturl Histor lf rtio. London: hurhill.
1845. pltils: Sequl tl
.,Vstigs
lf rtkltt.'' London: hurhill.
Darwin, . 1859. th origin of Speies.Lndon: John Murray.
Rus, . 1'996. od to : h opt of Prlgrss i uolutior Biolog.
amridg, A: Hrvard Univrsity Prss.
Sord'.I. A. 2000. Vitlri Sestiln:h trordir Publitio, Rptitl,
,,Vstigs
d Srt Authorshi1l lf lf th NtrI Histor of rtio.''
hiago: Univrsity of higo Prss.
Tnnyson'A. 1850. In morim. In R. H. Ross, d., l mlrim:
thorittiu t Bkr-ludsd Sors ritiism,3_90. Nw York:
-.R.
Nortn.
hanand volution
Th onpt of han is ssoiatdwith volutinary thory in svralways.
Ths assoiationsrang from ing f historil and philosophial impor-
tn to playing a ntral rol within thnial dats in volutionary and
molulr iology. Thr gnra1domains whr han is mployd in vo-
lutionary thory ar disussd:harls Darwin's us of han (for xampl,
Darwin 1859, 131), hn and mirovolution (random drift)' and han
and marovolution. Within ah mp svrl fin-grainddistintions ar
disussd.Howvr, tratmntof th mor thnial asptsof random drift
ar yond th sop of this ntry.
It is important to not that iologists gnrally do not us han in a
..Llnusd.''Th unausd sns
dp,mtaphysial sns,suh as maning
of han is most oftn raisd in philosophial disussionsof indtrminism,
suh as in quantum mhanis. For ampl, th randm rsult of flipping a
fair oin is prditd stohastially, ut it is dtrmind y physial laws.
Th rsult has a ausal story' so th undrlying prossstht dtrmin a
oin flip a not random, or stohasti,in a dp sns.Likwis in iology,
disussions of han rarly ntail th indtrminay of th urrdrlying
mhnisms.
Natural sltionis th privilgd mhanism of hang in Darwinian vo-
lution. It Was' in part' Darwin's us of han that distinguishd his thory
from rival thoris and worldviws. Darwin dsrid a natural pross' r-
lying on han mutations and natura1sltion,whih ould aount for th
adaptivomplity of th iologial world. In this pross' individuals of a
spisxhiit han variations, som of whih ar nfiial and othrs
dltrious.Barrs of sussful traits ar mo fit thn thir omptitors,
thus thy rprodu nd pass on thir gns to offspring at a highr rat.
ovr tim, th sussfultraits sprad to ll mmers of th spis.Th ar-
gumntthat hn variations ould offr th raw matrial for volving a
struturas ompl s a vrtraty smduntnal to the opponnts
of Darwinian vlution. Th famous nglish apologist William Ply (1802
[1819])rtiulatd this point of viw whn h drw th ontrast twn a
world that ws a result of han and natural prosss or th rsult of a
476 he d uoltil

dsignr.To Ply, th hoi was ovious; th intriat omplity of fa- population of alls
turs suh as th y was inompatil with anything ut a dsignr,a thsis nal population-y
ommonly rfrrd to as th Argumnt from Dsign. whr th gnotyp
Darwin's us of han variations distinguishd his idas not nly frm ltiv prssurs.
thos of rationists ut also from othr advoats of ompting thoris of Thr ar two g
volution, partiularlyJn-BaptistLmark (1809). Arding to Lamark, a atastroph or ot
mutations ar dirtd toward an organism's nds; thy aris with rspt t suh s a forst fir
th us and disus of an organism'sparts. For xampl, giraff nks r lon- ass it dos so indi
gatd aus giraffs strthd thm to rah trtop foliag. opposing this ganisms. h smal
modl, Darwinian volution lims that mutations ar undirtd, han gnration may hal
vnts;mutations for long-nkdgiraffswr just as likly as mutations for ltion. A similar for
short-nkd giraffs. Howvr, on nk-lngth mutations ourrd, th is, an ur wh
long-nkd giraffs had a highr fitnss valu, thus sprading thir long-nk poulation and sta
gns throughout th population. Th ritial point in Darwin's argumnt is Th migrating grou
that although mutations aris y han, it is not y han that som indi- ity with thm. Th
viduals surviv and rprodu whil othrs pish. Natural sltion distin- rat a nw popul
guishes twnnfiia1nd dltriousmutations. land prognitors,
hos of ths argumnts mrg in modrn ritiisms of Darwinian vo- rol. This pross l
lution that ar fousd on th sming surdity tht han variations importan ar link
ould vr produ intriat struturs or wll-ordrd iohmial pro. A sond form o
sss.Ths argumnts ar asd on mistakn idas of th rol of han th nxt, in all limi
that ho th argumnts in Darwin,s tim. Fist, han variations produ akin to Dozhanky
raw matrial that is moldd by natual sltion;without natural sltion qunis hang gr
to st th dirtion and spd of hang, hn variations ould not au- alll oms th '
mulat with nough spd to produ muh mningful hang in any stru- ours usth
tu or pross. Sond, han atually has a spifi maning with rspt ll frqunis.
to Darwin's han variations nd th gnti mutations tht aus thm. mon' and an unO
FIr,han mans that spifi mutations our at random with rsptto beoms xtrmly
th nvironmnt so that, unlik th l,amarkian argumnt' mutations ar aus allls may
not dirtd y nvironmntal fatursor th liklihood that thy will prov whih on will vr
usful. Som spifi mutations ar indd mor likly thn othrs-for x- Swall !right, r,
ampl, som mous oat olor mutations ar mor likly to aris than sponsil for fusin
othrs-ut thir potntial usfulnsshas no aring on thir liklihood of first to mphasizt
apparing. gud that thr wr
\Whnontmporary iologists rais th notion of han thy likly ar r- advantagous for tl
frring to rndom drift. Random drift rfrs to th hang in gnti ompo- of ths ominati
..shifting alan''
sition that ours in a population from on gnrationto th nxt ausof
.s7right
th random smpling of gns that ours in a finit population. A familiar rr, would
ampl of random gnti sampling is th s ratio within a family; in th- 1940s through th
ory, half of th hildn in a family should oys and half girls, ut it is not lution (Provin 198
unommon for a fmily of four hildrn to onsist of thr oys and on girl Th most influr
ausof th random draw of th s hromosoms.Th lassi illustration atd argumnt for
o f r a n d o m d r i f t w a s g i v n y T h e l d o s i u sD o z h a n s k y ( 1 9 7 , 1 2 9 ) ' w h r h thory of volution
dsrida snario of lindly drawing alls fom an urn. Th alls wr ol. wr disovrdan
ord diffntly, ut othrwis indistinguishal.If on lindly draws alls gl protin that s
from an urn and plas thm in sond urn, it is likly that th sampl wr two or mor

i; th intriat omplity of fa-
r aIrthingt a dsignr' a thsis
n Dsign.
rguishdhis idas not only from
Jvoiltslf ornpting thoris of
:k (1809).Aording to Lamark,
's nls;thy aris with rspt to
r ampl,girff nks r lon- assit dos so indisriminatl1,rvith rsptto ny prtiular rit in th or-
lh trtopfoliag. Opposing this
xutations r rrndirtd' han
vr just as likly as mutations for
k-lngth mutations ourrd, th
lu,tlrrtsspradingthir long-nk
| p o i n t i n D a w i n ' s i l g u n r n ti s
t is not y han that som indi-
.s prish.Natural sltion distin-
11tlt1tlons.
l d r n r i t i i s m s f D a r w i n i a n v o -
a s u d i yt h a t h n v a r i a t i o n s
rr wll_ordrdiohmial pro-
;taknidas of th rol of han
. First, han variations produ
lltin;withut naturl sltion
han variations ould not au-
:h maningfulhang in any stru-
:as spifimaning with rspt
:nti mutations that us thm.
rs oLlrat random with rspt to
. uttions
r n r k i n r g u m I . l tm
th liklihood that thy will prov
c mor likly than oths-fr -
ns r rnor likly to iris than
; no aring on thir liklihood of
notion of hn thy likly ar r-
: r sr t h h n g i n g n e t i o m p o -
gnrationto th nxt aus of
s in finit population. A familia
l s ratio within a family; in th-
[ oys and half girls, ut it is not
o onsistof thr oys and on girl
omosoms.Th lassi illustration
D o z h a n s k (y1 9 7 , 1 2 9 ) ,w h r h
'lls from an urn. Th alls wr ol-
rishbl.If on lindly draws balls
Ld urn, it is likly that th sampl
hn d uollttion 477
population of alls diffrs in frqurry'wit gard to olor, from the origi-
nal popultion-y han lon. In short, rarrdm drift dsisa snario
whr th gnotyp frquny of a population hangs in th :rsnof s-
ltiv prssurs.
hr two gnrl kinds of random drift: isd smpling as rsult of
a atastroph or Othr norrsltiviasing, and gnti drift. atastroph'
suh as forst fir' may kill a larg portion of a population, ut in most
ganisms. Th srll num of survivors tht ar lft to produ th nt
gnrationmay hav a diffrnt gnti omposition than th original popu.
ltion. A sinrilar fom of drift, thar is, n rupt hng in gnotyp frqun-
is, an our whn a srnall gror.rpof organisms migrat from th main
population and start a nw on' suh as ours whn an island is olonizd.
Th migtinggroup may only tk :r fration of th original gntivariail-
i t y w i t h r h m . T l r u n i q u g n i s u t u r o f t h f i l u n d i n g p o p u I a t i o n a n
rat a nw popr"rlationthat is ognizaly distint fom its oiginal' main-
land prognitors!vn though natual sltio my av playd a minimal
rol. Tlris pross has n alld th foundr fft, and argllmnts for its
importnar linkd to th influnof rnst Mayr (1954)'
A sond form of randorn drift ours grdually, fror on gntion to
th nt, in all limitd populations through indisrinrinatgamt sampling
akin to Dozhanky's analogy of th alls in tlr urn. Gnotyp nd alll fr-
qunishang gadually aross ]nrations'irnd, givn nough tim, on
alll oms tlr only gn in th populatiln (that is, it is fid). Fiation
ours austh dirtion of andom gnti drift is dtrmind y th al-
ll fqunis.Tvpially' a ommon lll is liklv to om mor om-
mon' and an unommon on is likly to om rar. vntually on lll
omstrmly ommon and thn Ad. Th han lmnt rmains .
aus allls ma qually ommon' thus rrr:rkingit impossil to prdit
whih on will vntually bom fid.
Swall Wright, who along with J. B. S. aldan and R. A. F.ishrwas r-
sponsilfor fusing Mndlian gntiswith Darwiniar-rvolution' was th
first to mphasiz th rol that random drift plays in volution. Wright ar-
gud that thr wuld ftrr diffrnt ominations f gns that would
dvantagousfor thir arirs and that random drift would influnwhih
of thsomirrtionswould mrg and sprad through a spis(Ii7right's
..shifting
alan'' thory of volution). Howvr' throughout his long a-
.!7right
rr, would vaillat on th importan of rndom drift and in th
1940s through tlr 1960s h dlgtdrandom drift to minr for in vo-
l u t i o n ( P r o v i n 1 9 8 6 ) .
Th most influntial appliation of gnti drift-and th nrost sophisti-
td rgumnt fo th rol of h:rrrin vlution-is th so-alld nutral
thoryof volutiln.As prtins and th DNA squnsthat odd for thm
wr disovrdand plord, iologistswr finding nrany variants of sin-
gl prltin that smd to hv quivalnt funtionirrg.In som ass' thr
Wr two or mor variants in :r singl ppulation and, in othrs' diffrnt
478 hne d uolutio

spis wr fixd for diffrnt vriants. otoo Kimura rognizd that ths Mayr, . 1954. hang
osrvationswr fats of a singl prolm unitd y th onpt of gnti Hardy, and . B. Ft
.$right's Alln and Unwin.
drift. H xtndd original mathmatial work on modling of gn-
Millstin, R. L.2000. (
ti drift to show how thspattrnswould mrg (Kimura 1983). This work
no.4: 603-624.
ld, in turn' to using DNA squnsas ..molularloks'' and to a rvolu- Paly'
.!.
l1802l 1819.
tion in undrstandinghow spis,gnra' familis, and vn ntir phyl ar Rivington.
.!.
r l a t d( K i m u r a 1 9 8 3 ) . Provin, B. 1986. Se
From a marovolutionary prsptiv'som iologists argu that vnts of hiago Prss.
suh as tintions, spis divrsifiation' radiations, and inrasing om-
plxity an st xplaind stohastially,without rfrnto a partiular
usal story. This is not to say that thr ar no undrlying auss to
Charlsworth, Br
marovolutionary vnts, ut that stohasti mthods sm to aptur ths Brin harlsworth il
vnts ttr than dtrministi mthlds (Millstin 2000). For ampl, is 50 yars. H is knowr
th inrasing mplxity of th organi world du to passiv or ativ imntal work has
trnds, that is, y han or dirtdvolution? Although it is diffiult to pro- for svral rasons:a
vid nssaryand suffiintonditions for iologial omplxity, it is gnr- tion of th onntior
ally agrd that lif originatd as simpl organisms and grdually am and numrial appro
muh mor ompl. Baus lif gan nar th minimum oundary o{ work of oths, and al
omplxity, rndom variation alon would produ mor ompl organ- Brian harlswortl
isms; this would a passiv trnd. An ativ trnd would mplifid y BA in Natural Sin
dvlopmntal routins or phnotypi novltis that driv volution toward honors in 1966.He m
inrsing omplity, yond that of hane (arroll 2001). orah harlsworth' l
A final sns of han in volutionary thory is that arriving at any pati- graduation' oth ha
ular nd point in volutionary spa is a han vnt. For xampl, Stphn .
studying undr J. .
Gould (1,989)fmously laimd that humans ar an improbabl nd point on Upon riving his
th volutionary tr;many unrlatd (han)vntsulminatd in th is- with Rihard Lwonti
tn of humans. Gould did not imply that thr ar not sltivrasons an offi with Td G
(mong othrs) xplaining th volutitln of humans. Rathr, h argud tht populations. B.. r
humans ar a rsult of a vast numbr of improal vnts.This is th sns lm; this ffort produl
of han, attriutd to Aristotl, whr indpndntausal hains omin fin fitnss and dsr
to produ an unintndd fft. A ar aidnt, for ampl' is n instan hangs. His ook,
of spartausal hains omining and produing an unintndd rsult. rmains th primary s
In 7971B.. was h
BIBLIOGRAPHY
tiatd xprimntal
Batty'J. 1984.hannd naturlsltion. Phil<lsrlph
lfSi51, no. 2: sions with D.. ld t]
183-211. hangs that afft tht
arroll' S. B. 2001. Chanand nssity:
Th volutionof morphologial
ifirs f rominatior
omplityand divrsity.Ntttr409 (Frury22|:1102_1109,
Darwin, . 1859. th rigi of Sples.London:John uray. immdiat rsult was
Dobzhansky,. |97. Gtisd th Origi of Sples. Nw York: olumia tht launhd D..'s
UnivrsityPrss. Th harlsworths
l,G. J. 1999.On th dul naturof hrrin volutionaryiologynd Maynard Smith (B..
paloiology.Plobiolog25 75*87. la at th National ln
Gould' s. 1989. WonderfulLif: The BurgssShld tb Ntur o/Hls/ry. Nw
olina bgan B..'s wor
York: W.'W. Norton.
Kimura, 14. 1983. h Nutrl Thrlrof olulr uolutio.mridg: tr' Jan, Was orn in
ambridgUnivrsityrss. Aftr attnding a sy
Lamark,J. B. 1809.Philosophizoologiqu'2 vols.Pris:Dntu. no-Darwinism in 19
 hrlsorth, Brin 479

o Kimura rognizdthat ths ayr, . 1954. hng f gntinvironmnt and volution. In J. uxly, A. .
unitdy th orrptof gnti Hardy, and . B' Ford, ds., t.'olutions Proess, 157_180. London: Gorg
Allrr and Unwir-r.
Ltialwok orr modling of gn_
illstin, R. L. 2000. han zrnd mrvo|ution,Philosoph of Sine 67,
r r g ( K i m u r a 1 9 8 3 ) .T h i s w o r k no.4: 603-624.
llularloks'' and to a rvolu- Paly,W. [1802l 1819. Nturl Thollg (ollted Wlrks' vol. 4). London:
milis,and vn ntir phyla ar Rivington.
Prvin,W. B. l986. S,ll Wrigt ld uollttior1,Biolog. hiago: Univrsity
of hiago Prss. -].Z.
lm iologistsargu that vnts
r l i t i o n s. n d i n r s i n g o m .
without rfrnto a partiular
' r no undrlving auss to harlsworth,Brian (b. 1945)
mthodssInto aptur thS Brian Charlswoth is on of th frrnostpopultion gntiisrsof th iast
u{illstin2000). For xampl, is 50 yars.H is krrown primarily for his thotialwork, although his pr-
w o l d d u t o p i r s s i v O r a t i v imntal Work has n sustantial. harlsworth's work hs n influential
ln? Although it is diffiult to pro- for svral rasons: willingnss to tak on diffiult prolms' an appreia-
l i o l t l g i l o m p l x i t y .i t i s g n r - tion of th onntion twn tlrory nd data, lvr us of mathmatial
rganismsand gradually am and numrial pproahs, honst assssmntsof th shotomings of th
lar th minimum boundary of work of othrs, ad an unusually road knowldg of th sintifi litratur.
l produ mor omplx organ- Brian harlsworth (B..) was born in ngland in 1945 and otaind his
/ trnd would mplifid y BA in Naturl Sins from Qun's ollg, amridg, with first lass
ltis that driv volution tward honors t |966. H mt his futur wif, Dorah Malrby (. |94, now D-
: ( a r r o l l2 0 0 1 ) . orah hrlsworth' hraftr D..), in his first ar at amridg. Upon
: o r r .i s t h e t a r i v i n ga t n y p r t i - graduation,blth harlswots Wnt on to PhD studis at amridg, B.C.
tn vnt.For ampl, Stphn studyingundr J. NI. Thoday.
s a an improal nd point n Upn riving his dgr in |969, B.. took a postdotoral fllowship
) vntsulnrinatdin th xis- with Rihard Lwontin at th Univrsity of Chigo. At hiago, B.. shrd
rt thr ar not sltiv rasons an offi with Td Gisl, wo was simulating volution in ag-struturd
'hutlans.
Rathe, h argud that populations.B.C. realizd tht h ould otain analytial rsults on th pro-
prolvnts,Tlris is th sns lm; this ffort produd a sustantialody of work that showd how to d-
dpndntausal hains ombin fin fitnss and dsri tlr rlationship twn gnti nd dmographi
i d n t ,f r a m p | .i s a n i n s t n hangs. His ook' uolutillt i g-Strtured Popultios (1980, |994),
oduing an unintndd rsult' rmainsth primry sour on th subjt.
| 1971B.. wils hird s a lturr at th Univrsity of Livrpool and ini-
tiatd primntal work on th voiution of rornination rats. Distls-
hilosophof Si51, no. 2: sions with D.. ld thm t study modifir thory' whih onrns gnti
hangs that afft th volutionary fors ating on othr loi, suh as mod-
:volrrtionof morphologial ifisof romination, dominan' mating systms,and nrutation rts' Th
.ur2v 2 ) :l 1 0 2 _ 1l 0 9 .
immdit rslllt was a ollaorativ sris of paprs on Batsian mimiry
rdon:JlhnN{r-rrr;ry.
of Spis. Nw York: olumi tht launhd D..'s arr irr volutionar biology.
Th harlswlrths movd to sussx in |974, thanks to fforts y John
iologynd
in vllutinary aynard Smith (B..'s hD aminr).A saatialyar in huk Langly's
la t th Nationirl Institutof nvironmntal Halth Sinsin North ar-
\Id th N,tturr;f Hrrr,. Nw olina ganB..'s work on tri1nsposal lmnts.Tli harlsworths' daugh-
tr,Jan, Was born in 1981.
ulruoltklz. amridg:
Aftr attnding a symposium at hiago,s Fild Musum on altrnativs to
vols.Paris:Dntu. no-Darwinism irr 1980, B.. oauthord a papr with Monty Slatkin and
h rlesulorth,D eb orh

Russ Land that sharply ritiqud hallngsto no.Darwinism. Ths hal- of Livrpool, D.. was
lngs prinipally stmmd from Stphn J. Gould's tnsions of Nils l- miriry y thir oll;
drdg and Gould's onpt o{ puntuatd quiliriunr. Tis papr hlpd in Batsian mimiry wa
atapult B.C. from a rsptd prt on mathmatial thry to a ladr in trn ould rought
t h f i l d o f v o | u t i o n .B . C . h s s i n t k n m a n y o } p o r t u n i t i tso r i t i i z B.. ld to ir ollaora
idas outsid orthodo no-Darwinism' frrn Garil Dovr's suggstiontht D..'s arr in volut
transposal lmnts might play a ausal rol in volution to rnt at- harlswort |975).
tmpts to rt a sinof IntlligntDsign. Undrstanding th r
Th Chalsworths movd to th Univsity of Chiago in 198.5,whr drstanding how volut
B.C. was dpartmnt hair fom 1986 to \997' The harlsworths' transi- att:rkd oth fats in
tion to th study of molular volution Was mntdy thir 1993 disov- stirnding of how gns
ry, with artin Morgan' that dltrious mutations rdu th n.runtof othr loi. This work w
variation in parts of th gnom with low rmination. This providd a gnOm volr-rtion.
novl planation fo a wll-stalishdpattrn that was thn asrid to th D.C. pursud th stt
hithhiking ffts of nfiial mutations. n-roststriking prolms,
| t991, B.C. was namd a Fllow of th Royal Soity,whih failitatda rorphisms that ford
rturn to th Unitd Kingdom via a Royal Soit Rsarh profssorship. and harlswort |979
B.. ontinus to work primarily in mlulr volution and th volLltionof brding systm itslf '
s hromosoms. s a powrful frgui
(Charlsworth 2006).
IBLIoGRAPHY D..'s alrivmntst
harlsworth,B.1.978.A modlfor volutionof Y hrnrosoms and dosag sity of hiago (1988_1
ompnsation . Prodigsof th Ntiol dmlf SiesUS 7.\: as wll as a srisof hol
5618-5622. low of th Royl Soi
Ilgsol |994. uolution i g-StruturdPlpultios.2ndd. mbridg: Biology and volution (
amridgUnivrsityPrss.
gnom volution, part
harlsworth'B.' R. Land,and Nl. Slatkin.1982.A no-Drwiniirn ommrltar
on marovolutj'on.uoluti on 36: 47449 8, and nourags studn
Chrlsworth, B., ' T. Morgn,nd D. harlsworth . 199.Th fftof thir minds to aompli
dltri<lus muttinsn nutralmlulrvariatiln. Gtis|4:
1289-|33.
B|LloGRAPHY
harlsworth, B., P. Snigowski'and !. Stphan.7994.Tevolutionary dynamis
of rptitivDNA in ukryots. Nture37|: 215_220. -D.H. hrlsworth'D. 2006..
R726-R735.
2007.Q&A:D
harlswrth' Dorah (b. L94\ harlswth,D., trdB' l
mimiry.1. Singllo
Dorah harlsworth (nalty) is a pionr in the study lf rdingsys. 1'979a.volution
tms and gnom volution. onntratingon how r'o|uriortmolds stions t979. Mdl for
of th gnom to allow th gns for omplx traits to o.inhritd so that 467-498.
th traits fntion proprly without th disruption that rmination and
sxual rpodution might othrwis produ.
Dorah Charlsworth (D..) otind hr undrgraduat dgr from htvrikosrg
ambridg Univrsity, whr sh mt hr husand, Brian harlsworth Srgi Srgvihhtv
(B..). Aftr taking hr dotoral dgr in iohmial gntis, D.. ws sian ntomologist, an
not al to otain a position in sin and workd s B..'s unpaid assis. ntis. Born into a w
tant for th nt 10 yars. Aftr B.C. tok a position at th Urrivrsitv th Univrsiry of os

o no-Darwinism.Ths hal-
iould'stnsions of Nils l.
1uilirium.This papr hlpd
lmtialthory to ldr in
l a n y o p p o r t u n i t i st o r i t i i z
]aril Dovr's suggstion that
cl in volutin to rnt at-
y of hiago in 1985' whr
l l . h h r l s w o t h st' n s i -
mntdy thir 1993 disov-
]tations rdu th anrount of
omination.This providd a
n that was tn asidto th
'yl Soity, whih failitatd a
oity Rsah profssorship.
volutionand th volution of
hromosoms nd dsag
US 75:
:dmof St:ies
Ppultks.2nd d. rbridg:
ommntary
A n-Drwirlian
.rth.1993.Th fftf
riatin.Getis|4:
l 9 9 4 . h v o l u t i n dyy n a m i s
2rs-220.
in th study of rding sys-
r h o w v o l u t i o nm o l d s s
traitsto o-inhritdso that
]Ption that romirration and
r Llndrgraduatdgr from
husand, Brian hrlsworth
liohmialgntis' D.. was
w o r k d s B . . ' s u n p a i d a s s i s -
< p o s i t i o r la r t h U n i v r s i t v
hetuerihou 481
of Livpool, D.. Was nouragd to work on th volution of Batsian
mimiry y thir ollagu Philip Shppard. on of tlr unsolvd prolms
in Batsian mimiry was how th svral gns ontrolling th mimti pat-
trn or'rld rought tgthr to inhitd as a unit; disussions with
B.. ld to a ollaorativ sris of paprs on this prolm that launhd
D..'s rr in volutionary iology (bginning with harlsworth and
l r a r l s w r t h1 9 7 5 ) .
Undstanding th o-inhritan of gns has a onvrs, whih is un-
drstanding how volution raks up a o'inhritd group. D.. and B.C.
attakd oth fts in svrl pprs' panding to a mor gnral undr-
standing of how gns at on st of loi afft th volution of gns at
othr loi. This work would vntually point toward thir roadr work on
gnom volution.
D.. pursud th study of o-inlrritan to som of th plant world's
most striking prolms, th volution of molular and morphologial poly-
morphisms that ford plants to outross and not inrd (harlsworth
and h:rrlswort|1I979a, 1979|. This work ld naturally to studis of th
rding systm itslf and how th volution of rding systms srvs
as a powrful for guiding th volution of gnom strutur as a whol
( C h a l s w r t h2 0 0 6 ) .
D..'s ahivmnts and rputation ld to faulty positions at th Univr-
sity of hiago (1988-1997\ and th Univrsity of dinurgh (1997-prsnt)
as wll as a srisof honors and rognitiotls,inluding hr ltion as a Fl-
low of th Royal Soity (2005) and prsidnt of th Soity for olular
Biology nd vlution (207).Sh ontinus to study rding systems and
gno volution, partiularl th volution of sx hromosoms in plants,
and nourags studnts to liv thy an aomplish whatvr thy st
thir minds to aomplish (harlsworth 2007).
BIBLIOGRAPHY
-D.H. hrlswrth,D. 2006.v<llutionof plantrdingsystms.urrntBillog\6:
R726-R735.
2007.Q & A: Dorahharlsworth. urrtBi-lbg|7: k264_R266.
harlsworth,D., rrdB. hrlswort. 1975.Thortialgntisof Batstan
mimiry.1. Singtlor'rs
n1odls. of
.|lurl Biolog55:283-303.
Thortil
] 979a.volutiorrand rkdownof S-alllsystms. rdit43: 41-55.
t i o n s l979. odl for th volutionof Amri
distyly. NturIist1|4:
467-498 -l.T. dD.H.
htvriko srgiSrgvih(1880_1959)
Srgi SrgviIrhtvrikov (sonrtimssplld Tshtwrikoff) was a Rus-
sian ntomologist, an prt on uttrflis'and a pionr of population g-
ntis. Born into wll-duatd,profssional family' htvrikov ntrd
th Urrivrsityof osow in 1900, graduating six yars latr. H ontinud
482 betuerihou

at th univrsity as rsrh fllow, riving an advnd dgrin |909. htvrikov mphsiz

.W.omn
Aftrwards, h taught ntomology at th Highr Shool for in spis.
l\4osow from 1909 to 1919 . In 7921,,Chtvrikov aptd a rsarh posi- htvrikov is st r
tion at th Institt for primntal Biology in Anikovo, nar osow. This translatd in 1961 as ..
was run y N. K. Kol'tsov. At th institut' htvrikov ontinud his natu. th Standpoint of od
ralist ollting, spially uttrflisand moths. H also undrtook pri- mntal onpts in popr
mntal and thortial rsh into th diversity lf lrrutations in natural forgottn fourth ontril-
populations. At th sam till, h tught oth gr-rtisald iomtris. (Ronald Fishr,J. B' s. ]
Chtvrikov ws rruitd y Kol'tsov for his krrowldgof flis. Ths ani- for omining ndll
mals wr rapidly oming th standard organism in gntis rsarh. Darwinian synthsis.ln
htvrikov playd a ky rol in stalishing a Drosophil resarch group at sour of varition in t
Kol'tsov,s Institut wlrn fly stoks from Thomas Hunt organ's famous la- N{ndlian) pattrn rath
oratory first arrivd tn |922, drstood as a srisof v
htvrikov's rsarh arr fftivly ndd in 1929, at its hight. dtrmining th rat and
was arrestd, proaly for suspiious orgnizing tivity' and anishd isolation an hav an im1
from osow. H was hlpd nithr y a history of arstsfor antigovrn- Th fat of htvri
mnt disturanswhil a studnt nor y los affilitions with th organ b a r r i r s i n s i n .F r
shool of hromosomal gntisand ndlian gntis.Josph Stalin's iso- journal itslf was poorl
.Wstrn
lationist poliis purgd influnsfrom Sovit sin and ultur. short summary irr rrgli
In gntis,this purg aompanid th ris of Trofirn Lysnko and a thory had to wait for Grma,
of inhritan that nrphasizd dialtis twn rganism and nviron- howvr, irtrlatdin st
mnt and th inhritn of aquird haratristis.Chtvrikov was on of J. B. s. Haldan in Lol
hundrds of primntal gntiistsithr sakd, imprisond, ild, or Dozhansky, first appa
murdrd. did not appar until 196
In il, htverikov movd to Svrdlovsk, working in a zoo unt1l1932' his famous pap. dit
He thn movd to Vladimir to tah mathmatisin a junior ollg.In 1935 halth; it was nv pu
h am a profssor of gntis at th Univrsity of Gorky. is ativitis in htvrikov's lvidsr
Gorky ar not wll known as h dliratly kpt a low pofil undr th som of whom migt
wath of suspiious authoitis. did undrtak som srall-salrsrh f6ff_Rssovsky,B. L.
into natural sltionusing silkworms. In |948, at th ag of 68, his positiorr Thodosius Dozhnsk
was offiially trminatd in anothr Lysnko-inspird purg of gntiists sophil gntis.
.Wstrn
linkd with idas. htvrikov rmaind mntally ativ until his
dath, although in his final yrs h was hamprd y povrty and por B|LIoGRAPY
halth. His rothr Niholas ard for him until his dath t th ag of 78. dams' . 1980.Srgi1
As an volutionary and population gntiist, htvrikov argud that nat- svnthsis.In ti. Ma1,r:
ural populations maintaind a large rsrv of rssivgnti mutations' Prsptius th Un
soaking thm up lik a spong. In an volutionary ontt' ths providd UnivrsityPrss'
raw matrial for nw varity. Thy fuld mirovolution. As an primn- Chtvrikov,S. 196l. on
stndpoint lf nlodrtl
talist' htvrikov usd inrdingthniqusto rvl th tnt of divrsity
S o i t 1 0 . 51: 6 7 _ l 9 5
in rssivmutations. is idas hallngdtraditional viws that mutatiorrs
simply wr laboratory artfats' rokn forms f th normal gnti infor-
mation. Instad,populations ould onsidrdas pools in whih many al-
lausn,Jns (1891
lls swam. Th frquny of ah alll shiftd aording to dmographi
hangs, suh as migration, and through natural sltion. Along th sam Jns lausn is rgardd
lins, whn h ws studying wild populations in thir natural haitats, tionary gntisof plant

lg an advanddgr in 19o9.
Highr Shool fo $omn in
:rikov aptd a rsarh posi-
in Anikovo, nar Mosow. his
- h t v r i k o v o n t i n u dh i s n a t u -
' t h s .H a l s o u n d r t o o k p r i -
lrsity of mutations in natural
o t h g n t i sa n d i o m t r i s .
s knowldgof flis. hs ani-
organism in gntis rsarh.
a Drosophil rsarh group at
nas Hunt Morgan's famous lab-
ndd in 1.929,at its hight. H
rnizing ativity, and anishd
i s t o r o f r r s t sf o r n t i g o v r n -
ls affiliations with th Morgan
ian gntis.Josph Stalin's iso-
.rom Sovit sinand ultur.
f Trofim Lysnko and a thory
t w no r g a n i s m a n d n v i r o n -
ristis.htvrikov was on of
sakd, imprisond, ild, or
, working in a zoo unt1|19z'
atisin a junior ollg.In 1935
r s i t yo f G o r k y . H i s t i v i t i si n
ly kept a low profil undr th
lrtksom small-salrsarh
1 8 , t t h a g o f 6 8 , h i s p o s i t i o n
l o - i n s p i dp u r g f g n t i i s t s
taind mntally ativ ntil his
ramprd y povrty and poor
ntil his dath at th ag of 78.
ist, htvrikov argud that nat-
f r s s i v g n t i m u t t i o n s '
tionary ontt' ths providd
i r o v o l u t i o nA. s n x p r i m n -
s to rval th xtnt of divrsity
t r a d i t i o n v[ i w s t h t m u t t i o n s
.ms of th normal gnti infor-
lrdas pools in whih many al-
iftd aording to dmographi
Lturalsltion.Along th sam
ions in thir natural haitats.
lsn 483
htvrikov mphasizd th importan of variation within polymorphi
spls.
htvrikov is st rmmrd as th authot of a |926 thortial papr'
..on rtain Aspts of th volutionary Pross from
translatd in 1961 as
th Standpoint of odrn Gntis.'' This is a lassi prsntation of funda-
mntal onpts in population gntis' and it arnd him rognition as th
forgottn fourth ontriutor t a trio of mathmatial population gntiists
(Ronald Fishr' J. B. s. Haldan' and Swall Wright) rsponsilin th 1920s
for omining ndlism and Darwinian natural sltion into a no-
Darwinian synthsis.In his papr, htvrikov argud that (1) mutation is th
sour of variation in volution; (2) inhritan follows a partiulat (i..'
ndlian) pattrn rathr than lnding or dialtis; () gns should un-
drstood as a srisof variant allls,and th frquny of thos allls is ky in
dtrmining th rat and dirtion of volutionary hang; and (4) gographi
isolation an hav an important impat on th volution of populations.
Th fat of htvrikov's writing is a good ampl of ommuniation
barrirs in sin. Fw rad his papr whn it first appard aus th
journal itslf was poorly distriutdoutsid osow. orovr, xpt for a
short summary in rrglishpulishd in tandm' thos unal to rad Russian
had to wait for Grman, Frnh' or nglish translations.Privat translations,
howvr' irulatd in svralrsarhntrsoutsid th Sovit Union (.g.,
J. B. s. aldan in London). Sletd passags' translatd y Thodosius
Dozhansky' first appard in print only in 1959, ^ full nglish translation
did not appar until 1961. During his ford rtirmnt'htvrikov rvisd
his famous ppr' ditating orrtions nd ommntary dspite failing
halth; it was nvr pulishd.
htvrikov'swidst influnam through his tahing and his studnts,
.Wst'
som of whom migratd to th inluding N. P. Duinin, N. V. Timo-
f6ff-Rssovsky, B. L. Astaurov, D. D. Romashov, and S. M. Grshnson.
Thodosius Dozhansky rdits htvrikov with introduing him to Dro.
sophil gntis.
lBLIoGRAPHY
Adams,M. 1980.SrgiChtvrikov,th ol'tsovInstitut, and th voluttnary
synthsis.ln . ayr and \W.Provin,eds.,ThuolutiorSthsis:
Prsptius o th IJifitkllf Biolog,242-278, amridg,A: Hrvard
UnivrsityPrss.
s. 1961.On rtainasptsof th volutionary
htvrikov, rossfrm th
standpointof rodrrrgntis.Prodigsf th mri Philosophil
Soit105:167_19.5.(Trnsltion of 1926papr.) -J.,
lausn,Jns (1891_1969)
Jns lausn is rgardd as on of th pionrs in th ologial and volu-
tionary gntisof plants as wll as on of th formost alifornia botanists
484 lus

of th twntith ntury. H is spially known for his synthti and intr- sts of rlatd x1
disiplinay attmpts to undrstand th totality of prosssinvolvd in th till gldulos,
origin of plant spis at a ruial tim in th history of volutionary iology. family), lausn,
Jnsn was orn in skilstrup, Dnmark, th son of farmrs and hous of otyps (gn
uildrs. His formal shooling took pla twnth ags of 8 and 14,ter- mhanisms of s
minating whn h had to tak on th rsponsiility of managing th family ologial fator
farm. H dvlopd an intrst in gntis and for aout ight yars duatd and arfully
himslf in th asi sinswith th aid of a shoolthr' gan to tak rst th lif in
an ativ intrst in th nw sin of gntis, studying Mndlian gntis inhritan) y s
along with Darwinian volutionary thory. In 1913 h ntrdth Univrsity y ringing to r
of opnhagn to study otany, gntis, and ology, oming in onrat world. Mor nal
with som of th shining lights in Sandinavin sin that inludd ompl intrpla
hrstn Raunkir (his major profssor in otany), P. Boysn Jnsn, Wil- polyploidy (hro
hlm Johannsn, and August Kogh. At on point h vn studid physis gnral thory ol
with Nils Bohr. His arly rsarhon gntisand ology in th Violaa Throughout tl
(th pansy family) ld to a mor dtaild undrstanding of hyridization in thir rsults in a r
plants and th phnomnon known as introgrssion,whry a smallr num- Institution. Thir
r of gns r introdud through th mhanis f hyridization. His hD spiation and 1
dissrtationon th Violaa' pulishd in \926, was on f th first mono- a viabl spis(
graphs to omin systmatis'ology, and gntisin any plant group. iosystmatis) t
was susquntly appointd assistant profssor to notd gntiist jvid gentis and
.Wing
at the Royal Agriultural ollg in opnhagn, ut his ral rak Dozhansky and
am with th granting of a Rokfllr fllowship in 1927_1928 that took i Plnts (1950)
him to th Univrsity of aliforni to work with . B. Baok on th gn- th nw fild of 1
tis of th gnus rpis. Whil thr, h mt with Harvy onroe Hll' a dis- lausn's owr
tinguishd alifornia otanist, who latr invitd him to partiipat in th the uoltio o,
nw intrdisiplinary projt h was organizing with th arngi Institution somwhat rif
.W.ashington
of to undrstnd mor prisly th mans y whih nw plant sngr Lturs
spis originatd undr natural onditions using nativ alifornia flora. rally n ovr
(Hall had arlir ollaoratd with logist Frdri lmnts on suh a ut nonthlss
projt.) Apting th offr' lausn arrivd in alifornia in 191, but in- ntury voluti
stantly found himslf th ladr of th nw tam that Hall had assmld th viw that v
whn Hall did unxptdly. nomna suh as
Th nw intrdisiplinary tam at th arngi Institution of !ashington In 1951,la
that was assmldat Stanford Univrsity inludd taonomist David Kk tird in 1956 ar
and physiologist \/illiam Hisey. Until th mid-1950s th tam of lausn, his wif, Anna I
Kk, and isy (as thy am to known) dsignda srisof lassial- rsarh positio
primnts to driv gnral prinipls for undrstanding plant volution un- in trms of stu
dr ontrolld, ut natural onditions. Adapting som of th lratd mtiulously
mthods of nintnth.nturyurpan workrs suh as Gston Bonnir, tory ttooks
and arly-twntith-ntury workrs suh as Gd,t Tursson, who sought to
undrstand th pross of adaptation, lausn, Kk, and Hisy prformd BIBLIOG
a numbr of transplant xprimnts involving thr distint loations along l a u s nJ, . 1 9 5 1
altitudinal gradints in northrn alifornia. On loation Was at sa lvl at UnivrsityP
Stanford, anothr at Mathr (at aout 4,600 ft)'and yt anoth at Timr- Frnh'. S. 19
lin in th Sirr Nvada mountain rang (at aout 10,000 ft).Prforming Biogrphi

)Wn for lris synthti nd intr-
ity of prosssinvolvd in th
historyof volutionary iology.
th son of farmrs and hous
wn th ags of 8 and 14, tr-
sibility of rrrnaging th falily
l for aout ight yars duatd
lhooltlrr. gan to tak
: s ' s t u d y i n gM n d l i a n g n t i s
1913 h ntrdth Univrsity
nd logy, oming in ontat
inavian sin that inludd
.Wil-
otny), P. Boysrr Jnsn,
point h vn studid physis
;s and olgy in tlr Viola
Jrstandingof hyridization in
ssion,whry a srllrnum_
rnis of hyridization' His PhD
|26, was on of th first mono-
]ntisin ny plant group. H
sor to notd gntiist jvtnd
Jopnlragn,ut his ral rk
wship in 7927-1928 that took
rith . B. Baok on th gn-
,ith Llarvy Monro Hall, a dis-
'.itd him to partiipat in th
l g w i t h t h C r n g i I I l s t i r u t i o n
th mns y whih nw plant
using nativ lifonia flora.
Fdri lmnts on suh
l in aiifornia io 791, r.rtin-
tam that Hall had assmld
rgiInstitution of \/ashington
:ludd taonomist David Kk
i d - 1 9 5 0 st h t a m o f l a u s n ,
Jsignda srisof l:rssial-
r s t a r r d i npgl e n t v o l u t i o n u n -
pting som of th lratd
krs suh as Gston Bonnir,
Got ursson,who sought to
r, Kk, and Hisy prformd
; thr distint lotions along
ln loation ws at sa lvl at
t),and yt anothr at imr-
rout 10,000 ft).Prforming
Iuse 48.'
stsof rlatd prirntaistudison nativ Cliforni plants suh as Pottt-
till gldulos, chille spis, and th Madiina (th hayfild tarwd
family), lausn, Kk, and Hisy plord hyridization and th forration
of otyps (gntiallydistint ologial rs of plnts). Thy hartd th
mhanisms of spiation and rfind th undrstnding of th gnti and
ologial fators that ld to th origin of lant spis.Thir wll dsignd
and arfully xutd transplant studis ar gnrally rgardd as laying to
rst th lif irr th irrhritnlf aquird hartristis(o Lanrarkiarr
inhritn)y showing th distintnssof th gnotyp and phnotyp, and
by ringing to rlif th importan of plrnotpi plastiity in th plant
world. Mor narowly within plant volution, tir studis highlightd th
omplx intrplay twnlryidization,apomiis, and phnomna suh as
polyploidy (hromosorrrduling), all of whih Wr ssnrialto driving a
gnralthory of plant volution.
hroughout th 190s nd 1940s, lausn' Kk' and isy pulishd
thir rsults in a srisof important paprs and monlgraphs with th Carngi
Institution. Thir work was rad widely y othrs inrstdirr mhanisms of
spiation and y thos intrstdin applying this undrstanding to dvlping
a vial spis dfinition usful ir-rfornrulating th ..nw systmatis'' (alld
iosystmatis)that sought to intgratapprohs and undrstandingfrom
gntis and ology to traditionl taonomy. Thy influend Thodosius
Dozhansky and Gorg Ld1,rdStins.Stins's Vrititllt d uol'utio
in Plts (1950) rought otany into th volutionarysynthsisand organizd
th nw fild of plant volutionaryiology.
lausn's own synthsislf plnt volution appard in 195] as Stgesi
the uoltil of PIt Spis'An imaginativ and pinring work, it was
somwhat rif and intodutory in natur ausit ws asd on th Ms_
sngrLturs h had givrrarlir at ornll Univrsity. Th ook has gn-
rally n ovrshdowd Stins'smor omprhensivarlir synthsis,
ut nonthlss rmins on of th lassi works in th history of twntith-
ntury volutilnayiology. Among its mor imporrant ontributions wes
th viw tht volution low th lvl of th gnus is rtiultdu to ph-
nomna sulr as hyridizatilnompanid y poIyploidy.
In 1951, Clausn was appointd profssor of iology at Stanford. r_
tird in |956 and did in Plo, Alto, lifornia, in \969. H was survivd y
his wif, Anna Har-rsn,who-lh rnarrid in 192\ thy had no hildrn' In a
rsarhposition for muh of his lrf' lausn did not lav rnuh of a lgay
in trnrs of studrrts,ut his insiglrtful studis, iraginativly dsignd nd
mtiulously utd, am lassi xampls usd in narly all introdu-
tory txtooks of gntis'logy, arrd volutiorr.
|BLIoGRAPHY
lausn,J. 1951. Stgsi th ulluti<l
lf Plt S1lt..is.Ith,
NY: orrrll
Univrsityrss.
Fnh,. s. l989. Jns hristirrIusn.Ntil Acdcntof Scines
Bilgrphil emoirs 58 7 5-107.
486 op

Hagn,J. 1984.primntalists and nturalistsin twntithnturyotany'

1920-1950.Jourlof th istor <lfBkllog 1,7:249-27.
Smoovitis,. B. 1999.Jnslausn,In J. A. Garratynd M. . rns,ds.,
merinNtiolBiogrph,vol. 5, 12_13.oxford: oford UnivrsityPrss.
Stbins'G. L. 1950. Vritiod ullutil
i Plts.Nw York: olumia
Univrsitv Prss. -vB.s.

op, dward Drinkr (1840_897)

dward Drinkr op was on of th most ativ and ontrovrsial palon-
tologists of lat.nintnth-nturyAmria. H plyd a lading ro1 in
opning up th fossil ds of th \stand dsriingth nw spisdisov.
rd thr. was also a mmr of what is alld th Anrrian Shool of
No-Lamarkism. H aptdvolution ut dismissd harls Darwin,s s.
ltion thory as inadquat to plain th pross of hang. H optd in-
stad for a thory in whih evolution was dirtd long prditllins.
op was born in Philadlphia in 1840 t<la walthy Quakr family.
Was taind in sintifi agriultur, ut soon aandond this for zoology
and thn palontology. H gan y studying living fish and rptils' thn vn th st of us an
movd on to studying thir fossil rmains, vntually gnralizing his work intuitiv snsfor ro
in palontology to inlud ll th vrtratsunovrd y his pditions wnt wrong' thy rall
plsiosaur,lsmosu
in th !st. was finanially indpndnt until h lost his fortun in a
m i l l i o n y a r sa g o ) .U n f
mining fraud in 1880, aftr whih h uilt up links with th Univrsity of ody, not ralizingtha
Pnnsylvania, oming profssor of zoology thr in 1895. H did in in this pitur is misint
1897. Plat2.)
op is rmmrdfor his ittr fud with Othnil harls arsh of Yal
Univrsity. Th two lashd ovr th dsription and naming of nw fossil
spis,ah ausing th othr of hasty work and numrous mistaks (s
figur). Thr ws also fir rivalry twn thir ollting tams, who volution alng prd
somtims am to lows ovr ass to th most produtiv fossil ds. ould rsult from ada1
vntually Marsh oustd op from his onntionswith th U.S. Gologial orgnisms in a nw n
Survey, whih gratly limitd his aility t ollt in th !st. Th ..fssil us of th ldy in n
fud'' am puli in 1890 through a vitrioli hang in th pags of th pross am know
Ne Y<>rkHerld. Jan-BaptistLamrk
ope was an arly onvrt to volutionism' ut am to th thory from th lif for that allor
diffrnt akground than Drwin. A dply rligious man, h Was on. rativity in natur.
vind that God dirtd th ours of organi hang. In his 1868 artil BlBLIoGRAPY
..On th
Origin of Gnra,'' h dpitd volutilnof ah group as th un-
Bowlr,P . J, 198.h I
folding of multiple paralll lins' ah advning at a diffrnt rat through
i th Dedsrou
th sam linar squnlf stags.volution was rapitulatdin th dvl- p , . D . 1 8 6 8 .o n a l
opmnt of th modrn mryo aus th flrs ontrollirrg dvlopmnt Nturl Sisof
simply addd on nW stagsin a onsistntdirtion. Th hangswr non- o s o r n ,H . F . 1 9 3 1 . o 1
adaptiv and th ovrll pattrn was produt of th divin will. Drihr op. Prin
lialla, D. R. |999. Th
In th 1870s op movd to a mor naturalisti intrprttin that om-
SitifiFrudof tl
ind orthognsis and Lamarkism. Fors intnal to th organism n-
surd that varition was dirtd, not random, and thus drov nonadaptiv
 op 487

n twntith ntury otany,

17 249-270. l...ja.j::.:
j]]'':

.rtyand . . arns, ds.,

' oford: oforl Llivrsity Prss.
P/lllrs.Nw York: oltrrnbi
-B.s.

)
'ltivand ontfovrsil palon-
r. H playl :r lading rol irr
:sriingth nw spisdisov-
s alld th Arr-rrianShool of
:dismissdharls Darwin's s-
)rossof hang. H optd in-
tdalong prditalelins'
wzrlrhQuakr family. coP oN Fosstl litl o lv J&sY

n aandorrd this for zoo|og

ng living fish and rptils' thn
vntually gnralizinghis work
:s unovrd his pditions
r t u n t i l h | o s th i s f o r t u n i n
up links rvith th Univrsity of
lgy thr in 1895. H did in
l othnil harls Marsh of Yal
tion nd namir-rgof nw fossil
'rk and numrous mistaks (s
'n thir ollting tams, who
l most produtiv fossil ds.
tionswith th U.S. Gologial
ollt in th $7st.Th ..fossil
lli hang irr th pags of th
L,ut am ro th tlrory from
,ly rligious man, h Was on.
Lnihang. In his 1868 rtil
lution of ah group as th un-
.ing at a diffrnt rat through
Lwas rapitulatdin th dvl-
fors ontrolling dvlopmnt
irtion.Th hangsWr non-
t of th divin will.
:alistiintrprtation that om-
s i n t n Ir o t h o r g a n i s n r n -
m, and thus dror, nonadaptiv
vn th st of us an mak mistaks.dward Dinkr op had a rilliant
intuitiv snsfor ronstrutingth monstrsof th p21st. But whn things
WntWrong' thy rally wnt Wrong.This pitur shows his onPtionof th
plsiosaur,lsmosurusplturus,from th lat rtaous(aout 70
million yars go)' Uflrtuntl,lp Put th had at th wrong nd of th
ody, nlt ralizingtht th ral r-rinrl had trmndouslylong nk' whih
i n t h i s p i t u r i s m i s i n t r p r t ds a t r m n d o u s l yl o n g t a i l . ( F r m o p 1 8 6 8 ,
P l a t 2 . )
volution along pdtrmindlins. A similar pross of paralll volution
ould rsult from adaptiv volution ontrolld y th haits hosn by th
organisms in a nw nvironmnt-haratristis aquird y th onsisrnt
us of th dy in a nw Way wr passdon to Sudinggnrations.This
pross am known as Lamarkism, although op WaS not awar of
Jan-BaptistLrark's work whn h dvlopdhis ida. op argud that
th lif for thirt llwd organisms to adapt ws a manifstation of God's
rtivity in rratu.
BIBLIOGRAPHY
Bowlr,P. J. 1983. Th clips of Dru,'iism:Ati-Drluiiuolutio Thoris
i th Ddsroud i 900. Bltimor:Johns Hopkins UnivrsityPrss.
lp,. D. 1868.On a nw largnaliosaur,Prodings of tbcdemof
N trISis of P hildelpbi Z'
Osorn, . I...l931. lp:str Nturlist;h Life d Writigsoi drd
Drihr op. Printon,NJ: PrintnUnivrsityPrss.
.Walla,
D. R. 1999. The Boehutrs'Ruge:Diost'trs,Greed d the Grtst
-P'l.B,
SitifiFrud of the Gildd g. Boston:Houghton ifflin.
488 roltl

surprisingly larg
row, JamsFranklin (.1'916)
lthal mutations. T}
Ovr th ours of a arr that spans ight dads,Jams Franklin row has ous ffts ar not m
mad svral sminal ontriutions to oth thortial (mathmatial) and - dividuals haror a
primntal population gntis. Althugh his primary rsarh organisms rssiv natur of 1
hav n fruit flis of th gnus Drosophil' row has also mad important sion. row ontinut
ontriutions to th study of humans and agriultural rops. Although row humans.
has workd on prolms ranging from th gntis of DDT rslstn to - Crow was th r
amining gns that violat ]Vlndl'slaw of fair miosis to dvloping thory Kimura dvlopd tJ
aout th bnfits of sual rprodution, his most prominnt ontriutions that th vast majorit
rvolv around th rol of mutations in volution. with rsptto sl
Born in January 1916 in ollgvill, Pnnsylvania, row grW up mainly asily gnratsman
.liihita,
in Kansas, whr h rivd his undrgraduat dgr from whthr and what f<
Frinds Univrsity, a Quakr shool. I 197, row wnt to th Univrsity of Laying th foundati
xas at Austin for graduat work, initially to study with prominnt gn- Crow to dtrmin
tiist Hrmann Josph (H. J.) ullr, who would latr win th Nol Priz this variation was tl
for his studis of X-ray mutagnsis in Drosopbila. Mullr, howvr, was to variation and ran
about to migrat to th Sovit Union' so John Thomas (J. T.) Pattrson . row dvlopd an
am row's major advisr. Just prior to th Unitd Stats' ntry into.!7orld that vry mutatior
\WarII tn 1941, row landd a faulty position at Dartmouth Collg, whr modl, th xpt
.W.artim
h stayd until 1948. and th drafting of his ollagus at Dart- was hetrozygousa
mouth ld row to add ourss as divrs as parasitology, navigation, and quals four tims th
mryology to his initial tahing rsponsiilitis of gnral zoology and g- mutation rat. rol
'Wisonsin'
ntis. In 1948, row movd to th Univrsity of whr h has r- ook, z lntrodut
maind for si dads. y gnrationsof st
Mutations ar not an unmid blssing.Although mutations ar th ulti- apparan in t970
mat sorrr of vriation and ar thus ndd for volution to prod, dl- In addition to hir
trious mutations far outnumr nfiial mutations. row and his studnts mad svral othr
usd th gnti thniqus availal in Drosophil melnogstr to masur studnt undr th t
th input of nw lthal mutations apparing on singl hromosom ah Austin, row was
gnration.From this valu, thy alulatd that th lthal mution rat pr isolating arrirs
gnti lous (gene)pr gnration was around thr pr million. utations study th gnti
nd not hav lthal ffts to b dltrious; in fat, row and his ollagus mappd largly to
hav shown tht nonlthal, dttaldltriousmutations aris at last 10 D. ldrihi-2.
tims mor oftn than lthal mutations. Rnt studis suggst that th rat row has also h
of mildly dltriousmutations is vn highr than thos mutations that hav Mndlism. Invest
ffts that an masurd in th laboratory. qual sgrgationh
Crow's group and othrs hav shown that most dItriousmutations ar part a fair gam.
almost ompltly rssiv;that is, th ffts thy hav in htrozygots mioti driv syst
are far smallr than th ffts thy hav in homozygots. Baus muta- Dr>sophilmelo
tions ar mostly rssiv,thy ar largly shltrdfrom sltion and thus at studnt, Yuih
ar not asily wdd out from th population. Ths dltrious muta- row, his studnts
tions, whther thir ffts ar lthl or mild, drasth man fitnss of gntis,and volu
th population that harors thm. ullr alld this rdution of fitnssth row's influn
.With
gnti load of th population. ullr and othrs, row invstigatd tion to th sors
the tnt of th gnti lod in oth Drosophil and humans, finding it to dirtly trained, rr

cs,Jams Franklin row has
:tial(mathmatial)and -
p r i m r r s a r ho r g a n i s r n s
:ow has lso mad important
rlturalrops. Although row
tis of DDT rsistan to -
miosisto dvloping thory
l o s t p r o m i n n t n t r i u t i o n s
n
lvania'row grW uP mainly
undrgraduat dgr from
row Wnt to th Univrsity of
study with prominnt gn.
rld latr wirr th Nol Priz
phil. u||er, howvr, ws
Thomas (J. T') Pttson .
.!orld
ritd Stats' ntry into
at Dartmuth ollg, whr
rg of his lllagust Drt-
larasitology, navigation, and
s of gnralzoology and g-
i Wisonsin,whr h has r-
rougmuttions ar tlr ulti-
lr volutiorrto prod, dl-
ttions.row and his studnts
bil mlogstrto msure
n a singl hromosom ah
t th lthal mutation rat pr
r h r p r m i l l i o n . u t t i o n s
[at' row nd his ollagus
rs mutationsaris at last 10
stdis suggstthat the rat
an thos mutations that hav
lst dltriousmutations ar
; thy hav in htrozygots
omozygots.Baus muta.
rdfrom sltionand thus
ln. Ths dltrious muta-
jrasth man fitnss of
I this rdution of fitnssth
d othrs,ow invstigatd
/ a n d h u m a n s ,f i n d i n g i t r o
ro 489
surprisingly lrg. Th vrg human harors th quivalnt of svral
lthal nrutatiorrs.Ths lnutations r rstly rssiv,nd thir dltri-
ous fftsar not manifst unlss individuals inrd aus unrltd in_
dividuals lraror diffrnt st of rssiv dltrious mutations. Th
rssivnatur of ths mutations is tlrr.rsth sis of irrdingdprs-
sin. row ontinus to hav an intrst in mutations and thir ffts in
hurrans.
row was th mntor and longtim ollaorator of otoo Kimura.
Kimura dvlplth nutrl thory of molular volution, whih argus
that th vast majtlrity of rnolr:lrvriants ar ithr dltriousor nutral
with rsptto sltion.An impotant fatur of th nutral thory is tht it
sily gnatsmny tstirlprditions that n allow iologists to dtt
whthr and what forms of sltionar Opratingat prtiuiar gnti loi.
Laying th foundation for th nutral thory was work that Kimur did with
row tl dtrmin how muh vriation would ptd, providd that
this variiltion was th rsult of a alan twn nutral mutations addirrg
t variatin and random gntidift roding variation. |n \964, Kimura and
row dvlopd a importilnt mathmatil modl in whih thy assumd
that vry mutation that ariss is uniqu. In this so-alld infirrit allls
rodl' th ptd htrozygosity (th proportion of th population that
rvas htrozygotlst a sit) of rrutr:alallls r.rndrmllttion/drift aln
quals flur tims th produt of tlr fftivpopulation siz nd th nutral
n.lutationrat. ow and Kimura also pulishd a highly influntial tt-
lok, ll [trodtio tl Pltil Getics Thor, whih hs n usd
y gnrationsof studnts and pratitionrs of popultion gntissin its
a p p r a n i n 1 9 7 0 ,
In addition to his work on th ol of mutation in volution, row lras
mad svrl othr ontiutions to volutionary gntis. As a graduat
studnt r-lndrth tutlag of J. T. Pattson t th Univsity of Ts t
Austin' row was an arly invstigator of th gnti asis of rprodutiv
isolating arrirs twn Drosopbil spis.row was among th first to
study rh gnti asis f hyrid fmal inviaility, firrding tht this trait
mappd largly to th hromosom in hyrids twn D. mllri and
D. Idrihi-2.
row has als had n tiv inrstin gntionflit and th volution of
ndlism. Invstigationsof gnti systms tht violat Mndl's law of
qual sggtionhv hlpd our undrstanding of why Mndlism is in
part a fir gam. Th st knowr.rand st haratrizd of ths so-alld
nrioti driv systms,sgrgationdistortion on th sond hromosom of
Drosolbil mlogstr, was dis<lvrdin row lortory y his gradu-
at studnt, Yuihiro Hirazumi, during th 1950s. In susqunt yars,
row, his studnts,and ollagushav workd tln th gntis' molular
gntis'and volution of th systm.
row's influnon th fild as a ther and mntor is immns. In addi-
tion to th sors f graduat studnts and postdotorl fllows that h has
dirtly traind, row hs inf]undth fild with his various ttooks and
490 rustc.ts
..Histlrial Insts' mollusks, a
historial ssays. Along with \/illiam Dov, row has ditd h
Persptivs''stion r Getis sin 1987. (As oi mid.2008, ow as r of spis, ut rr
writtn about 250 atils.)A longtim mmr of th National Aadmy of divrs animal group l
Sins(NAS), rw has haird many ommitts fr th NAS, inluding adquatly onvy th
on xamining fornsi uss of DNA fingrprinting.H has srvdas hir of known forms lik lo
th Dpartmnt of Gntis and ating dan of th diirl Shool t th fmiliar amphipods'
..sd shrimP''), to nl
Univrsity of Wisonsin at adisn, as wll as prsidnt f th Gntis So-
ityof Ami. ompris only a smal
sPis.Thy r pri
BIBLIoGRAPY or moist trrstrialh
row, J. P. 1979. Gnsthat violatMndl's r'ils.Sitifimrin 240, no. 2.. li, pill ugs, sow u;
\4_1'46. th roughly 150 millir
1987.Populationgntishistory:A prsonaliew.tl Ruiof G- Paifi annually migrt
tis27: I-22. tims that nur of
1993.Mutation'lanfitnss'and gnti|oad,ford Surueof
groups (.g., larg
uolutitlr Biollg9: 342.
1997.Tlrighspontnous mutionrat:ls it halthrisk?Prodigs shimp) hav adapt
oi t,|itili dmof SisU5 9,l: E3ii0-386. ths rustaansun(
row, J. F., and M. Kimura. 1970. ltro,llttilto PopultilGtisThor. nystd in a hard p
Nw York: Hrpr & Rw. drying, only to mrg
Intrvirv with Protssor JamsCrow. 2006.Blossa1,.s 2[i' nl.6: 660_678|doi
rustaans ang
10 . 1 0 0 2 / b i s . 2 0 4 2 6 ] .
Kimura,M., and J. F. row. 1964,The numrof alllsthatan b maintaindin and nthi spisli
a finit1.lpultin. Gtis49 725-78. ns spidr rabs an
Simmons,. J., and.}.F. rolv' 7977,utittionstftingfitnssin Drosopbil and lostrs hav
populations.llul Rui,of )tis11 49_78' -N../. tiipatd that many
th 68,000 spisu
vry rmot haitats
rustaans sa-and nw findin1
rustaans ar a strphylum of arthropods' as ar uniramians (insets,rnilli- spis. Aording tt
pds, and ntipds) nd hlirats (spidrs' nrites, tiks, arrd hlssh hav n dsrid
ras), lthough' i som lassifiation shms,arfhropods ar onsidrd to usd lassifition'tl
polyphylti (i..,not dstrddfrom a singl ommon anstor),and rus- tirly nw lass f r
..whirt is av in th Bhams
taans r lvatd to phylum sttus.Th smingly simpl qustion'
a rustaan?''is suprisinglydiffiultto answr' and this is atlywhatmaks Tlrmosatlaa,N
this group of animls so intriguing. As arthropods, rustaans ar asilly limtrs) of praari
sgmntdanimals with jointd appndagsand a stiff yt flil xosklton th dp sa just sin
mad of hitin that must priodially slrd, or roltd, to aommodat ordrs mpris only
mtamorphosis and growth. Yt rlativly fw fatr"rrs distinguish rustaans disjunt distriution
from othr arthropods-a uniqu larval form alld a nauplius and a h:rd r- splaogriphaans
gion omprising fiv body sgmntson whih ar fourrd two pirs of ntnna in Brazil, South Afrk
(usually iramous, or two-brnhd), :r pair of m:rndibular appndags' and populations rprs
two pais of spializdfding appndagsalld maxilla-ut vn thsfw widsprad; hangs
faturs ar not ommon to all rustaans.Lik som othr rthropods, th trind in, nd latr
rustaan multisgmentd tunk postrior to tlr had, or phalon, may or rustans fd i
may not b furthr rgionalizlinto distint stions of fusd sgmnts'suh as mals. Som ar filt
..tail''), thir watry nviro
a thorx (or phalothora) and adomn (.g.,a lostr's and ap.
pndagsan vary grtly in ntrmr and dgrof spializatin. prd2ltors.Th ltt
 tstcns 49 1

row has ditd th ..Histoial Insts'mollusks, and hliratsd rustaanswith rgard to num-
. (As of mid- 2008, row has r of spis,ut rustaansar unquivoally th rnost morphologially
r of t National Aadmy of divsanimal group on arth. h fw, vry gnral faturslistd aov in-
nitrs fr th NAS, inluding adquatly onvy th rmrkal divrsity of a grrr-lpthat inluds wll.
rting. has srvd s hair of known forms lik lostrs, shrimp, as, and arnls, as wll as lss
of th dial Shool at th familiar amphipods, isopods' fairy shrimp' watr flas, and ostraods (or
..sd
s prsilntof th Gntis So- shrimP''), to nam ut a fw (s figur). In ft, th familiar forms
ompris only snrll propt>rtionof th roughly 68,000 known rustaan
spis.Thy ar pimarily a marin grop ut som also oup frshwatr
or moist trrstrialhaitats ('g.,trrstrialisopods, ommonly alld wood
. Sintifimri 74, no.2: li, pill ugs, sow ugs' or ..roly-polis,''and trstrialras). Intstingly,
th oughly 150 rnillion rd trrstrialrs of hristmas Island in th Indo-
nal virv.l Rt,'itulf G. aifi annually migrat from inland forsts to th s:l to spawn; mny mor
tims that numr of juvrrilsmigrat in th rvrsdirtion. Sonr aquati
Ioad. Oford Suru of
groups (.g., larg ranhiopods rfrrd to as fairy, tadpol, nd lam
: ls it a hlth risk? Prodigs shrimp) hav adptd to living in tmporv ponds or ..phmralpools'';
8380-8386. ths rustaansundrgo urapid iif yl and pr<ldudorrant mryos
n to tllultirlCtis Thor. nystd in a had pottiv as that n withstand tndd priods of
drying, only to mrgand ontinu thir dvlopmntwhn a pond rfills.
ss28,no. 6: 660-678 [doi:
Crustaansrang in siz from lss than a millimtr (minut zooplankton
lllstht n nr:rintindirr and nthi spis living within marin sdirnents)to th Yry larg. Japa-
ns spidr ras an hav [g spans ding 3 mtrs, or roughly 9 feet,
tfftingfitnss in Drosopil and lostrshav bn aught that d 20 kilograms (40 pounds). It is an-
9-78 -N../.
tiipatd that many rustaansawait disovry (at last as many spis as
th 68,000 spisurrntly known), spilllyupon ftrrthrplor:rtion of
vry rmot hiritats lik frshwatr nl marin av systms and th dp
sa-and nw findings ar not nssarily ptd to onfind to mr
s ar uiramians (insts,rnilli. spis. Aordirrg t artin and Davis (2001), almost 200 nw familis
rs, mits, tiks, and lrorssho hav n dsibd in th 20 vars that lrv lpsd sin th lst widly
s' arthopods ar Onsidrd to usd lassifiation,that of Bowman arrd Abl (1982). Rmarkly' an n-
l ommon anstor).and rus- tirly nw lass of rustaans'th Rmipdia' was disovrd in a martn
ringlysimpl qustion, ...Whatis av in th Bahrnas as rntly as 1981, and nw ordrs (Splaogiphaa,
.,and this is atlywhat mks Thrmosbanaa'Mitaa, ilnd Bohusaa)of minut forms (0'5 to 2 mil-
pods' rustaans:lr asiall), limtrs)of prarid rustnshav n disovrd in rmot avs and
d a stiff yt flil osklton the dp sa just sin th mid-1950s. Not surprisingly,som of tlrsnwr
J, or moltd, to aommodat ordrs ompris only a fw' rirr spis.\what is surprising is th rmarkaly
ftursdistinguish rustaans disjunt distriution of som of ths nrvly disovrd groups, sulr as th
:alld nauplius and had r- splogriplrirans-only our spisr known, disovdin singl avs
Lrfound two pairs of antnna in Brazil' South Afria, and nrthwstrn Australia! It is susptdthat ths
'f mandibular appndags, and populations psntrlis of an anint marin group that was on mor
d mailla-but vn ths fw widsprad;hangs in s lvls might hav ld to individuals tllrringn-
ik som othr artlrropods, th trirrdin, and ltr adapting to, grounlwtand a'',haitats.
th had' or phlon' may or ustaans td and rpr<lduin alrnost vry \4l1lyimginal for ani-
tions of fusd sgmnts,suh as mals. Som ar filtr fdrs and siv atria or tiny zooplankton from
.g., a lostr's..tail''), and ap- thi Watry nvironmnt' othrs savng for thir food, nd lany a
of soilization. prdtos. h lattr mer1,h:rr,s()m rInkl adpttions, srrh as th
492 rsts
..lalv'' .
*:;;;.Sflffi'v
r1,-|],\.
modifid
smsh mllusks
rustilan hs a t
) ('.,' gthr, produ
'r @I1
d -_--,' shrimp''). Th sol

.g
stun or vn kill sr
also notl for >
mits. Nutnrltts
spollg lll1d f()rm
phnornllon of
ntly mmtt
Ss ar usu
known) nd su
as watr fl:rs,wil
as parthnogns
vlop frorn unfrti
nvironmnt o s<
swimmir-rglarva 1
morphosirrg intl t
spis, sh as f
mrg (dirtdv
Th rlv rrr
liotl ,virrsgo) sa
xhiitillg n inr
this ..plosin''O
tonishingi bif l
phyla hav long si
an quivalrlt[rllr
a topi <lfd:rt
plosion tlf ody p
haps gratr oxy
mtaboli rtsan
gn.dupliation t
what wr tl-rnr
s
progrms <lfanim
Rprsntativs of som si rr-rstantyps. () opoda; () ostrod; nonlthl l-najor:
() nostraa; (d) Notostraa; () ladoa; (f) Lptostraa; (g) Arraspidaa; mnt onstitutdi
(h) irripdi; (i) Stomirtl1.lod;
(j) ysida; (k) Tanaidaa; (l) lslpo1a; filld by adapt
(m) Amphipoda; (n) uplr;rusiair;(l)Dndrrnhiat; (p) arida;
an adptiv radiat
(q) Astaid; (r) Anomur; (s) Ahlata; (t) Thalassinid; (u) BrahyLrra;
Fossil vidnr
(v) Polyhlida. (From Polr 2004, Figur 2.)
suphyl (or arth
taly, on oi th ft
vi]nthat rus
this tirlr' Th las
or ..sa r-nonky
som of th ail

s. (a) oppoda; () Ostralda;
f) Leptostraa; (g) Anaspidaa;
<)Tanaidaa; (l) Isopod;
l n h i t (l p ) r i d
halassinid;(u) Brahyu;
49
modifid ..law,, of a stomtopod mantis shrimp tht allows .^:':
smash mollusks or spar fish with grat for and spd. Anothr typ of
rustaanhas a rndifid lw with pinrs that' whn quikly rought to'
gthr, produ loud snapping nois (hn thir nam, ..snapping
shrimp''). Th sond is assoiatd with a ..shok wav'' strong nough to
stun or vn kill small pry, suh as fish. Som spisof snapping shrimp ar
alslnotal for hiiting soia1havior mor typial of ants, es,and tr-
mits. Numrous offspring of a singl fmal ( ..qun'') will oist in a
spong and form asts to a for th ylung and prott th olony. This
phnomnon of usoiality in snapping shrimp ws dsrid rltivly r-
ntly y mmtt Doff (996) and is a first among rustaans.
Ss ar usually sparat (although som hrmaphroditi spis ar
known) and sual rprodution is th typial mod. Som rustaans, suh
as watr flas, will asionally rprodu asxually via a mhanism known
as parthnognsis,whry fmals will prldu young (also fmal) that d-
vlop from unfrtilizd ggs. rustaan ggs may rlasd into th aquati
nvironmnt or so.tims roodd, ut th young usually mrg as fr-
swimming larva that will undrgo a sris of molts and stags for mta-
morphosing into th adult form (indirt dvlopmnt). Th fmals of fwr
spis, suh as frshwatr ras, rood thir ggs until miniatur dults
mrg (dirt dvlopmnt)' molting as thy grow largr.
Th arly amrin priod of gologial history (approimatly 550 mil-
lion yars ago) saw th mrgnof numrous multillular animal phyla
hiitirrgan inrdil divrsity of ody plans. Palontologistsnow liv
this ..plosion'' of lif forms ourrd ovr only 5 to 10 million yars (an as-
tonishingly rif amount of ti"l,volutionarily spaking). any of ths
phyla hv long sin gon tint, and nvr again in th arth's history has
an quivalnt urst of volutionary innovation ourrd. Th rasrnrmalns
a topi of dat and spulation.Possilontriutors to th amrian -
plosion of ody plans inlud hangs in th arth's gohmistry (.g.'pr-
haps gratr oygn availaility favord marin organisms with highr
mtaoli rts and lrgr sizs)and suddn, larg-salgntihangs (.g.,
gn-dupliation vnts and a rshuffling of gn rgulatory pathways in
what wr thn rlativly simpl organisms).Thraftr' prhaps th gnti
programs of animals had rahda lvl of omplity that prludd furthr
nonlthal major hangs. Anothr viw is that th pr-amrian envlron-
mnt onstitutdn ologial taula ras of mpty nihs simply waiting to
filld y adapting organisms. Thn' as now, suh nvironmnts Dromot
;
an adaptiv radiation lf spis.
Fossil vidnonfirms that rustaansWr on of numrous arthropod
suphyla (o arthropodous phyla) prsnt in th amrian and, mor no-
taly' on of th fw that hav survivd to this day. vn mor striking is th
vidnthat rustanshad alrady divrsifidinto svrlmain lineagsy
this tim. Th lasssOstraoda (sdshrimp), Branhiopoda (rin shrimp,
or ..s monkys''; fairy, tadpol, and lam shrimp; and watr flas), and
som lf th aillopoda (spifiallythostraans
[arnalsIand wormlik
494 rustns

parasiti pntastoms)hav amrian rprsntativs.Shrimplik fossils of rustan) that proa

th lass alostra (omprising th familir shrimps, lostrs,urndras) rltivl rrndiffrrrtiat
ar known from th Dvonian (apprlximatly400 million 1'arsago). R- o s t s g m n t s o r p i
..living
markaly' prhaps th oldst surviving animal spisis a rustaan of whih rmains ontr.
th rdpol shrimp Triops criflrlis.This spisapprs tody irs
f<rssil,'' basi ground pln, ut r
it did 22 million yelrsago. Tlr rmiringrustaan lasss (Rmipdia, holdovr of a pimitiv
phaloarida), whil primitiv in apparn'ar ptlorly rprsntdas fos- from a more drivd an
sils. Th virtually simultanuspparanin th fossil rord of most of th rustans oftn rsult
rustaan lasss maks ronstrutitln of tlr vttlutionar rlationships tint rgions, alld tagr
among thm from fossil vidn impossil; sintists annot dtrmin and adomn) and dv.
whih of th major rustangroups or lasssaros first' sond, or third, tions. But mong taf
or whih major grup spifillygav ris to anothr. th urstof morphlog
wo faturs of rustaans-thir anint origin and thir wildring ously rsrvd for th o
arra of morphologial divrsity (gratr than that of th mor spios omntal iology sugg
insts)-add to th diffiulty ot infrring thir phylogny (i..' thir volr'r- surviving ling, most
tionary intrrlatiorrslrips). Th goal of volutionaly ronstutionis tl d- a b i l i t y t o d v l o pd i f i r
iphr th phylognti pattrn to dtrmin whilr group of organisms is omplmnt of gns).(
anstralto (that is, gav ris to) anothr group or groups of organisms.Tra- volutionary tiks. Va
ditionally, s,vstmatistshv lookl for pattrnsof shd morplrologial fa- hang rsultd irr nw
turs among groups of organisms to rollstrut a phylognti (family) tr of ustaan ody pla
that dpits th groups' vollltionary rlationships.Baus th major rus. th appriation of th
taan linagsdivrgd from ommon anstor so Iong ago' linagshv prolm this divrsity:
had a long tim during whih to volv and tl divg from ah othr mor- lationships within this g
phologially. Disrning th morphologil fatursthat xtnt mmrs of D r i . r m t i a n d r l t i
eah linag still sha, and that would indiat n anint ommon ans- homoti gns.oftn l
ty' is thrfor diffiult. For ampl, rr-rals wr long thouglrt to rguiatory gns invoi\
mollusks rthr than rustaans(or vn rthropods!).Thir highIy drivd homoti gns' th tir
and spializd adult morphology lis any affinity with rust;rans,and it ryoni rgions that th.
wirs only whn th larvl stgsof arnals,rosrvd in tlr 1830s tht lls wirhin a dvlopin
thy wr rognizd as rustaans.In fat, harls Darwin himslf pro- tial orgaolzation of th
vidd th first omprhnsivrviw of anal taxonomy' and in doing so for ampl, ths imp
h am mor assurd of his yt-unpulislrdidas on volution. Phylog. ody sgmntation(.g
nti ronstrution is furth amprd y parasiti rustaan taa (.g., rax' o adorn) an
pntastomid ..Worms''' and asothoraidans, rhizophalans, and tanttrlo- mandils, swimming l
arids) that hiit gratly modifid morphologis that ar diffiult to fit into nor mutation in on of
..natural
a lssifiati<ln''(i..' orr that fltsvlutionay rltdnss)of with a dramatially no
th rustaa. ody sgmntsof an t
For all ths rasons,littl onsnsushas arisn aout th volutionary r- pa'' would radially e
lationships mong th lsssof rustaans(.g.,on th T of Lif wsit, sgmntsof th rusta
,.rttstaa''
http://www.tolw.org,th link to lads t an unrsolvd various pattrns is sus;
phylogntitr for rustansas a whol and unrsolvdtrsvn within gis osrvd in rusta
most of th rnajor rustan linags).Atrthoritisvn disagr aout th sgmntsmay also fus
numr of taxonomi lasss l-rdthir lI-tstitul-lts, s is fltdy th xampl, proaly a
numr of diffrnt lssifitionshmsurrntly il-lus. in diffrnt spis of 1
\What aunts for rustaans' rmarkl divrsity of ody plans? organization sn in lot
..ster''
l-ik oth rthrpods, rllstaallsar drir,dfr<lmn anstor (tlr thorx' and dominal

ntativs,Shrirplik fossils of
L rs h r i m p s 'i o s t r s ' n d r a s )
y 400 million yrs ago). R-
I spisis rustaan ..living
;.This spisappars today as
:rustaanlsss (Rmipdia'
, ar poorlv rprsntdas fos_
th fossil rord of rost of th
:h volutionary rltionships
. , ;s i n t i s t s n n o t d t r m i n
s aosfirst, sond, or thid,
anoth'
origin nd tirir wildring
rn har of th nlr spios
ir phylogny (i..' thir volu-
ionary ronstrutionis to d-
w i h g r o u p t l f o r g r l i s m si s
D or groups of organisms.Tra-
s of shrdmrphologial fa-
t phylognti(family) tre
;hips.Bausth rnajor rus-
stoso long ago, linagshav
divrgfrom ah othr mor-
rtr.rrs tht tant rr-rmrs of
.t an anlnt olmon ans-
ls wr long thought to
ropods!).Thir highlv drivd
[finltvwith rustans'and it
re osvdin th 1 830s that
hls Darwir-rhimslf pro-
: I t x < l n o m y'n d i n d o i n g s o
l idas on volution' Phylog-
rasiti rustaantaa (.g.'
rhizophalns,anl tantulo-
isthat ar diffir-rltto fit into
s evlutionaryrlatdnss)of
;naout th volutionary r-
].' on th Tr of Lif wsit,
a'' lds to an unrsolvd
l unrsolvdtrs\,nwithin
itisvn disagraout th
'tunts' s is rfltd y th
]ntlyin us.
l divrsity of ody plans?
from n anstor (tlr ..stm''
rustces 49.'
ustaan)tht proaly had flil oskltonnasing numrous and
rlativly un{iftrntiatdsgmnts(a ondition trmd srial homonomy).
ost sgmntsr paird, jointd' and fairly sirnplappndags,tlr natu
of whih rmains ontrovrsial.Th rustaanlass Rmipdia rfltsthis
asi ground pln, ut viws diff ovr whthr th rmipd ody plan is a
holdovr of a prirnitiv ondition or a rvrsion to th primitiv ondition
fonr a mor drivd anstor.Sr"rsqunt evolutionary hang amorrg arly
rustaansoftrrsultdin th Ompartmnt:rliztionof sgmntsinto dis-
tint rgions,alld tagmata (singr"rlr,
tagma; xanrplsar th hd, thora,
and adomn) and dvlopmnt of appndgs modifid for diffrnt fun-
tions. But among xtant arthropod suphyla' rustaans alon parallld
th burst of morphologial innovation obsrvd in tl-rarly amrin prvi-
ously srvdtor th origin of disprat phyla. Rnt findings from dvl-
pmntal iology suggst tht arly rustaans' mor so tharr any othr
surviving linag, most likly possssdgrat dvlopmntl plastiity (th
aility to dvlop diffrntly undr diffrnt onditions' vn from th sam
ornplmntof gns).rustaansrral to draw from an amazing ag of
volutionry tiks' Vrious pr()ssspromotin. dramti morplrologial
hang rsultd in nw linags nd ontriutd to th astounding divrsity
<lfrustaan ody plans today. Undrstanding ths prosssan add to
th appriation of th singular sop of rustaan divrsifiation and th
prlm this divsityratsdurirlg attmptsto ronstrutvolutionary r-
latilnshipswithin tlris group'
Drmati and rlativly suddn voltionry hang an b afftd y
hlmoti gns. Oftn rfrrd to as Iox gns in animals, ths ar mastr
rgulatory gns involvd in an organism's dvlopmnt. Th rrumr of
homoti gns' th timing of thir prssion' nd th othr gns and m.
ryoni rgions tlrat thy in turn afft dirt th fat of various groups of
lls within a dvloping mryo irnd ultimtly ontol th idntiry,and spa-
tial organizatiorrof th various ody parts sn in th adult. In rustaans,
for ampl, ths important rgulatory gns rgult th dvlopmnt of
ody sgmntati()n(.8.'th numr of sgmntsthat mak up a had' tho-
a' or adomn) and th numr and kind lf appndags (ntnna,
nrandils'swimnring lgs, t.) tht dvlop on tos sgmnts.vrl a mi-
nor mutation in on of an organism's homoti gns an rsult in n aduit
with a dramatizrllynovl ody pln. For arpl, th dorsal fusin of th
ody sgmntsof an anstral rustaaninto som sort of shild or ,,ara-
pa'' would radially ltr th appaanof its dsndant.Partiularly for
sgmntsof th ustanhad rgion, trmd th phalon, su ftrsion in
vaious pattrns is susptdto aollnt for th divrs arapa mrpholo_
gis osrvd in rustaantaa. Rlativly undiffrntiatdanstral trunk
sgmnts may also fus into tagmata. Variations i Ho gn prssion, for
xampl, proably aount for th diffrnt numrs of trunk sgmntssn
in diffrnt spisof putativly pimitiv rmipds,and also for th ody
o;anizationsnin lostrs:for ampl, sgmntsfr"rsing into distint had,
thor, and adominal tagmat,irnd sgmrrtsof th thorax zrringwalking
496 (]rustcs

appndags(th first pair of whih ar modifid lws or ..hlipds''sn in today ar th volurio

nany of th ttr-known rustaans),wlrras thos of th adomn ( plays a onfusing oml
lostr's tail) ar appndags modifid for swimming. Variation in Ho n a t i o n t h t m k sd i s
gn prssion likly aounts for arnals laking adominal sgmnts To irumvnt ths
ntirly (on of th rasons thy wr originally ovrlookd as rusraans). than th traditional
rnparativ studis of gn prssion pattrns for th Hox-lik DistlJess pars th larvl morph
and rrgraild dvloprnntal rgulator gns hv sigrrifiantly;rdvand tions that arly dvl
our undrstnding of th arly morphognsis of rustaarr lims and s p i a l i z t i o n ss n i n
sgmntation. mor auratly rflt
Htrohrony is anothr pross that affets th volution of morphology, a o v , i l I u s t r t tsh i s
nd on that has Iikly ontriutd to th divrsity f rustaandy plans. unit groups suh as
Bror.rghton y mutations in th grrsontrolling th timing of dvloprnn- phala within th bar
tal vnts witlrin an organisr,th htrohroni pross may quikly rsult siti Asothoraida as t
in th apparan of novl morphologil diffrnsbtwn anstorsand n usd in attmpts
dsndants.vn slight hangs in th rlativ dvlopmntal and growth primitiv as wll as m(
rats of diffrnt body parts in an mryo an rsult in a dramatially altrd f -i|v i n g l i r h o t r o p
adr"rit.For ampl, a mutation that tnds th priod during whih a gn pothss aout th .rf
tht rgulatsntnnet1 growth is ativ n rsult irr arr offspring with antn- and th vlution of th
na many tims longr than thos of its parnts. If th mutation is hrital, Anothr approah, r
thn susquntoffspring will also grow long antnna.In som rustaans plin of phylogntis
(e.g., anostraansJsuh as fairy shrimp and rin shrimp, also alld sa shard y diffrnttypt
monkys), antnn ar oftn rquird for sussfLrl opulation. Mals with nzym ytohronro
long ntnna may al to mat 1Orsussfullytlran ons with shortr vry rustaan.Th
i1ntnna'or thy might oniy al to mat with partiular v;rriantamong sPis of rustaans
th fmals in a population. If popultion o-lsrprodutivlyisolatd, diffring dgrs of sin
its gns ar no longr mid ak into th parnt group at ah gnration, rustaans ar mor
and sltion an afft it indprrdntlyof th parnt group; it may drift proah to infrring v
awzry,volutionrv spaking, t om a nw spis.Padomorphosis is a p r i s o n o s p i st h a
flrm of htrohrony in whih suall1'mtur adr-rltsrtain som or all f morphologially divrs
thir lrval (or juvnil) faturs.It an ris ithr through arr alratd u s t a a n sw ' hih alm
rirt of sxul maturation (progrrsis)or through dlrationof th dvl- pially prolmati. Fr
opmnt of nonrprodutiv organs (notny),ut th prosssar livd a group of wormlik pa
to hav ontriutd to th rlativl,vfast volution of divrs arthropod ody oganisms possssfw
foms during th :rrnbrian.Progrrsisapps to tlre mor omnron of lationship with som
th two htohoniprosssy whih rustilanody plans divrsifid, nlids, myriapods, tar
and it is thought to hav playd a ky rl in th volution of many of th within thir own phylu
maxillopodan rustaans' whih xhiit izarre and highly rdud ody strutur rvald sim
plans. In ontrast' ladorans, lld ..watr flas,',ar susptdto hav taans (fish li), a f
volvd by notny from a lam-shrirnp anstor tlr:rtam suz.rllyma- nular l8S riosom
tur ur ypassdtlr rr:ruplirlrval stagf dvloprerrt. o t h r p o t n i lr l a t i v
Som sintistsspulatthat th gn-rgulatorypathways ontrolling d- Th DNA ppoh
vlopmnt, whthr involving homoti gns or ons ontriuting to ht- shipsamong organism
rohrony, Wr mor flxil among th rlativly primitiv, lss omplx ial similaritis du n
arnbrian nstorslf modrn rusransand thr onditions in th distant dvlopmnt of similar
pst wr thrfor nror fvoral for th rapid voltrtion of divrs ldy tatswih similrslr
pins than tlrv lrav n vr sirr.Th main lLlstaanlinagsosrvd indpndntly volvd

l lws or ..hlipds,'sn in
as thos of th domn (a
wtmrning. Variation t Ho
laklng adominal sgmnts
y ovriolkdas rustans).
Lsfor th Hox.lik Distl-less
hav signifiantly advand
;is of rusta linrs and
the volutionof morphology,
sity of usranody plans.
ng th timing of dvlopmn-
l possmay quikly rsult
rnstwn nstors nd
dvlopmntaland growth
sultin a dramatially altrd
e priod during whih a gn
rlt in an offspringwith ntn-
;. If th mutatin is hritabl'
ntnn'In sorrustans
lrin shrimp, lso alld sa
ssfulopulation. als with
;sflrllytlrn ons with shotr
th a partiularvriant among
oms rprodutivlyisolatd,
n tg r o u t l rg n r t i o n '
' parnt group; it may drift
spis.Padomorphosisis a
adults rtain som or ll of
:ithrthrough an alrtd
Lghdlratiorrof th dvl-
:t oth prosssirr livd
n of divrsarthropod ody
s to the mor ommon of
aanody plans divrsifid,
:hvolution f many of th
.r and highly rdued
ody
flas,''r susptdto hav
or that anrsually m-
:vlopmnt.
ory pathwaysorrtrollingd-
or ons ontriuting ro ht-
ivly primitiv, lss ompl
t h a t o d i t i o n si l r t h d i s t n t
id volution of divrs ody
rustanlinagsosrvd
rustcs 497
today ar th volutionary rsult of this dvlopmntal plastiity. ah dis-
plays a onfusing omintion of primitiv and dvand faturs,a ombi-
nation that maks disrning volutionary trnds diffiult.
To irumvnt ths diffiultis, systmatists hav turnd to mans othr
than th traditional omprison of dlt nrorphology. On approah onr_
pars th larval morphologis of diffrnt taxa and is asd on th assump-
tions that arly dvlopmntal stags ar fr of th uniqu morphologial
spializations sn in adults and tht shard laval faturs may thrfo
mor aratly rflt ommon anstry. Th s of barnals, mntiond
aove,illustratsthis point, and larval morphology has furthr n usd to
unit groups suh s th Arothoria, hoaia, nd parasiti Rhizo-
phala within th arnal infralass irripdia and to rogniz th para-
siti Asothoraida as th sistrgroup of irripds.Lrval faturshav lso
n usd in attmpts to disrn shad anstry mong hrmit ras arrd
primitiv as wll as mor advand tru ras. Th rnt first disovry of
fr.living lithotrophi (nonfding) rmipd larva is posing nw y-
pothess aout th ffinity of this nigmati group with othr rustaans
and th voltion of th rustaannaupliar lva.
Anoth approah, on that has volutionizd and rvitalizd th disi-
plin of phylogntis, rlis on omparisons of DNA squns for grrs
shard y diffrnt typs of ustaans.For xampl, th gn oding for th
nzym ytohrom oidas suunit I ours in th mitohondrial gnom of
vy rustan. Th DNA squnsfor this partiula gn in diffrnt
spis of rustaans an b dtnrind and th squns ompard. Th
diffring dgrs of similarity among th DNA squns an tll us whih
rustns af mor losly iatd than othrs. Th DNA-squn ap-
proah to infrring volutionary rlationships is patiularly usful for m-
parison of spis that shar fw morphologial haratrs, aus vn
mophologilly divrs spiswill hav mny gns in ommrn.Prasiti
rustaans' whih almost always hav highly modifid morphologies' ar s-
pially prolmati.For xampl, th volutionary affinitisof pntastoms'
a group of wormlik prasits,had gnrtdmr'rhspulationausths
organisms possss fw morphologial harateristis that might suggst a r-
lationship with sotn othr group. Thy had n variously allid with an-
nlids, myriapods, tardigrads, onyophorans' and mits and evn plad
within thir own phylum, ut a omparison of sprm dvlopmnt and ultra-
strutur rvald similaritis twn pntastoms and branhiuran rs.
taans (fish li), a finding that was rrooratd y DNA analysis of
nular 18S ribosomal RNA gns and mitohondrial gns for ths nd
othr potntial rlativs.
Th DNA approah also is usful for dtrmining phylognti rlatiolr-
ships among organisms that hiit a high dgrof homoplasy (morpholg-
ial similaritis du not to shard anstry ut instad to onvrgn,i..'
dvelopmntof similar morphology in sponsto volution in similar habi-
tats With similar sltivrgims).For xampl, urrowing rustaanshv
indpndntly volvd modifid digging appndags and a stramlind ody
498 rustns

form; diffrnt urrowing spis may ppar morphologially similar ut pattrn thory,'' also r
not shar a rnt ommon anestor. volutionary affinity among spis irr losr affinity to t
may also misonstrudif any of thm has undrgona rent volutionary all thr viws onsid
rvrsal to a mor anstral morphologial fatur. Th DNA approah to and omplling mol
infrring rlatdnss avoids th problms prsntd y ths instans of rustaan nstor' I
,
morphologial homoplasy. Thrmosanaans r minut rustaans found som ar alling th
in groundwatr and springs' and they hav variously n plad within th Sustarrtially mor da
suprordr Praarida (in part austhy hav a rood hamr, a fatur th National Sin
found in som praarids)and in a ompltly sparatsuprordr,th Pan- Arthropod Phyl<lgn
arida (austhir rood hmr has a distintiv dvlopmntl origin 120 diffrnt gnsfrt
and loation). Similarly, mysid and lophogastid shrimplik rustaanshav pnding and r high
bn onsidrd ithr to ompris an ordr of praarids or to sprat
ordrs and ludd from th rarida. In ah of ths ass, DNA vi- BIBLIOGRAPI
dn has n usfirl in rsolving long-standing phylognti unertainty Bowman,T. ., and L.
and has shown that morphologial faturs (.g.,th rood pouh and fa- l,. G. Al,d.,S1
turs of th arapa' forgut irulatory systm' and ys) shard y som tsiologof rust
so-alld praarid spis may in fat hav volvd indpndntly or n a l m a n ,W . T . 1 9 1 1 .
Duffy, J. . ] 996. us
lost ntirly.
orty,R. A., and R.
Som rustaan spis (.g.,on ntir infralass of rustaans rlatd to ssoiationSpi
arnals, th Fatotta or ..y.larva'') ar known only from thir larva N{artin,J. W., and G.
(adult forms hav nvr n found). othr spies ar known from only mal rL|std. Sin
or only fmal forms, so knowldg of ky morphologial faturs is similarly Anglsounty.
laking. In th mphipod family Luothoida' mls may undrgo a adial Poor,G. . B. 2004..
l dtifi tiltt' 'ol
transformation during a singl molt, suh that pr- and post-molt mals wr
Shrm,F. 1986.rus
rronously lassifid as not just diffrnt spis ut as longing to diffrnt
familis. Individuals of som spis of aridan shrimp ar squntial hr-
maphrodits, ginning lif as mals and latr transforming int fmals (a
uvir, Grgs
ondition trmd protandry). Taonomi rognition and assignmnt is sp-
ially ompliatd y a spisthat has supr-malswith nlargd gnitalia Although Gorgs
(ths nvr transform into fmals) in addition to th mor typial mal- playd a prominnt r
phas hrmaphrodits that will vntually transform. In all of ths ass voluntarily. In rtros
DNA vidn is somtims th only mans y whih som spis an lishd yond doubt
idntifid auratly and plad within an volutionary hirarhy. that this had ourr
Finally' in addition to th intratility of dtrminingrustaanrlation- drmind th ida of
ships, th qustion of rlationships among rrrstaansand th othr major lassify organisms ir
tant arthropod suphyla (Hxapoda, or insts;Myriapoda' or ntipeds philosophr ihl I
and millipds; and hliriforms, omprising spidrs, mits, tiks, and volution than Jan-
horssho ras) is prpling, for many of th sam rasons notd aov. tion thory. owvl
Varius altrnativphylognishav n proposd on th asis of diffrnt uvir's allgd ont
intrprtationsof morphologial vidn.For instn' th ..uniramian hy- tt. som of uvir
pothsis'' onsidrs arthropods to polyphylti (i..' unlikly to hav had a th sin of volut
singl, ommon anstor) and sparats hlirats (.g., spidrs) and rus- drstood in light of
taans (possssingiramous appndgs)from uniramus insts' myriapods, uvir was primr
and onyophorans. Th ..mandiult hypothsis'' taks a monophylti (sin- him was lassifiati
gl anstor) viw of arthropods and units rustans' insts, and myri- prviding unifid
apods within th andiulata. Yt a third viw, asd on an ..arthropod was losly rlatd t

r morphologially similar ut
ionarr' irffiniy alllolrg spis
l d g o n a n t v o I u t i o n
.I'h
:atur. DNA approah to
sntdby ths irrstnsof
s a rinutrustaans found
riously n plaed within th
rva rood hamr, a fatur
. spratsuprrdr,th Prn_
stintivdvlopmntal origin
id shrimplikrustaanshav
lf prarids or to [r spat
ah of ths ass, DNA vi_
d i n g p h y l o g n t i u n
:.g.,th rood pouh and fa-
m, and ys) shard y som
:volvdindpndntlyor n
alassof rustaanSlatdto
known only from thir larva
:isar known from only mal
phologilftursis similarly
, mais may undrgo a rdial
pr- and post-molt mals wr
s ut as longingto diffrnt
:an shrimp ar sqntial lrr_
r transforming into fmals (a
gnitionand assignmntis sp-
'-mlswitlr nlargd gnitalia
ion to th mor typial mal_
ansform. In all of ths ss
y whih som spis an
utionaryhirarhv.
trminingrustaanrlation-
l s t a a n s n d t h o t h r m a j o
rts;yriapoda' or ntipds
;ing spidrs, mits, tiks, and
h sam rasons notd aov.
posdon th basis of diffrnt
r instan'th ..uniramian hy-
:ti(i..'unlikly to hav hd a
:rats(.g',spidrs) nd rus-
uniranrousinsets,myriapods'
sis'' taks a monophylti (sin-
)rustaans'insts, and myri-
iw, asd on n ..arthropod
uuier 499
pattrnthory,''lso viws athropodsirs monplrvltiut plasrustaans
ilr losr affini' to Irliatsthan to insts arrd ryripods. Proporrnts of
y ll thr viws trnsidr insts and ryriapods tl sistr ta, ut now nw
and ornplling nrolular vidn stlggStstht insts ar drivd from a
rustai]11 anstof' thr1,n-rakingth rusta a paraphylti taon that
som ar alling th ..anrttstaa.' nd making insts ..flyingrustans.''
Sustarrtiallymor dta ar ndd to rsolv ths otltstanding issus.In 2001
th Ntional Sin Foundation wrdd ovr $1 milliln to fund tlr Dp
Arthropld Plrylogny projt to ltirrovr 60,000 DNA ss fronr roughly
120 diffrnt gnsfrom 85 arthropol spisfor just this purpos. Rslts r
pnding and ar highly antiipatd.
|LloGRAPY
r t a i n t y Bowmn,T. ., and L. G. Al.1982.lassifiation of th rntrusta.
In
L. G. Al,d.'Sstemtis' th FtlssilRord,,ldBiogogrdph.o|. 1'Th
Biolog ()frL|sttle, 1_27.Nw \rk: AadrrriPss.
alman,W. T. 19l |, Th Li| of rust. Londorr:Mthun& ompny.
Duffy,J. . |996' }iusoiality in a orl rfshrimp.Ntur 381:512_514.
Forty,R. A., nd R. . Thms'ds.l998' rthropodRltiosips. systmatls
AssoiationSilVo[r-rrrr Sris.5.5.
Llndon:hpmanarrdHal[.
Martirr,.|.\W.,nd G. . Davis. 2001. UpdtdIssifiti<'ltt of tb Rt
tst.SinSrisNo. 39. Ls Arrgls: Ntural History usumof Los
A n g l s o u n t v .
Polr'G. . B. 2004. ri Dplt!(',rust lf Sthrustrli: Guid ttl
Idti|itir.ollirrgwood, Vi.: SIRo Pulishing.
-7.S.
Shram, F. 1986. rustdl.Nw York: oford Univrsity Prss.
uvir, Gorgs(\769-182)
Althlugli Glrgstrvir pliitly rjtd th rlodifiation of spis, h
playd a prominnt rol irr th arly history of vllutionarysin,lbit in-
voluntarily. In rtrospt,h mad two major ontributions. First, h st-
lishd yond dout th rirlity of th tintiorrof spis' and h showd
that this had urrd frquntly irr tlr histr of th :rrth'Sond, h un-
drmind th ida of th grat hain of ing;h provd it was impossil to
lassif1'organisms in a srial ordr, ithr logial or hronologial. Frnh
philosph Milrl oultsid rht uvir did rrrorfo th mrgnof
volution than Jan-BaptistLamark or arl1,dvoats of th trnsmuta-
tion thory. H.vr,this rtrosptivvluation is historially mislading.
Cuvir's llgd Ontriutionstl volutilnmust plad irr thir own on-
tt. Som of uvir's tlrtlisand rnthods Wr ind{inorporatd into
th sin of vtllution, ut th ntirty of uvir's work an ttr un-
in light of a diffrt onption of th history of lif.
drsttlo.l
trvir was primarily a zoollgistnd n an21romist. h main problm for
him was lassifition. H saw oparativ natomy s ing apal of
providing a unifid thory of living ings. Ctrvir's omparativ anatomy
,as losly rlatd to physiolog and to th study of pst oganisrs.From
500 uuir

th ginning of his ar to th nd, h ombind antomial dsription

and lassifiatinof animals, prsnt or tint' with physiologial hypoth-
ss aout thir funtitlnal organization. Bausof his major disoverisand
mthodologial innovtions in this road fild of rsarh,h has oftn n
onsidrd th founding fathr of oth omparativ and stratigraphi pal-
ontology. In oth filds, h had prdssorsand omptitors, ut h playd
a disiv rol in ah as. Historians of zoology (.g.'Russll L\9161 19s2)
insist on th first aspt' whil historians of palontology (.g., Rudwik
1976) emphasiz th sond aspt. In uvir's work, th two dimnsions
an hardly dissoiatd.Nvrthless' thy ar rfltdin th titles of the
two major synthti ooks that h pulishd (uvir 1812,1817). Although
this is somwhat artifiial, th two aspts'stratigraphipalntology (sp-
ially th thory of xtintion) and omparatlv anatomy, will amind
sparatly.
In th sond half of th svntnthntury, svralsintistslaimed that
fossils ould intrprtd as rlis of xtint spis.But th qustion r.
maind opn whthr ths spis had indd disappard or still istd
somwhr on th arth. For instan, l,amark, who dsrid a hug numr
of nw invrtrat fossils' rjtd th ida of tintion. uvir proposd to
rsolv this qustion y using fossils of larg trrstrial animals as tst. In
ontrast with marin animals, larg trrstrial animals ould hardly rmain un-
known y 1800. In 1796, uvir gav th first dtaildproof of suh a as.
On th asis of drawings rivd from Paraguay, h idntifid a giant fossil as
an unknown animal rlatd to th sllths. alld this animal gtherium.
Th laim tat egthriumad disappard was disivly supportd y th
pris dsription of th fossil, and its assignmnt to a known fmily (th
sloths). Latr, uvir umulatd vidn aout th disapparanof num-
br of othr larg trrstril animals (mammoth' mastodon, Pliherim, and Part of Gorgsuvi
many spis of fossil lphants,roodils, dr, t.) in diffrnt parts of mummifid organism
th world. By us of anatomial dsriptionsof unprdntdprision' h him on his ampaign
pointd out is idnti
providd ruial support for th laim that ths animals no longr istd.
that thris no hang
Bausmigration ws unlikly' only two hypothssould aount for th
disapparanof ths animals. At th vry tim whn Lamrk gan pub-
lishing his transmutation thory, in 1801 uvir statd thy had n ithr disappard in th n
ntirly dstroyd or modifid in thir forms. Cuvir rjtd th modifia- of frshwatr and m:
tion hypothsis aush thought thr was no vidnto support it. He siv vnts of lvat
advoatd th tintion hypothsis aus h thought thr was vidn in juntions twn t]
its favor. ummifid animals rought ak from gypt y tinn Goffroy ludd that suddn 1
Saint-ilair (Napllon'spdition) did not provid th slightstlu of th tion of sussivtr
modifiation of animals twn antiquity and th prsnt (sfigur).If no from othr plas.
vidn of small hangs was availal' how ould Lamark and othrs - As shown y ar
trapolat to largr hangsovr long priods of tim? In ontrast, uvir and tion...wasprimar
Alxndr Brongniart's osrvations(1s0s) of the sussivstrat aove th vir's gnral rprs
halk in th Paris asin thus suggstdth arupt disapparanof loal fu- rvolutions, whih t]
ns. Ovr long distans (gratrthan 100 kilomtrs),thr was onstant migrtion (in and or
suprposition of th strata' ah strtum with its haratristifossils. that that ould hav dst
 ier 501

ind natomial desription

:,with physilogial hypoth-
o f h i s m j o r d i s o v r i sa n d
of rsarh,h has oftn n
rtiv and stratigrphi pal-
rd on-rptitors, ut playd
g y ( . g .R, u s s l l[ 1 9 1 6 ] | 9 8 2 )
palontology (.g., Rudwik
.'swork, th
two dimnsions
r rfltdin th titis of th
J u v i 1 8 1 2 , 1 8 | 7 \ . A l t h o u g h
rtigrirplripalontollgy(sp.
'r antomy' will xmind

svralsintistslaimd that
s p i sB. u t r h q u s t i o n r -
d disppardor still isted
who dsid hug nutnr
)xtintion.uvir prop<lsdto
:rrstrial nimals as a tst. In
i m . r i s o u l dh d I y m a i nu n -
dtaildproof of suh a as.
y, h idntifida giant fossil as
Ildthis niml t'gthrim.
irs disivlysupportd y th
-
nnt to a known family (th
,'t-
rt th disapparanof num-
mastodon,Plotrim, and Prt of Gorgs uvir's argumnt agtrinst volution was sd on th
:r, t.) irr diffrnt prts of murrnifid organisms rought ak y Napolon's sintists,whrl went rvith
him on his amp:rign tl gypt. his is th skltlrrof n iis, whih uvir
I r.rnprdntd prision, h
pointd out is idntil to th skltons of iiss tlda,v,thus apparntlv showing
animals no longr istd.
tht thr is no h:.rngeovr tim. (Fom uvir 1804.)
'othssould aount for th
rwhn l-amark gan pu-
:r statd thy had lr ithr disappardin th nt stratum. Futhrmor,thy osrvd an altrntion
3uvir rjtdth modifia- of frshwatrand marin forr:rtions.This ould b aountd for y sus-
to vidnto support it. H siv vnts of lvation and dprssionof th ontinntal aras. Baus th
, h o u g h t h r w a s ' " . i d n i n juntions twn th two kinds of strata Wr rthr sharp, uvir on-
rr gypt y tinn Goffroy ludd that suddn gographi hangs had n rsponsilfor th tin-
'ovidth slightst
lu of th tion of sussivtrrstrialfaunas and for a rpopulation by xisting faunas
th psnt(sefigur).If no frorrr othr plas.
uld Lamark and ohrs - ..uvir,s
As shown y artin Rudwik (1976, 117), rjtiorrof volu-
tin-r? In ntrast' uvir and tion . . . was primarily a dfnsof titio' not of spial crtion.,' :u.
sussivstrata aov th vir's gnral rprsntationof th history of lif onsistdin a srisof loal
lt disapparanof loal fau. rvolutions,whih thmslvsimplid th tintilnof som spisand th
ntrs),thr was a onstant migration (in and out) of othrs. nvr mntiond ir gnral atastoph
its htristifossils, that that ould hav dstroydall xisting spison arth. Suh a h,vpothsiswas
t02 uuir

mntiond latr y othr sintists (Alid d'Origny in Fran, William 181'2.Rherhssur

Bukland in ngland) who also dvoatd vnts of ..spialration'' fol- 1 8 1 7 [ 1 9 8 3 ]L. e R g
lowing th ..atastrphs.''uvir had no thory to aount for th divrsit sruird bs I,histoir
ompr.4 vols.Paris:
of living ings. For hirr.r,this qustion was yond th limits of th positiv
ontmporarns.
sinof his tim. uvir, G., and A. Brongniar
Although Cr,irhad no tho11,for th origin and divrsity of living or- nvironsd Paris.lol,lr
ganisms, h lrad a thOry of lassifiationthat rlid on thr rational prini- Fouault,N|. |966. Les mots
pls. Th first prinipl, th prinipl of th onditions of istn, stats Pris:Gallimard.
..th Rudwik, r|"[.t976. Th
tht diffrnt parts of ah irrgmust oordinatd in suh a Way to
PIeontolog.higo:
r n d r p o s s i l t h i s t n o f t h b i n g a s a w h o l ' ' ( u v i r 7 8 1 7 , 1 : 6 \ . Russll,. S. 1916 ||982];. Ft
This prinipl dos not rfr primarily to trnal onditions' ut to th n- AimI orpholog.Lo
ssar oadptation of ir-rntionsand organs within tlr animl ody. Tlr hiagPrss.
sond prinipl, th prinipl of orrltion of pirrts, is arl appliation of
th formr prinipl. It says that a givn funtion nnot vry without or-
rsponding m<ldifiationsin all rltd orgns. In prati, ur'ir usd this
pinipl t infr th shp trf a nurnr of organs in a fossil fron-rth shap
of on organ. uvir usd his first two prinipls for dnying th possiility
of th modifitrtion of spis.organs and funtions in an org:rnism ar so
hrmorriously oordinatd in an orgnism thi1t any hng in :r givn prr
would rquir simultantlushangs in mny or all of th othr parts.
Cuvir's third prinipl, orrowd from botanist Brnard d Jussiu, was
th prinipl of suordination of hratrs.It statsthat som haratsar
mo fundamntal than othrs' in th sns that thy dtrmin ;r st of possi-
ilitis for th rst of th organism as whol. This prir-riplld Cuvir to dis-
tinguish four diffrnt mranhmnts, or funtionirl typs of animls:
vrtrats'mollusks, i1tiulats' nd radiats.Ths typs Wrnot rduil
on to anothr. Th notion of major htrognous strutural-funtional
arrangmnts ondmnd any attmpts to l:rssify nimals-and evn mor
organisms in gnral-aording to a sriirl ordr. For uvir thr was no
possilasnlingsrisof mplity (as rsarhdy Lamark) nd no ul-
timat unity of ody plan (srudidy tinn Goffry Saint-Hilair; s th
lphtialntry ..tinnGoffoy Saint-ilair'' in this volr'rm).This was
a disiv rgumnt against th hypothsisof ommorr dsntof organisms.
Nvrthlss,in a onvolutd way, th notion of mbranhmntsplayd a
rol irr th mrgne of harls D:rrrvin's and Alfrd Russl \Walla'snotion
of ranhing volution.
As notd y most ommntatos, uvir prfrrd fats to idas, and h
lookd systmtiallyfor lws. H did not admit ithr modifiation or sp-
ial ration of spis,bauslr ould not find fats supporting ithr hy-
pothsis. orrlativly, h livd that rpatd vnts of tintion and
lol rvolution had o1,drgular laws of ntur.

BILIoGRAPHY
uvir,G. 1804.6moirsur Llbis dsninsgyptins.
lsdu usum
tiold' HistoirtrIl
A(1804):116-1.5.
 uuir 50

rbigny in Fran, William 181'2.Rehrhessur ls ossmsfossilsdes qudrupdes.Paris:Dtrvill.

s of ..spialration'' fol- 1817 |1983l.L Rgiml distribd'prsso rlrgistiopor
to aountfo th divrsity seruird bs l'histoirturellds im et d'itrlductio I'tomi
ompr.4vols. Paris:Dtrvill.Rprint,Paris:ditions ds rhivs
r d t h | i m i r so f t h p o s i t i v e
ontmporains.
uvir,G., and A. Brngniart.1808.ssaisur la gographi minralogiqu ds
nd divrsity of living or- nvironsd Paris.Jourl des ins 23: 421458.
d t l n h r r a t i o n a lp r i n i - Fouult,t^. 1966. Les mots et ls choss:U rhologieds sishumis.
ditions of xistn,stats Pris:Gallimrd.
ordinatdin suh a Way to Rudwik, N{. |976. Th ing of Fossils:pisodsin the Histor of
Plotolog.hiago:Univrsityof hiago Prss.
h o l ' '( u v i r1 8 1 7 , | : 6 ) .
Form d Fution:A otributilto th Histor of
Russll,E,.S. 1916 L1'9821.
onditions'ut to th n- Aiml orpholog. London:John urray. Rprint, hiago:Univrsityof
:hin th animal ody. Th hiago Prss. -J.G.
parts, is an appliation of
l annot vary Without or-
l prati'uvir usd this
s in a fossil from th shp
for dnying th possiility
ions in an organism are so
rny hang in a givn part
rll of th othr parts.
st Bnard de Jussiu, was
'stht som haratrs ar
y d t r m i n a s t o f p o s s i -
;priniplld uvir to dis-
rtional typs of animals:
rstypsWrnot rduil
1ous strutural.funtional
y animals-and vn mor
:. For uvir thr was ntl
rd y Lamak) and no ul-
lffroy Saint-Hilair; s th
,'
in this volum).This was
mon dsntf organisms.
mranhmntsplayd a
. d R u s s |l W a l l a 'ns o t i o n

rrd fts to idas, and h

ithrmodifiation or sp-
fats supporting ith hy-
l vnts of xtintion and
t.

ens.lsd usm
 known and appriat
as a man of solid wort
ntifi laims a fair an
Darwin's first systn
h was traind y ar
th world is formd 1
tastrophs),th grat
Lyll' whos thr-volt
grly dvourd y th
thsis, arguing that tht
Darwin, Chrls (1809_1882) Lyll hypothsizdtha
harls Rort Darwin, th nglish naturalist and volutionist,was orn in ing transitory and a
th town of Shrwsury on Fruar 12, 1809, th sam day as Araham th distriution of lan
Linoln aross th Atlanti. H did on April 79,1882, at his hom in th vil. kind of watr d, wh
lag of Down (Knt). H was th author of o th origi of Spies' pl- rings on lvation in
lishd in 1859, and h is onsidrdth fathr of volutionary thinking. Darwin aptdth
.
Darwin's patrnal grandfathr was physiian asmus Darwin, author o{ in the Lyllian mod.
Zoonomi (1794_7796), on of th arlist works prolaiming volution. islands? Lyll argud t
His fathr' Rort Darwin, was also a physiian as wll as a mon man' ar- argud that it was im
just rak th sa surf
ranging mortgags twn aristorats with land and a nd of ash and
usinssmnwith ash and a nd of saf loans. Darwin's matrnal grandfa. lands nd thn thy si
thr was Josiah \Wdgwood, th man rsponsil for industrializing th pot- th sa surfa,ven a
try trad in Britain. Darwin mntd th !7dgwood onntion furthr pltly nth.
.W.dgwood. harls Darwin's g
whn h marrid his first ousin, mma and his wif had 10
hildrn, svn of whom livd to adulthood. on was Gorg Darwin, th In Gln Roy, a small v
minnt physiist and authrity on th tids. indntations around t
harls Darwin was duatdfirst at Shrwsury Shool, on of ngland's as thy wr knowni
lading privat shools, and thn snt to dinurgh Univrsity in th foot- around wat that on
stps of his fathr and grandfathr, intnding to train as a physiian. Un- lat 1830s, Drwin a
h,ppy in this dirtion' h nrolld at hrist's ollg in th Univrsity of ara had mvd upw
Camridg, intnding now to an Anglian lrgyman. Whil at univrsity' th Swiss sintist Lo
oth in Sotlnd and ngland, Darwin startd to show a strong intrst in watr Was a lak, da
sin,and y th tim h graduatd in 1831, senior sintistshad startdto appriatd his mista
spot oth th intrst and th talnt. Th sustantial Darwin-!7dgwood for- thinking in a atastr
tuns mant that Darwin nvr had prssingfinanial ndsand ould always of God.
indulg his passion for mpirial inquiry. Darwin would not I
Divrtd from th lrial arr y th offr of a voyag on HMS Begl, As a young man prpz
undr th aptainy of Rort Fitzroy, Charls Darwin spnt fiv yrs going tsgl voyag his lit
around th oast of South Amria and vntully irumnavigating th strongly in a God wh
glob. Although h was ngagd s a gntlman ompanion to th aptain, trfr with his rati
Darwin soon bam th full-tim ship's naturalist, aumulating xtnsiv th writing and puli<
olltions that h shippd hom' thus adding to th alredy good opinions lik many mid-Vitor
of his Camridg mntors. Darwin lso mad tnsiv travls inland. Th ism. h Anglian b
diaris h wrot on th voyg Wr th foundation for a popular tavl ook aus it insistd that (
titl'edTh Vog of th Bgle, first pulishd as part of th offiial rord of of organisms, suh at
th ship's travls and thn indpndntly. Thr is no dut that ing Arhdaon William I

-t04

t and volutionist,was born in
09, th sam day as Araham
9,1882, at his hom in th vil-
th rigin of Speis, pu-
of volutionarythinking.
tn rsmus Darwin, author of
works prolaiming volution.
an as wll as a mony man' ar_
lnd and a nd of ash and
rs. Darwin's matrnal grandfa-
l for industrializingth pot-
Wdgwood onntion furthr
wood. H and his wif had 10
on ws Gorg Darwin, th
sury Shool, on of ngland's
nurgh Univrsity in th foot-
g to train as a physiian. Un-
's
ollg in th Univrsity of
:lrgyman.Whil at univrsity,
d t o s h o w a s t r o n g i n t r s ti n
sniorsintistshad startd to
antial Darwin-.$dgwoodfor-
rarrialndsand ould always
r of a voyag on S Bgl,
D r w i n s p n tf i v y a sg o i n g
ntually irumnavigating th
a n o m p a n i o nt o t h a p t i n ,
,ralist . u m u l a i n g t n s i v
t o t h a | r d yg o o d o p i n i o n s
: tnsiv travls inland. Th
rtionfor a popular travl ook
as part of th offiil rord of
.hr
is no dout that ing
Dri, hrls 50.'
known and appriatd y th gnral puli mntd Darwin's fputation
as mn of solid Worth. This status hlpd latr to giv his ontroversial si.
ntifilaims a fair and fvoral haring.
Darwin's first systmati intrsts as a sintist wr in gology. Although
h was traind y ambridg profssor Adam Sdgwik, who livd that
th world is formd y oasional priods of grat rrrptionand turmoil (a-
tastrophs),th gratst influn on Darwin was Sottish gologist harls
Lyll' whos thr-volumwork, Priipls of Gllog (1830-1833), was a-
grly dvourd y th voyaging naturalist. Lyll proposd th uniformitarian
thesis,arguing that th gologial world is govrnd y unrokn law. Also,
Lyll hypothsizd that th world is in a kind of stady stat' with all hangs
ing transitory and funtion of altring limat rought on y hangs in
th distribution of land and sa. H lookd on th surf of th glob as a
kind of wtr d, whr dposition in on ara auss ssidnand thus
rings on lvtion in anothr ara.
Darwin aptd this viw ompltly, and his first sintifi triumph was
.Why
in th Lyllian mod. dos on find irular orl rfs around tropial
islands? Lyll argud that thy ar th tops of tint volanos, ut Darwin
argud that it was improal tht all of th volnos would oinidntally
just rak th sa surfa.Rathr, h argud, th oral grows round th is-
lands and thn thy sink, with th oral growing vr upward to rmain at
th sa surfa' vn aftr th islnds thmslvsmy hav disappardom-
pltly bnath.
harls Darwin's gratst sintifi rror Was also in th Lyllin mod.
In Gln Roy, a small vally in Sotlnd' thr a thr horizontal' traklik
indntations around th sids. !(/hat is th rason for ths paalll roads'
as thy wr known? All agrd that thy Wr th rmains of ahs
around watr that on filld th gln, ut whr is th watr now? In th
lat 1830s, Darwin argud that it had n sawatr ut that th whol
ara had mvd upward and nw th watr was drind. Shortly thraftr
th Swiss sintist Louis Agassiz visitd th ara; h at on saw that th
Watr was a lak, dammd y i that had sin mltd. vntually Darwin
appriatd his mistk' ut this took tim us Agassiz wrappd his
thinking in a atastrophi thory that rlid havily on th intrfrn
of God.
Darwin would not hav ojtdto th supposition that thr was a God.
As young mn prparing for th lgy' h ws a sinr hristian. on th
Bgle voyag his blifs hangd and h am a ommittd dist, living
strongly in a God who workd through unrokn law and who dos not in-
trfr with his ration. This position lastd for many yars, right through
th writing and puliation of te rigi. Thn, toward th nd of his lif,
lik many mid-Vitorian intlltuals,Darwin's rligion fadd into gnosti-
ism. Th Anglian akground was lways vital to Darwin, howvr, b-
aus it insistd that God's dsigning powrs an sn in th adaptations
lf organisms, suh as th ys for sing and th ns for smlling. Lik
Arhdaon.silliam Paly, whos I'Jturl Tholog (1802 [1819]) Darwin
506 Dnui, hrls

rad as an undrgraduat, Darwin livd that th most signifiant fatur thinking pulily. Th rt
of th living world is tht it is ompl and dsignlik,t th nfitof indi- also (aftr a dd or t
vidual organisms. ism. lWhat is lar is th
Lyllian gology dirtd Dwin to thinking aout organisms; fossils wr tion than for Darwin's <
vital vidn of subsidnand lvation. Thus primd, in 1835, whn th to dny th lttr outr:
Begl rahd th Galipagos Arhiplago' a group of islands in th Paifi, auss) ranging fom L
)arwin was astoundd to larn that th dnizns, partiularly th irds and tis) to saltationism (r
th giant tortoiss' r diffrnt from island to island. volution from a om- som kind of guidd v
mon anstry is th ovious answr' spiallyto sot-neonwho livsthat Darwin had intnd
God's glory is shown through the working of law. Howvr, it was not until would now writ, filling
h rturnd to ngland in th following spring of I87 nd was onvind only on of ths oo
that th Gal6pagos varitisa truly diffrnt spisthat Darwin mad th Plts dr Drllesttc
lap to tansformism, as it was thn alld. inadquat hypothsis
Darwin rea|ized tht h ndd a mhanism to plain his nwfound - that partils ar givn.
lif. As a graduat of th Univrsity of amridg, h was awar of th tri- y h s r g a n s .T h i s
umphs of its grtst sintist, Isaa Nwton. Darwin thrfor sought for to whih Darlvin alwa
th organi world a kind of quivalnt of Newton's univrsal law of grvita- Apart from a sris
tion. H found his answr in Sptmr 1838. H startd with th althu- tl ook on orhids, a
sin lulation that populion prssurswill tnd to outstrip supplis f plants, yt anothr on
food and spa. H notd that thr smsalways to b a sour of variation yars Wr on th hum
in natural populations. Hn, arguing analogially from th susssof th and a kind of supplm
rdrsof animls and plants, Darwin onludd that th will e a natu- mls (|87z\.In th or
ral form of sltion, with som winning and othrs losing, and suss ing aout humans, othr tl
a funtion of th winnrs' puliar faturs. Vitally, this natural sltion, o that h thought his th
..survival th topi unintrsttng
of th fittst,''as it am known, lads not just to hang but to
hang in th dirtion of adaptation. thory on th tal ant
Darwin soon Wrot up his thinking as a skth of a thory' ut for rasons mnt about humans.
still undetrmind h did not pulish. Undoutdly a factor was that h fll In this,Darwin was'
sriously ill nd for th rst of his lif ws an invlid and a rlus. (Th x- th impliations for Ho
t aus of th illnss is unknown. Som spulatthat it was psyhologial, pl wantd to talk a<
th strss of having suh an volutionary thory. Othrs think it might hav prssur from frinds a
n th rsult of n illnss,hagas disas,pikd up on th Bgle voyg.) that lWalla had on
For mny yars, Drwin sat ol.l his volutionary thinking and was divrtd to plain hulnan vo
into a massiv taxonomi study of arnales. physial and soial, rs i
.!alla,
Finally' in Jun 1858, Alfrd Russl a young naturalist witing tion was th ky ausa
from th Fr ast, snt Darwi a short ssay ontaining th sam ids Dar- ary mhnism, sxu
win had 20 yars for. Thus spurrd, trats of Darwin's thinking and Darwin livd that l
.lfalla's mining th diffrns
ssay Wr pulishd at on. Darwin now rapidly Wrot up his
thory, pulishing o the origitl of Speist th nd of 1859. Although h did not
Te rigin was immdiatly ontovrsial, ut it is important not to xag. last dad.H ontinu
grat this ft. In Britain and n th ontinnt, and vn in Amria, many growing orrspondn
popl, oth sintissand th gnral population, quikly aeptdth ida rs ross th wrld as
of volution. Darwin himslf hd laid th groundwork for this y arfully with rspet and aff
ultivting mn-notaly anatomist Thomas Hnry Huly and otanists h did' y gnral a
Josph Hookr in Britain and Asa Gray in Amria-who would dfnd his from Nwton and a p

th most signifiant fatur
lik, t th nfit of indi-
ut organisms1fossils wr
primd' in 1835, whn th
up of islands in th Paifi,
s' partiularly th birds and
land. vlution from a om-
) somonwho livs that
v. Howvr, it was not until
of 1837 and was onvind
lis that Drwin mad th
to plain his nwfound -
g' h was wr of th tri-
)arwin thrfor sought for
l n . su n i v e r s l w o f g a v i t a -
{ startd with th althu-
tnd to outstrip supplis of
,s to souof variation
Llly from th susssof th
d that thr will a natu-
rs losing, and suss ing
lly, this natural sltion, or
.ds not just to hang ut to
r of a thory, but for rasons
:dly a fator was that h fll
valid and a rlus. (Th x-
rtthat it was psyhologial,
. othrs think it might hav
d up on the Bgle voyag.)
y thinking and was divrtd
' a y o u n g n a t u r a l i s tw i t i n g
:rtainingth sam idas Dar_
ts of Darwin's thinking and
n now rapidly wrot up his
r ndof 1859.
t it is important not to xag.
' and vn in Amria' many
ln, quikly aptd th ida
:rdwork fr this y arfully
{ n r yH u x I y n d o t n i s t s
ria_wh would dfnd his
Dri, hrls 507
thinking pulily. Th rligious tndd to mor rlutant, ut thspopl
also (ftr a dad or so) mst proaly aptd som form of volution-
ism. What is lr is that thr was onsidraly mor nthusism for volu-
tion than for Darwin's own mhanism of natural sltion.No on wanted
to dny th lattr outright, ut most wantd to supplmnt it with othr
auss' ranging from Lamarkism (th inhritan of aquird haratris-
tis) to saltationism (volution y instantanous mjor rrw variations) to
som kind of guidd volution whr God plys a ontinuing and ativ rol.
Darwin had intndd t'e rigi to a tsr for a sris of ooks h
would now writ, filling out in dtail all th laims mad rifly in te origi.
only on of ths ooks was vr writtn, Th Vritio of Aimls d
Plts under Dlmstitil (1868\, notl hifly for Drwin's somwhat
inadquat hypothsis aout hrdity, alld pangnsis. (Darwin argud
that partils ar givn off all ovr th ody and olltd in and transmittd
y th s ogans. This would thus nsur Lamarkian ffts, hypothsis
to whih Darwin always susrid.)
Apart from a sris of inidntal works that Darwin now authord-a lit-
tl ook on orhids' anlthr ln liming plants, a third on instivorous
plants, yt anothr on arthworms-th major laors of th post-rigi|1
yars wr on th human spis,rsulting in The Desnt of (1871)
and a kind of supplmnt,he prssio of th motios i d i-
mls (7872|' |n te rigiz, dliratly, Darwin had said virtually nothing
aout humans' othr than an unamiguous ommnt at th nd suggsting
that h thought his thory also applid to humans. It was not that h thought
th topi unintrstingor unimportant' ut h wantd first to gt th si
thory on th tal and not gt vrything at on lost in th gnral argu-
mnt out humans.
In this, Darwin was only partially sussful.At on, vryon sizd on
th impliations for Homo spiesof th ..monky thory.''This is what po-
pl wantd to talk aout. For a whil, Darwin stayd aloof. Thn, undr
prssurfrom frinds and supportrs,and prtiularly glvanizd y th fat
that \/allahad om a spiritualist and had turnd to non-natural auss
to xplain humn volution, Darwin argud in dtail that human natur'
physial and soial, is a produt of unrokn law. As always, natural sl-
tion was th ky ausal featur, ut Darwin supplementdit with a sond.
ary mhanism, sual sltion, whr organisrs ompt for mats.
Darwin blivd that sxual sltion was partiularly important in dtr-
mining th diffrns twn human ss and ras.
Although h did not liv to a vry old ag, Darwin's halth improvd in his
last dad. ontinud to work until th nd, inluding answring an vr.
growing orrspondn.Aknowldgd both y his ountrymn nd y oth-
s aross th world as On of th grat sintists of all tim, he was rgardd
with respet and afftion. hr Was thrfor no grt surpris tht whn
h did, y gnal aIaim,h was urid in $7stminstrAy, just aross
from Nwton and a pa or two away from Lyll.
508 Dri, rsmus

BlLloGRAPHY args. H was also i

Brown,J' 1995. hrles Drtui:A Biogrpb.ol. 1, Voglg.Nw York: th topi. For his twtl
Knopf. h wrot a small trt
202.hrlsDri: A Biogrph.Yo|, 2, Th Por rlf PIe.Nw York: FI orspndd v
Knopf. on aquaintan v
Darwin, . 1839.lournl of Rserchesito th GologdNturl istor of thinkrs of his day, D
th Vriousoutriesvisitd b HS Bgle.London: Hnry olurn.
th Frnh Rvoluti
1859. th rigi of Spisb s of NturI Seletio,or the
Prsrutioof FuourdRsi the Strugglfor Lif, London:John urray. tion of an normous .
1868. Th Vritioof imls d Plts uder Domstitio.London: aus h was a rilli
J o h nM u r r y . mn of his day; King
1871. Th Dstof ' d Sltioi Reltio /o Sex. London:John physiian.
urray.
Darwin's intrstir
1872. Th prssillf the motirlsi n d imls.l,ndon:John
urray. tiularly thos thrwr
Darwin,.[1794*1796]l801. Zooomi;or,-hLtasof rgiI'if.3rdd. tunnls for anals.Bu
London:J. Johnson. h atually startd wl
Lyll, . 1830-1833. Priniplsof Golog:Big n ttmpt to plin th (7794-1796), arrid
Formr hgsi the rth'sSrfb Rfrto sesNr,ui ral of his poms. A1
pertio.3vols.London:John Murray.
dards, Darwin mak
Paly,W. [1802l1819.Nturl holog(olltdWlrks,vol.4). London:
Rivington. attntion to th simil
Rus' M. 1979. Th DrtuiiRuolutio:SiRd i Tooth nd ltu. is alld homolgis)
hiago:Univrsityof ChiagoPrss. a amo for what h
208.hrls Dui. Oford: Blakwll. floatd th ida that
.Wall,
A. R. 1858. On th tndnyof varitisto dpartindfinitlyfrom th
is alld Lamarkism
riginal tp.lourl of the Proedigsof th l-iSoit,Zoolog 3:
5_62. -^4.R. nism of sual slt
Truly, howvr, f
th hang of organi
of hang in soity
Darwin, rasmus (\7 3!_\802)
vand, that is to sy
rasmus Darwin, th grandfathr of harls Drwin, was on of th bst- monrh (uttrfly)t
known early volutionists. A physiian' h livd in Lithfild in th nglish Darwin's latr pom
idlands and latr in Dry, also in th Midlands. H fathrd thr sons
Organ
with his first wif, thn two daughtrs with a mistrss, and svn mor hil-
was
drn with his sond wif. H was opn in his nthusiasm for sual ativity'
First fr
and to this nd h didd' as a young man' to giv up drinking lst it impir
Mov
his prowss. A grat nthusiastfor Linnaan taonomy, asd as it was on
Ths
sexual haratristis,Darwin was known to th widr puli as a pot wh
Nw p
hymnd th virtus of plant sulity' Th Lous of tbe Plts (1789) ing .sh
on of his majr litrary ahivmnts.
And
rasmus Darwin was los frind of industrialistsinluding pottr Josiah
.s(dgwood,
whos daughtr marrid rasmus's third son, Rort, and on of Thus t
whos hildrn was harls. Knly intrstdin mahins and thir uss, whit
rasmus Dawin was a mm of th Lunar Soit, a group of mn who Th V
mt monthly to disuss mattrs of intrst. This group inludd hmist Th l
.Uatt,
Josph Pristly, industrialist Matthw Bolton, and Jams famous for Th
his work on improving th stam ngin. Darwin was himslf an invntor, Whos
inluding a horizontal windmill' an untippal arriag, and anal lift for Imp
 Drtl,i, rsrs 509

args.H ws lso intrstdin agriultr.rr, writing sintifi tratis on

o|. 7, Vtlgig.Nw York: th topi. For his two middl hildrn, whonr h st up in thir own shool'
h wrot a small tratison fml duation.
2, h Poer of Pl. Nw York: H orrspondd with popl far and wid on topis of mutual intrst.
on quintan Was Bniamin Franklin. Lik many mor advand
eolog d Nturl Histor of
thinkrs of his da, Darwin approvd of th Amriirn Rvolution, as wll as
L o n d o l r :H n r y o l u n .
lf Nturl Sltio, or th
th Frnh Rvolrion,ntil it got out of hand. A mn of grat girth, a fun-
for Lif, Ltltllon:John N4urry. tion of an normous irpptitth was nvrthlss wlom hous gust -
s udr D nrcsti tit.ltl. Londln: aus l"lwas a rilliant onvrsationirlist. ws on of th wisst mdial
mn of his da-v;King Gorg III unsussfullyggd him to th royal
i Rltion,o Sex. London: John
physiian.
Darwin's intrst in volution sms to hav n spakd y fossils, par-
M d imls. London: John
tiularly thos thrown up in th ours of boring through mountains to mak
' Ls of Orgi Life.3rd d. tunnlsfor anals. But it was not until virtually th last dd of his lif tht
h atually startd writing on th sujt.is murjorpros work, Zoonomi
n Attt|1pttrl pli tb (|794-1796), arrid disussion of th topi' anl tlr sujt ntrs into sv-
?rto uss No in
ral of his poms. Although his thinking is somwlrat rud y today's stan-
td Works, vol.4). London:
dards, Drwin r.rrksirrlportnt points' still vlid' spially drawing
ttntiontl th sinrilritistwndiffntkinds of organisms (wlrt today
t' Rd ill ot'th nd ltu. is all]homologis)and to th possiilltv tlrt n-rrologial
dvlopr.nt is
a amo for whirt happns to spis througlr th ags. ausally, Darwin
flatd th ida that aquird hratristisr-night inhritd (what today
o dp.rtindfinitly from th
is alld Lamarkism) as wll as offring a vrsion of his grandson's mha-
: Lil Soit, Zoolog
-.R. nism of sxual sltion.
Truly, howvr, flr rasmrrs Darwin, as for r,ry othr arly volutionist,
th hang of orgnisms ovr tim was takn to a rfltion of and prof
of hang in soit'vovr tim-hang from a sinlpl stat to on mor ad-
vand,that is to s.1'harrgof a progrssivnatltr.As 1rusd to say, fom
Darwin' was on of th st- monrh (utflv)tl t-t-tot-tarh
(king). This lrstlrrough larl in on of
ld in Lithfild in th rrglish Darwi's lilt poms. T mpl of Ntur.
llands. H fathrd thr sons
)ganiLif nathth sholsswavs
mistrss'and svn mor hil.
W:rs orn nd nurs'd in Oan's parly avs;
n t h u s i s mf o s u l a f i i v i t y ,
First forms minut' unsn y sphri glass,
l g i v u p d i n k i n gl s ri t i m p i
Mov on th mud, or pir th wtry mss;
txonomy' asd s it was on
Ths, as sussivgnrationsblom,
:hwidr puli as a pot who
Nrv powrs aquir, and larg lims ssum;
,uesof th Plts (1789) ing .Vhrr
ountlssgroupS of vgtationspring,
And rathing ralms of fin' and ft,and wir-rg.
trialistsinluding pottr Josiah
's
third son, Rort' and on of T h r r st h t a l l O a k , t h g i n t o f t h w o o d ,
. d i n m h i n sa n d t h i r u s s , \Whih ars Britnnia's thunds on th flood;
.Whal,
Soity, a group of mn who Th unmasurdmonstr f th main'
This goup inludd hmist Tlr lordly Lion, monarh of th plin,
r, and Jams l/att, famous for Th agl soaring in th ralms of eir,
rwin was himslf an invntor' Whos y undazzld drinks th solr glar,
arriag'and a anal lift for Imprious man, who ruls th stiirlrowd,
510 Dri, rsmus

Of languag' reason' and rfltionproud' Dnai d th n

.i7ith row rt whtl sorns this arthy sld,
Nlind nd Behuiot
And styls himslf th imag of his God;
I is Drouin d the ,
Aros from rudimnts of form and sns,
h u i o r ( l 9 8 7 ) . R o r tJ
An mryon point, or mirosopi ns!
(Darwin 1803, 1: anto I, lins295_14) that hovr around biolo
t l l i g n .r s o n ,h b i
In arguing thus, rasmus Darwin was gling against a ilially inspird ar frquntly usd to rn
aount of origins, ut th atual disagrmntwith Gnsis Ws nvr a ma. and thr is nothing lik
jor impdimnt in th ys of hristians, most of whorn y his day W not an ths human ha
litralists (in th sns of tody's rationists).or ojtionalws th thmslvs funtin as
fat that progrss' whih dpnds on th fforts of individual humans, was swfs to ths two qust
sn to onflit with th ida that salvation an om only through Provi- as rasmus Darwin' Pi
dn, or God's unmritd gra. That onflit did mak volutin a radil through hrls Drwi
ida' ompoundd y th fat that th vil dltrin ws sn tl hav ld to latr Darwinians suh z
th Frnh Rvolution. Darwin himslf ws dist, somOnwho livsin Baldwin, and William.
God s an unmovd movr. Th fat that volution is groundd in th rul of ntury.
law would hav n, for him, a proof of God's xistn and powrs rathr As always' harls D
than a rfutation. no th sol' mhnis
In th nd, rasmus Darwin's rputation was dstroydy a savagparody human traits. Howvr,
of his potry writtn y politial onsrvativs,inludirrgGlrgnning, a uralisti. Som of Drw
British statsman during th Napoloni ra and latr, rifly, prim minis- did not. !7alla gan
..Th Lovs
tr. In th funny of th Tringls'' (1798), Dawin's spultions gnratd y natural s
aout plant suality and inlinations (in an arlir pom' The Loues of t joind with Hrrt Spt
Plts) wr mokd. Thr Wr rtainly thos whl studid Drwin's idas A ommon lif am
with r, inluding th agd Immanul Kant in Konigsurg, who rad a i a n t h o y r u s h dn i n
Grman translation o{ Zlllmi. Nvrthlss, gnrlly, nthusiasm frr his vrs and purgd nturt
thought wnt into stp dlin. Prhaps somwhat unfairly, for postrity h on oth ounts. Prha
taks sond pla to th latr, and no mor sophistiatd,writings of Frnh lifs of fundamntlis
iologist Jan-BaptistLamark. l i s t s .D a w i n a n d h i s d
thnti moral sns.
BILIoGRAPHY
Th sond issu o
anning,G., H. Frr,and G. llis' 1798.Th lovsof th triangls. Ati-Jobi, nisms for volutionary
16 April, 23 April, and 17 May.
topi has n usually
Darwin,F,,1789.ThBotiGrd(PrtlI,The I-ousof tb Plts).l-ondon:
win, ut Rihards shov
J.Johnson.
[1794_1796]181. Zlomi;or, h I's of rgi l'if. 3d d. ntury to th lat tw(
London:J. Johnson. onrns' ut latr Dar
183.Th Tm1llof Ntr. Llndon:J. Jlhnsln. did, at last not until t
King-Hl,D. 1963. rsmsDr,i:Grdfthrof hr!sDuitt. Nw Ylrk:
win, som animals' wh
Srinr.
Nil, . 1987. Udrth Brof Sic:rsmsDri d His ge. mnt movs out from l
anhstr:anhstrUnivsityPss. haviors. Ovr sv
Rus,N4. 1999.str lf stris:ls utllutirl Srlilrstrtio| into nw haits. Howl
amridg,MA: Harvrd UnivrsityPrss. -.R. tndd to rjt th Ba
vry minor signifian
Baldwin fft hs
Darwin originally thor
 D:rrwin and th mrgr-rlf vllutionary Thlris 511

)tionroud' Drt,,innd th E,mergncof uolutionr Theories of

his arh1, sod, ind ad Bhuior (Rort J. Rihads)
rf his God;
t and sns, |nhis Drcui d th mrge of uolutirr Thoris of id d B-
:optns! huior (|987), Rort J. Rihards ddrsssth most diffiult st of idas
s2 9 5 - 3 1 4 ) that hovr around ioiogizrlvolutiorr-th volution of rrrind,insrint, in.
tllign,rason' haits' nd havior.Ths idas ar diffiult austhy
lg aginst a ilially inspird
ar frqr.rntly
usd t invstigatthmslvs.Rihards also addrsssmorals,
t with GnsisWas nvr a ma-
nd thr is nothing lik mols to intfwith th study of nrorals'
of whom y his dy Wr not
an thshumn haratristisvolv via natural sltion' and an thy
). or ojtionalWas th thmslvsfuntion as a mhanism in volution? Rihrds tras th an-
l r t so f i n d i v i d u a l h u m a s ' w s
swrsto thstwl qustionsin th works of suh ightenth-nturysholars
an oll-lonly tlrrough Povi_
as r:rstnus Darwin, Pirr-Jan irnis, and Jan-Baptist d Lmark,
t did mk vllutilna radil .Walla,
through harls Darwin' Alfrd Russl and Hrrt Spnr, and
)trlnWas sn tO hav led to
latr Darwinians suh as Gorg J. Romans, ' Lloyd organ, James I\4.
list,solnonwho livsin
Baldwin, nd \WillirnJanrs, ad on up toward th nd of th rwr.rtith
rtion is gou,nddin th rul of
ntury.
l's istnand powrs rthr
As always, harls Dawin viwd natural sltionas th hif, although
not th sol, mhnism oprating in volution, ir-rludingth volution of
s dstroydy a sirvagparody
human traits. Howvr, ll of thsmhanismswr for Darwin totally nat-
; .i n l u d i r l G
g o r g a n n i n g ,
uralisti. Som of Darwin,s ontmporarisfollowd Darwin's lad, ut most
r n d | t r ' r i t . l yp. r i m m i n i s -
did not. \Valla gan thinking tht tlr pulirly humarr traits ould
(1798 I' D wit-t'sspuiiltions
gnratdy naturl sltion.Latr h andond Drwin on this sor and
larlir pom' Th I'oues of the joind with Hrrt Spnr.
'swho studidDarwin's ids
A ommon lif among prsnt-dayhistorins of iology is that Darwin.
rt in Konigsurg, ',vho rad a
ian thory rushd nintnth-nturylif in spiritully domintd uni_
s, gnrlly,nthusiasmfor his
vrsand purgd natur of intlligntdsign nd purpos. Rihrds disagrs
wat unfairly, flr postrity h
on oth ounts. Prhaps latr Darwinians did not hold th unsophistiatd
phistiatd,writings of rnlr
lifsof fundamntalisthristians, ut thy wr hardly ralitrnt mtri-
alists.Dawin and his disiplsttmptdto infus human natur witlr an au-
thnti moral sns.
Th sond issu onrnd suh things as hal.litsfuntioning as mha-
s lf th triallgls.nti-Jobi,
nisms flr volutionary h:rng.Amng historians of sin irr tlr pst, this
Lrlus f th Plts). London: topi h:rs n usually limitd to parodis of Lamark and latr Jams Bald.
win, ut Rihards shows tht suh lifswr ommon frlm th ightnth
us of ()rgtti I'if. 31d. ntury to th lat twntith ntury. In fat, it was on of Darwin's main
onrns' ut latr Darwinins did nt pay as muh attntion to it as Darwin
hnson.
did, at lst not until th nd of th nintnthntury. Aording to Bald-
f hrls Dri. Nw York:
win, som animals, whn thy ntr a nw nvironmnt or tir old nviron-
rsmusDttri d His g. mnt movs out irom undr thm, zrrradjust to th harrgy larning nw
havios. Ovr svral gnrations ths haviors might transmutd
l Soil ostrutillt? into nw haits. Howvr, in th twntith ntury, volutionary iologists
-^4.R.
tdd to jtth Baldwin fft as not oLlrringat all or s ing of only
vry minor signifian.During th past fw dads or so, howvr, th
Baldwin fft has xprind a rsll!]n.ay it is as impotant as
Darwin oiginally tlrought.
512 Dtukis

In Drtc,i nd tb mrgec of uolutionr Theories of id d B- mhins. mphasizing t

huior, Rihards spouss adial philosophy of histoy-a seltion m n t l i r l s i g h t sD
. awkirl
modl. Historians of sin ar wll awar that the gnalgy (who got what nattlral sltion atd;
from whom) of idas is important, ut ronstruting onptual gnalogis that inludd languag ar
is quit diffiult. Usually historians hav to sttl for similarity of idas, and of volution y natural s
too oftn, when sirrrilarity and gnalogy onflit, thy sid with similarity. ation of salint ulturirl
Rihards dos not. not only spouss a sltion modl of sintifi faturs of ulturl volu
hang ut also Ptlts it into prati. His history is all th ttr for it. prsntd th idil of m
stratnss of th fun
lLIoGRAPHY
limd to making a n
Hodg,J., ad G. Radik, ds.2003' T,hmbridgeompioto Dri. volution of human ult
amridg:amridgUnivrsityPrss. thusiastially' with mix
Rihrds,R. J. 1987. Dri d th l|mrgcof uolutiottrThorisof id In 1986' Dlvkins put
nd Bhuilr.Chiago:Univrsityof hiagoPss.
Wtbmker, in wlrih h
1992. Th ig of uolution:Th orphologiIostrutiond
IdeologiIR>nstrutioof Drlt,i'sTheor.Chiago:Univrsityof hiago v o l u t i o n a r r h l r yv i
Prss. th arly triumphs of th
|999, Te pistmologyof historialintrprtation:progrssivityand 1995' h was appointd
rapitulationin Drwin'sthory.In R. rathand J. ainshin,ds., li Undrstanding f Si
pistm()logd B ioklg, 64_9.Cmridg:CambridgUnivrsityPrss'
talnts' h has subsqu
Rlts,R. 2003. Dluin nd Design:Does uoltion ue Purpos?ambridg'
A: Harvard UnivrsityPrss. plaining asptsof sin
Wr,B. ., and D. J. Dpw. 2003. Li,uolution
d Lrig:Tb Bldu,,iEffet th puli disussionso
Ronsiderd. mridg' A: MI Prss. -D,L.H. N4ost rntly, his roa
tmt to rais onsiou
rligious lif,has attr
Dawkins, Rihard (. I94|) a l a d i n g r o l i n w l r a ti s
Rihard Dawkins is a British zoologist and volutionary thorist st known th lay puli ar Dawk
for his onpts of th slfish gn, th xtndd phnotyp, and th mm. kin sltion and th v
Born in Knya, Dawkins was duatd at Oxford' whr h has livd and
workd sin 1970. F{is 1976 Th Slfish Gee is on of th lassi works of lLIoGRAPHY
no-Darwinism, unifying and laifying th mgingonsnsusmong suh Dawkins,R' 1976' Th S
volutionary thorists as !(/illiam Hamilton, Gorg Williams, Rot Trivrs, |982. The tnd
1986.h Blid \
John aynard Smith, and his dotoral suprvisor, thologist Niko Tinbrgn.
204.Th cst
This ook artiultrs th thory of ntural sltion in suh vivid and assi-
26,Th G<ldD.
l trms that it is oftn mistaknly rgardd as mrly a ..popular'' ook. Gafn,., and M. Ridlv
Dawkins's ntral laim is that th most fundamntal' and hn most x- tb W W hittk.
planatory, prsptiv on all volutlon by natural sltion is th .,gn-y
viw,,' in whih nfits to spis, linags, groups' and vn individual or-
ganisms ar sn to susidiry to th primary nfiiaris of all adapta- Th Descent of
..slfish'' gns
tions' th thmslvs. ovr tim' vlution designs and uilds Reltio to Se (|
organisms (..survivalrahins'')that nfitoalitions of gnsy improving
thir prospts for rpliation. T ah of gns dos not stop at th skin of Tb Dstof ,
th oganism: sol gns ontrol th dsign and onstrution of an xtndd ws harlsDarwin'sv
phnotype' harnssing faturs of th nvironmnt and vn othr spis. h had nunwillingt
Thus th bav'sdm and th spidr's w ar just as important pats of th whrrAlfrd RusslW
phnotyps of thos spis as thi ys and mouths' nd parasits oftn - that hr-rmans ould not
ploit th haviorl ontrols of th host spis' hiiaking othr gns' survival tht h had to gtinvol
 IrDsntof Man .'1.]

r Theories of ind d Be. mahins. mphsizing th univrsal appliation of harls Darwin's funda-
ophy of history-a sltion mntal ir-rsights,Dawkins drW attntion to th possiility th;rt whn gnti
rt th gnalogy(who got what naturl sltion ratd a spis,Hoto spies, with an xtndd phenotyp
ruting onptual gnalogis that inludd languag and thnology' this in fft stablishd a nw mdium
rtl for similarity of idas, and of volution y natr'rralsltion:human ultur, in whih th diffrntial rpli-
flit, thy sid with similarity. ation of salint ltual itrr-rs'alld ms' uid aot-tntfo rrrany of th
sition modl of sintifi faturs of ultural volution that ar Othrwis prpling. Dawkins initially
ry is all th ttr for it. prsntd th id of mms as a sort of thought xprimnt to illustrt th
bstratnssof th fundametal ida of natural sltion. nd h has not
laimd to making a major ontrir-rtionto th sintifi invstigation of th
gompioto Drtui. volution of humn ultur. Othrs, howvr, hv takn up th onpt n-
thusiasti:rlly,with rid rslrlts to dat.
{ ullutionrTboris of ind
In 1986, Dawkins pulishd anothr pdggialtour d fore,h Blid
Prss.
phologiclonstructiod Wthkr,in whih h illr.rstratd many of th most lusiv impliarions of
r,.hiago:Univrsityof higo volutionary thoy via his pionring Blind \Wathmakrsoftwar, on of
th rly triumphs f th Artifiial Lif 1ovmentin thortial iology' In
rtation:pogrssivityand 1995,e was app<lintdto th first Charls Sirnonyi Profssor of th Pu-
. andJ. ainshin,
ds., li Undrstanding of Sin t Oford. In this idal rol for a writr of his
l ambridgUnivrsityPrss.
't Hu Purpos?mbridg' talnts,lr has susquntlypulishd a srisof luid ooks and atils-
plaining sptsrfsinto h gnralpuli. L{ also has takn tlr lad in
d Lrni g:'', u o.:([:,n. th puli disussions of ontrovrsial issus of sin poliy and politis.
"'
Most rently, his roadsid on rligion, Th God Dlusio (2006), an at-
tmpr to rais orrsiousnssout th waknsssand follis of traditional
rligious lif,has attratd oth nthusiastipris and vilifiation' playing
a lading rol in what is bing alld ..thnw tlrism.''Lss wll known to
llutionry thorist bst known th lay puli r Dawkins's important analyssof fundamntal prolms in
ld phnotyp, and th mm. kin sltionand th volution of ornrnuniation,among othr topis.
ford, whr h has livd and
z is on of th lassi works of |BLIoGRAPHY
lrging onsensus mong suh Dawkins, R. 1976, Th SIfsll G. oiord: oford Univrsit1'Prss.
lorg!illiams, Rort Trivrs, 1982.Th tededPhotp.Oford: Oford Univrsitrss.
or' thologistNiko Tinrgn' 7986. Th tslid WtbmaAr.Nlv York: \W.W. Norton.
;tion in sh vivid nd ssi- 204.Tb ttt,stlr's
?ll/c.Boston:HlughtonNlifflin.
206.h (]ldDlsitl.Bostorr:ughton ifflin.
as mrly a ..popular'' ook.
Grafn,A., nd M. Ridly,ds.2006.RihrdDtukis:Htttu Sitist hgd
lamntal, and hn most - th W |XtTik. ofrd: Oxford Univrsityrss. -D.D.
ral sltion is th ..gn's-y
:oups, and vn individual or-
rry nfiiaisof all adapta.
The Descent of NIn, nd Selectio in
:, volution dsigns and uilds
'litionsof gnsy improving
Reltio to Se (harlsDarwin)
Lsdos not stop at th skin of Tbe Dst of , d Sltili Reltio to Se, pulishd in 1871,
d onstution of an xtndd was hrls Darr.in'swork on th evolution of th human spis.Initially,
mnt nd vn othr spis. h hd r"rnwillingto gt into this ast of tlr volutionary dbat, ut
iust as importnt parts of th whn Alfrd Russl !7all (1870), who had om a spiritualist, dlard
loths,and parasitsoftn - that humans ould not hav n produd y natural mans' Darwin flt
hijking othr gens' survival that h had to gt involvd.
514 Th Desntof n

Littl of th si disr-rssionof human volution is originl,

s Darwin r- promotlng an unsllpPol
lid havily on th rports and findings of othrs. Lik Thomas
Hnry Hu. Vitorin, h is a vry l
ly (in Euiden s to 's Pl i Ntur, 186) for
him, Darwin f f t so f m d i i n ,h i l
hif mod of rgumnt was to ompar humans with othr
animals, show- stop using it. H also t
ing that th diffrns ar a lot lss grat than ln right imagin.
Thrfor, ovrom othr popls
thr is no rason to think that humans ar not part of the naturl
world, whit popl mor ap1
and, givn th shard faturs with th animals' mh rason
to think that or grrrally,althouE
humans ar thir dsndirnts.Dwin did not think tht
humns wr d. tht do sound lik So
sndd trom animals aliv today, nd h was inlind to support
th ida of on many sours, and
an Afrin origin for humankind.
Volkish sr-rtimntsof t}
Darwin was prtiulrJy intrstedin humn fason nd
soiality. H ar- last as signifiant, or m
gud that morality (nd riigion) ar nattrraloutoms of
volution. Baus th Nazis grrrailydid
in arlir writings Darwin had livd that natural sltion
works always nss of humankind, linl
1r th bnfit of th individual rathr than th group' h
was onrnd to disusss ths hrgs
show that a gnuin morl snsof r for othrs ould nvrthlss
volv.
In som rspts' thought that a group kind of sltion might hav pro- lLIoGRAPHY
-h
dud morality' ut h also gav th suggstion tht morlity D r w i n , ' 1 8 7 l . T h D
.oulJ .
nlightnd slf-intrst. Hr h antiipatd th modrn
mhanism of ..r- John urray.
ipoal altruisnr..' pIld:No Itlligcll,
!alla'sapostasy also plains th rathr odd ntur of Darwin's N.p.: Pnrisdi (
ook, F r i t l l a n d S '. l 9 9 ' . N .W
li
baus muh of it was not aout humans at ll. It was rath
about Dar- 19.1-9. London:
win's sondary mhanism of volutionary hang, that is,
sxual sltion. G r u r , . . | 9 8 1 .D r u
Hr, organisrrrsin a spis ompt for mats, and hn H i t l r ,A . | 9 2 5 . i K l
thy dvlop
adaptations (.g.,antlrs of the dr, tail fthrs of th paok) Flu.ly,T . H. 7863.uid
to id in
this omptition. !alla argud that faturs lik humn hailssnss Norgat.
and
grat intllignould not hav bn produd by natural Krshaw,I. t999. itler.
sition nd so Rihards,R. J. 2008.
must b th rsult of spirit fors. Darwin rplil that
sual sltion uolutior hough
auss many human fatus'suh as intllignand ody Rus, . 2008. hrlesD
shap. For marry
yars' popl wer inlind to dismiss sxual slrion Trivrs,]t' L. 1971.Th
s somthing d-
manding tratmnt in its <lwnright' nd thought of it as somthing Biollg46: 5-57,
st in- .Wlla,
orporatd with natural sltion.With th ris of intst in A. R. 1870.Th
anima] soial otributiltls ttl tb
havior, sxual sltion is now somthing of grt thortial
and mpiri- Mmillan.
l intrst.
Wikt,R. 2006. FrlmL
In th ours of th disussion, Darwin was ld to mak Rism i Crm.
som vry on-
vntional Vitorian judgmnts aorrt th superiority of rrrals
ovr fmals,
and uropans ovr all othr humans. H also worrid that modrn
mdi-
in was having dltrios ffts on human ding stok,
ut flt al
Th Duelopment
to dfnd apitalism aus it lads to lisurd p.opl. -be
of ultur whos Dulopmt of l
works bnfit us all. Rntly, on th asis of rhs sorts
of rmarks, th just a fw months aftr
Amrian rligious opponnts of volution (th rtionists
and thir asso- Th ook hivd n
iats' th so-alld Intllignt Dsign Thorists), frustratd
in thir at- win Studis, th histor
tmpts to pov tht thir ilially asd lifs are gnuin
sine, hav Studis nr spi
trid a diffrnt tk. Thy argu that Darwin lads straight
to th vil id- ntnnial of th puli
olgis of th twntith ntury, partiularly NationI Soialism
(ssp. soon grWino majo
ially \Wikart 206 and th Bn Stin-narratd fiIm
plld: No du to two fators. Fi
Itellige llou',ed' 200s). This is a lassi xampl of
ad sholarship xtrnrly importnt s

llution is original, as Drwin r-
hrs.Lik Thomas linry u-
r' 186) for him, Darwin's
nans with othr nimals' show-
n o might imagin' Thrfor,
) not part of th natural world,
nals,ruh rason to think that
rot think that humans wr de-
s inlindto support th ida of
lan asonand soiality. H ar-
outomsof volutiorr.Bause
naturl sltionwoks lways
th group, h was onrnd to
lthrsould nvrthlssvolv.
Lndof sltionmight hav pro-
]stion that morality ould
th rodrnmhnism of ..r-
r odd natur of Darwin's ook,
at ll. It was rathr aout Dar-
hang,that is, sxual sltion.
mts' and hn thy dvlop
rthsof th pok) to aid in
:s lik human hirirlssnssand
rdy natural sltin and so
n rplid that sxual sltion
nnd ody shap. For many
ual sltion as somthing d-
Lrglrtof it as sornthing st in-
ris of intrstin nimal soial
of gratthortial and mpiri-
las ld to mak som vry on-
iorityof mals ov fmals,
lso worrid that modrn mdi-
rn rdingstok, ut flt al
sud popl of ultur whos
of ths sorts of rmarks, th
t h r a t i o n i s tas n d t h i
:orists)' frustratd in thir at-
lifsar gnuin sien, hav
in lads straight t th vil id-
y National Soialism (s sp-
-nrratd fr|m plld: No
i arnpi of d sholarship
Th Dvlopmntof Darwin's Thory 515
promoting an unsuppotal tho|ogial thsis. Although Darwin is vry
Vitoria' he is a vy liral Vitori:rn. For all that h worrid aout th
fftsof mdiin, h immdiatly strssdthat lr did not think w should
stop using it. H also thought that th hif rasorr whit popl tnd to
ovromothr popls is baus othr popls suum to th disassof
whit popl mor rpidly than whit popl do to th disass of othrs.
or gnrally' although thr indd pssagsin ei Kmpf (1925)
that do sound lik Soial Darwinist' Hitlr drw somwhat haphzardiy
on many sours' and th nti-Sritism of his youth in Vinna nd th
.Wagnr
Volkish sntimnts f th group promoting th mm<lryof wr at
lastas signifiant,or mor (Fridlandr 1997; Krshaw 1999).In an as,
th Nazis gnrally did not muh lik volution aus it strssdth on-
n s so f h u m a n k i n d , i i n k i r r gA r y n s w i t h J w s a n d G y p s i s . ( R i h a r d s 2 0 0 8
disusssths hargs' putting thm in ontt.)
|LIoGRAPHY
Darwin,. 1871'.The Dstof , lld Seltilli RItil,oSe.London:
Iohnuray.
plld:No lntllignclltl'd,2008.NthrrFrnkowski,dirtor.90 mirr.
N.p.:PrmisN{diaorporation.
Fridlandr,S' 1997. Nzi Germd the |es.Vol. 1, Th Yrsof Prseution,
193_9.London:Widnfldand Niolson.
Grubr,. . 1981.Dri o .2nd d. hiago:Univrsitof hiagoPrss.
Hitlr,A. |925. in Kmpf.Munih:VrlgFranz hNahfolgr.
Huxl,' H. 1863.uir'lc s ttl 's Plcei Nture'Lon]on:l/illimsand
Nrgat.
Krshaw, |. 1999.itlr.o|.1,1889-1L).J6: I]ubris.Nr'York:Norton.
Rihards'R. J. 2008. h Trgi Sseof Life: rst lIhld th Struggllur
uolutiorThoght' hiago:Univrsityf hiago Prss.
Rus,M. 2008. brlsDnui. Oxford: Blakwli.
Trivrs,R. L. 1971.Th volutionof ipro:rl altruism.QurtrlRuituof
Biohg 46: 35_57.
Wll,A. R. 1870.Th limitsof nattlrlsltionas applidto man.ln his
otributios to th Thorof NturlSltio, 332-71' London:
amillan.
Wikart,R. 2006. From Dui to Hitlr: uoltiorthis, ugis,d
Rismi Grm\,,Nw York: Palr]rar. \4amilln. -.R.
The Deuelopmet of Dui's Theor (Dov ospvat)
Th Deulopmet of Dri's Thor ws pulishd in 1981. It appard
justa fw months aftr th dath of its author, Dov Ospovat, at th ag of 33.
s s o - Th ook ahivd an important milston in what am known as Dr-
win Studis,tlr histoil study of harls Darwin's lif nd work. Darwin
Studisam a spial ara in th histlry of sirrshortly aft th 1959
ntnnialof th puliation of hrls Darwin's o the origin of Spis.It
soon grw into a mior fild of historial rsarh.Th xplosiv growth was
due to two fators' Fist, until th 1950s Darwin was l.lotrognizd as th
xrmlyimportant siIrtisttht h is today. Th modrn synthti thory
516 Th Dvlopmntof Darwin's Thry

lf volution (th thory fomd v omining Drrvinian natual sltion This nw viw lf Dar'
with Mndlian gntisand, m<rrrntly' with rnolular gntis)gan pholog and mryoltlg
during th 1930s nd 1940s. At Iirst it dm<lnstrtd tht a numlrr of diffr- ginning in th 1980s, wl
nt influns' inluding natural sltion and gnti drift, ontriutd to and mbryologial dvlo
volutionary hang' ut th rlativ strngths of ths influns Ws un- th stag fo roadr his
known. By th 1950s it Was gnrlly ptdthat r-raturalsltionwas th gathrd y historian D;
dominant voltionry for. This onsnsusoinidd nily with tlr l- Th D r'itti Her itg
rtions of th ntnnil of the rigi. B:lt'ts Drwin Was th fathr of portant was Ospovat's s'
natural sltion,h now am th fathr of th synthti thor of volu- o m l n i t t d t o t h n o t i r l nt
tion. Th nw interst in Darwin oinidd with th disovry tht Darwin jor intrst to historins
hd lft an immns sto of prsonal paprs that inludd prsonl orr- 2 0 0 3 ,2 0 0 4 ) .
spondn, rsrh nots, and prliminary drfts f his ooks and artils.
Urrlik Gr;orMrrdl (rvhospps wr dstoydy lris stlssor))Dar- lLtoGRAPY
win's paprs offrd immns opportunitis fo rsarhin th histlryof si- Amundson,R. 200.5.TDe/
n and, indd, in th lif story of a sintist whos importan was Rlotsof uo-duo.al
suddnly ing rognizd. Burian, R. . 2004. list
Th first wav of Darwin Studis onntrtd on his voyag on MS tis. amridg:lr
K o h n , D . , d . 1 9 8 5 . h D
B g l a n d h i s n o t o k s o f 1 8 7 - 1 8 9 , w r i t t r rs l o n a f t h r t t r r r r d .
Prss.
T.his priod ovrd th girrnirrgslf Darwin's volutionary thought and O s p o v tD , . 1 9 8 1. T b D e u
his dvlopmnt of th prinipl of natural sltiln.This phas of Dar- UnivrsityPrss.
win's lif was importnt to sintists aus natural sltion was th as- R i h a r d sR, . J . \ 9 9 2 ' h N
pt of Darwin's thought that fit most losly wirh th synthti tlrory of d ldollgiIR ls
hiagoPrss.
volution. Th priod from 1840 to l859 was oftn rgardd as a pr-rzzl. If
200, he Rlmtltt
D r w i n h a d d i s o v r d n t u r l s l t i o r r f o r 1 8 4 0 , w h , v d i d h d l a y of Glth. l.rillgl: LII
pulishing until 1859? ospovat;lrovidd an llswr to that qustionnd' 2004.MihalRus
simultanously, gan th study of Darwin as a lnAn lf his ag-a Vito- 2s-38.
rian naturalist. Rus, . 1996. od to ]
arnridg, MA: rv
Ospovat was bl to provid this nw prsptiveaush was tlr first
2004.h r<lmatt
historian to gain assto Drwin's paprs from th so-alld lak <l,th
isttt lf tsiolog.r'7:
st of portfolis that Darwin had ssmldin th 1850s in ppratiOnfor
writing his major work on volutiln.(This work ',vas,of ours' intrruptd
.Walla's
y Alfd Russl odisovry of natural sltiln,and the ri.
Dinosaurs: Th mor
gi lf SpiesWas th rsult. S th main ssay ..Th History of volutin-
ry Thought'' by ihal Rus in this volum.) Th fist pratitionrs of Bas of thir immt.t
Darwin Studis hd sn ntural sltiorras Dirrwin's major lrivmnt first ol to mind wl-r
and intrprtdim as uniqu gnius.Ospovat, in ontrast,dpitd him as pld with th fat that
having bnfitdfrom wid adingsin th sinof his day, inluding pal- ntury ngland) wlrr
ontology' systmatis,morphliogy, and mryolgy. This plaind Dar- t h i r h i s r o r i e l l y i n gu
win's hsitation. vn though n.rtural sltin ws in pla, Darwin's t t , d i n o s u r sh a v p I
vidnthat volrrtionhad atuall .ltrrrd Was rlarivlwak in tlr rly our volution. Nvr
yars' It was gradually strngtlrndduring th two dzrdsl>for1959. d v l o p i n g m o s i
..Darwin was and undrstandingvol
th first rputal lraturalist to argu for transmut.ttion who
was l to tak advantagof th dvlopmntalviw of natllr that mrgd Dinosaurs ar a unlq
in th mid-nintnth ntury. This, as nruh as lris nw thory [natural thir rt postu and '
sltion],sprtsDrwin's work frlm [Jan-Baptistll,amark's'' 1Ospo. most sLlssfultrst
v a t 1 9 8 1 .i 6 8 ) . spirnnd th piod fror

rg Drwinian natural sltion
with molular gntis)gan phology and mryology was influntial on svral philosophrs who, -
stratdtht a numr of diffr-
d gntidrift, ontriutd to
hs of ths influns was un-
:d that natural sltion Was th
; oiniddnily with th l- The Drtuiin eritge, whih ws dditd to Dov Ospovat. Vry im-
:ausDrwin was th fathr of
lf th synthtithory of volu-
with th disovry that Darwin
s that inludd prsonal orr-
draftsof his lks and artils.
lstroydy his sussor),Dar-
:or rsarhin th history of si-
intist whls importan was
ltratd on his voyag on HMS
writtn soon aftr h rturnd.
win's volutionary thought and
l sltion.This phas lf Dar_
s natural sltion ws th as-
ly with th synthti thory of
'as oftn rgardd as a puzzl. If
-l840,
lfor why did h dlay
Ln answr to that qustion and'
l as a man of his ag-a Vito-
sptivaus h was th first
:romth s-alldlak ox' th
l in th 1850s in prpaation for
work was, f ours, intrruptd
rtural sltiln,and o th ori-
..Th History of volution-
:ssay
lum.) Th first pratitionrs of
as Darwin's major ahivlnt
Ovat,in ontrast' dpitd him as
;inof his day, inluding pal-
nyology. This plaind Dar-
ltion was in pla, Darwin's
d ws rlativlywak in th arly
g th two dads for 1959.
to argu for transmutation Who
ntalviW of natur that mrgd
tuh s his nw thory [natural
an-Baptist]Lamark's'' (ospo.
Diosurs 517
This nw viw of Darwin as a bnfiiary of nintnth-ntury mor-
ginning in th 1980s' wr onrnd with th rlation twn volution
and mryolgial dvlopmnt (Amundson 2005; Burian 2004).It also st
th stg for roadr historial undrstandings of Darwin, suh as thos
gathrd y historian David Kohn into th monumntal 1985 anthology
portant was ospovat's strongly supportd laim that Darwin was firmly
ommittd to th notion of iologial progrss, a topi that has n of ma-
j o r i n t r s tt o h i s t o r i a n s i n r n ty a r s ( R u s 1 9 9 6 , 2 4 1R i h a r d s 1 9 9 2 ,
2 0 0 3 2. 0 0 4 ) .
BIBLIOGRAPHY
Amundsln, R. 2005. Th hgig Rol of tb mbro i ullutior Biolog:
Roots of uo-duo. amridg: mridg Univrsity Prss.
Burian, R. ' 2004. 1listmllogil Pprs o Duelr\lmet, uolutio, d G-
tis. amridg: amridg Univrsity Prss.
-l
Kohn, D., d. 985. h Drii Hritge. Printon, Nj: Printon Univrsity
Prss.
-he
ospvt' D. 1981 . Deulopmt of Drtuin's hor. mridg: ambridg
Univrsity Prss.
Rihards, R. J. 1992..h nig of uolutil: The orphologil ostrutio
d ldologil Rlstrutio of Dri's hor. hiago: Univrsity of
hiago Prss.
2003. The Romti lptil of Lif: Sci d Phiklsopb i te g
of Goethe. higo: Univrsity of hiago Prss.
2004. ihal Rus,s dsign for |iving, Jourl of tb Histlr of Biolog 7:
2s-38.
Rus, M. |996, r-ld to : h opt of Progrss i uolutior Biolrlg.
amridg' : Harvard Univrsity Prss.
2004. Th romanti onption lf Robrt J. Rihards. Jurl of th
Histor of Billog7: -2' -R..
Dinosaurs:Th modl systmfor volution
Bausof thir immns popularity, dinlsaursar oftn th organisms that
first om to mind whn on thinks aout animals from th past. This, ou-
pld with th fat that thy wr disovrd at th pla (mid-nintnth.
ntury ngland) whr modrn volutionary thory took root, has ld to
thir historially ing usd s xampls in volutionary dbats.In this on-
tt, dinosaurs hav playd oth positiv and ngativ rols in argumnts
aout volution. Nvrthlss,dinosaurs rmain an important atalyst for
dvloping a mor sintifiallylitrat soity, a rquisit for appriating
and undrstandingvolution.
Dinosaurs ar uniqu linag of rptils st apart from othr groups y
thir rt postur and distintiv hip and hand strutur.Thy ar also th
most sussful trrstrial animals to vr inhbit th ath. Thir tnur
spannd th priod from 225 to 65 million yars go. Furthrmor' if ids
' 18 Dilsrtrs

ar in ft a linag of rnivorous dinosaurs (sfigu on pag 520), thir l a R i h a r d O w n , w l

rign atually tnds anothr 65 million yars to th prsntday, whr thy 1863), ah dsrid
dominat th arial ralm with ovr 9,000 spis. tionary thinking, faild
Dspit thir grat suss' dinosaurs had huml ginnings. For in- a l v i a n t r i u sS. u t
stan' although w tnd to think of dinosaurs as giilnts,th arly forms wr Frsh off an Xtnsivs
atually qrrit small (50 to 100 pounds). Ths nivorous ipds wr this spimn's impotir
rathr linor playrs in sozoi ommunitis. Ovr tim thy divrsifid t u r s n d p u l i a t i r l n is
into vrity of terrstri:rlnihs nd in :rll majo linags gigantisnr o- r h <lp t r.r,.is l
urrd, whr som spis am to wigh thr or mor tons. Som di- lutionar missing lirrk.
nosaLlrs'inluding all arnivorous forms, wr ipdal, whras hrivorous So how do dinosaur
forms also inludd quadrapdal taa. On a road sal, dinosaus appar to tim that rhoptr1
hav n quit sophistiatdin thir bhavioral rprtoirs.Physiologially known as ompsogtt
thy do not appar to hav ben lik living rptils. Biologially thy w nosaur provd for th f
perhaps mo akin to today's irds and mammals, thus possily plaining Own hd ontndd
thir grat suss. tutly pointd orrt th:r
Sintifi rognition of nonavian dinosr.rrsourd in 1824 whn that of irds. H tln
Willim Bukland dsrideglosurus (a larg arnivorous thropod di' oth ird nd rpril
nosu similar to Trrtosurus re| from rtaous sdimnts in ngland. rahd th sm on
This vnt was followd by a flurry of nw disovris of othr dinosaurian Llps (= Drptt,sr
giants tlrroughout urop. With thir ovious rptilian affinitis' it was ini- h poind out rhatth(
tially livd ths animals w simply sald_upvariants of living saurian Llps wr alsll vi
groups suh s iguanas and roodils. It was not until 1842 tat this para- Ggnbaur,s findings a
digr-rrws laid to rst whn rnownd Brirish natomist Sir Rihrd Own rmarkal similriti
showd that dinosurs wr a uniqu rptilian linag in and of thmslvs. Using rheo|)trr .
..farfully grat
nmd ths giants th Dinosauria, or rptils.'' own, ommon twn bird
who was a rationist at th tim, pointd out that attributSof ths ani- s a p l o n l u s i ( ) n
mals, suh as rt postur irnd sustantial fusion of th hip vrtra,ar thropod dinosaurs.
not forrnd in xtant rptiIs'ut ist in living irds irnd mammals. Hn, h This viw of avian
onlr"rddthirt dinosaurs W anatomially nd physiologially mor ad- lontologists.Ironiirl
vand than today's rprils.H als ddud that living rptilsar dgnr- only rntly rrd
at forms of a formrly ir{virnd goup' and h qustiondth plausiility of ntury' palontologis
progrssivism and trans-tutationof spis. any sholars liv that tils from whih th l
own ratd th Dinosauria to dunk volutionary thoris that wr at tivs), roodilians, h
th forfront of sintifi dat at th tim. rptils),saurishian d
Svrl dadslatr, dinosaurs wr again to figu prominntly in volu- irds indpndntlv r.
tionary dirts.In 1860 quarrymn in a limestonqu[l.yin Solnhofn, G- irds and dinosaus s
many' disovrd tilthr from rhoptr' his finding showd that within thopods tlr
irds livd at th tim of th dinlsaurs.In 1861, rwo yars aftr hrls Dr- I n t h 1 9 6 0 s ,H u l
win pulished n th rigin of Speis' a nw spimen of Arhoptr, John Ostrom disovr
known as th London spimn, rvald th tru, ovrall natur of this ra- r of th Dromaos
tur. It was undutdly ird aus it sportd wings, fathrs, and a nosaurs with sikI-l
wishon. Howvr, ther was mor to it. It possssdtth, a rptilian tail, narly 200 dinosuri
..missirrg
and lawd hands. Hr was th sptaulr link'' (an animal in- i n l u d i n g n u m r o r t sf t
trnrditbtwn two inlportnt groups) Darwin nd lris followrs ndd arls (s figur on p:r
to prov tht volution Was a rality. Am:rzingly, this rvlation was not ogy whry shard-d
immdiatly ralizd. Grmn palontologistAndras.!agnr(in 1861) and t o s t a l i s h I i t t i o n s
 Dinosurs 519

sfiguron pag 520), thir latr Rihard own, who purhasd th spimn for th British usum (in
to thprsntday, whr thy l863), alr dsrid th spimn. Both' prhaps du to thir antievolu-
lS. tionary thinking, fild to attritany volutionary signifianto th atypi-
humbl ginnings. For in- al avian attriuts. Susquntly,Thoms Hnry Huly ntrd th pitur.
ts giants' th arly forms wr Frsh off n xtnsiv study of avin anatomy and systmatis, h rognizd
lsrnivorous ipds wr this spimrrimportan nd usd it s modl organism in his famd l-
ls. ovr tim thy divrsifid tursand puliationsin support of volution (.g.,Huxly 1s6s). To this day
m a j o r l i n a g sg i g a n t i s m o - rhoptery is lratd as on of th most lar-ut ampls of an vo.
hr or mor tons. Som di- lr'rtionrymissing link'
i p d l 'w h r a sh b i v o r o u s So how do dinosaurs figur into th rheoptr stor? Aout th sam
oad sal,dinosaurs appar to tim that Arbeoptry, WaS disovrd, a hikn.sizd thropod dinosaur
.al rprtoirs.Physio1ogially known as ompsogthgus was also disovrd irr Grmany. This tiny di-
ptils.Biologially thy wr nosaur provd for th first tirrrthat not irli dinosar.rrsw lphant-sizd,as
nals, thus possily plaining own had ontndd. Dvlpmntal iologist arl Ggnbaur in 1864 as-
tutly pointd out that th ankl strutur of ompsognthgus was akin to
urs ourd in 1824 whn that of irds. H lrrluddthat it was n niml of doul lationship-
argarnivorous thropod di- both ird and rptil. Amrin paleorrtologistdwin Dinkr op latr
)toussdimntsin ngland. ahdth sam onlusion rgarding th lrg arnivorous tyrannosauroid
s o v r i os f o t h r d i n o s a u i a n Llps (= Drptrlsurus)'In a srisof paprs pulishd from 1866 to 1869
r p t i l i . r na f f i n i t i s ,i t w a s i n i - h pintd out that th longatd nk vrtraand light skull strutur in
d . u p v a r i n t so f l i v i n g s u i a n Lelps wr also avian faturs. Shortly thaftr, Huly, inspird y
not until l842 that this pra- Ggnbaur'sfindings and apparntly unawar of Cop's rsah,notd th
anatomist Sir Rihrd Own rmarkai similritis btwn th hips of eglosrus and tant irds.
l lingin and of thmslvs. Using rhopter as guid, h was al to idntify many attriuts in
arfully grat rptils.'' own, ommon twn irds and dinosars. By th 1880s h had rahd an in-
ut rhet ttriutsof ths ani- sapal onlusion-I.lot only r ids rptils' thy r in fat living
usion of th hip vtra' ar thropod dinosaurs. rbeoptrr is simply a fathrddinosaur.
irds and mammals. F{n,h This viw of avian anstry is now almost univrsally aptd among pa-
and physiologially mor ad- lontologists.Ironially, dspit ing naly a ntury and half old' it has
that living rptilsa dgnr- nly rrrlyn doptd on a road sal. By th turn of th nintnth
1qustiondth plausiility of ntury' palontologists instad favord th ompting thory of stm rp.
;. any sholars liv that tils from whih th mjor lads of ahosaurian rptils (dinosur rla-
,lutionaythoris that wr at tivs)' roodilians, hrivorous ornithishian dinosaurs, ptrosaurs (flying
rptils),saurishian dinosaurs (thropods and long-neked hrivors),and
to figur prminntly in evolu. irds indpndntly ros. It was flt tht th ommon ftures shard y
ston qllarry in Solnhofn, G- irds nd dinosurs stmmd fom mor anint ommon nstor than
r. Tts finding showd tht within thropods thmslvs.
51,two yars aftr harls Dar- In th 1960s, Huly's harg was rnwd whn Amrian palontologist
3w spimn of rbeopter, John ostrom disovrd wll-prsrvdspimnsof Dinonhus, a mm.
tru,ovrall natur of this r- r of th Domaosauria (small to rndium-sizd, agil rnivoros di-
sportd wings, fathrs' and a nosaurs with sikl-lik laws on thi ft). ostrom and othrs idntifid
possssdtth, a ptilian tail, narly 200 dinosaurian attriuts shard twn irds and dromaosaurs.
..missinglink'' (n animal in- inluding nmrous fusd sral vrtr,hollow lg ons, and long for.
)arwinand his followrs ndd arms (s figur on pag 520). Furthrmr, ladisti analyss-a mthodol-
zing|, tlris rvltion was not ogy whry shard-drivd(volutionailyhangd)haratristisar usd
AndrasWagnr (in 1861) and to stlishrlationships-showd tht th gnsisof ids am from within
520 Diosurs

portd glidr, appars tl

irlrptor, th smal
asal dromaosaur to
volution of irds. Ad<
s a u i a n a n s t r yf o r i l
nonavian dinosaurs is r
lik irds, and that t
nosaur skltons. \Vith
has all ut ndd. Dinol
ologial ordr spara
nosaurs ruld, from mo
astroid or othr ata
outsid today.
Dinosaurs hav plyt
volutionry thory y 1
port of volution-rl
omparison twnth fist trd, Arbelpter,and th dromaosaurian
gnsis.But do thsan
dinosaur, Vlocirptlr.Not th shard faturstwnth ird (lft) and te
nonavian dinosau kisht)' whih inlud upright postur' soktdarnivorous ary undrstanding in th
tth' fathrs'thr or mor fusd hip vrtra,thr main tos and fingrs'n ampls of human and
s.shapdnk, trmly long forlims,an longatdand rtrovrtdpuis (hip srv as a valual vhi
on)with a oot shape'a stiffndtil, urvdfmur (thighon), longatd ntifi mthod and at th
mtatarsals(foot ons),lavils(wishons), a prforatataula(a tunnl heopter story is n(
through th hip ons twnwhr th thighonsattah),n ntorital dtail, although argua
fnstra(a hol in th skull in front ofth y sokt),and hlllowbons,just to
rat. In fat it is just o
nam a fw.
nosaurs rahd lph
for vriant dvlopmn
land situations.'W kn
th Dinosauria. Statistiallyit is trmly unlikly that irds ould hav d- dntal attris (th v
sndd from any othr group of animals. tion with floral hangs
Today, aptanof this thory is almost univrsal. Nvrthlss,during hangs, ontinntal dr
th last 15 yars a fw holdouts (gnrally avian palontologists and or- at th tim of an astro
nithologists) lung to th notion that avian anstry still might found in world.
som arlir stm rptil. Thy ontndd that similaritis twn dino- Th aformntiond
saurs and irds might somhow rflt indpndnt adptations to similar ral attntion in th
nvironmntal sltivfors. Th ru of thir dissnsionWas th asn Through dinosaurs th
in dinosaurs of svral ritial dfining attriuts of irds. Ths inludd roadly promoting v
th avian wishon (fusd lavils or furula) and fathrs,s wll as th gualy on of th most
fat that th smallst dromaosaurs' thought to th anstors to irds, influn in promoting
wr gratr than 25 pounds in mass and smd unlikly andidats for volution gain broadr
giving ris to th muh smallr rcheoptry. Spurnd y thir n- tn than not our hildrt
tntions, palontologists n mass fousd on th study of ths faturs.In and thr is no ttr
quik sussion, ah was rvaled in dinosaurs. Ramination of young' urious, opn m
thropods showd that furula xistd, ut thy had not n prviously r- ar told will b tt.
ognizd. Fathrs Wr found in nonvolant dinosaurs from China. Prsumaly ar told is a rality or :
th fathrswr for display and/or thrmorgulation.on spimn,a pur- important sintifi pu

r, and th dromaosurlan
:s twn th ird (left) and te
h t p o s u r .s k t d a n i v o r u s
t' thI main tos and fingrs, an
lngatd and rtrovrtd puis (hip
,d fmur (thighon)' longatd
a prforatatula (a tunnl
ronsattah),n antrital
; o k r )' n d h l l o w o n s ' j u s t t
rnliklythat irds ould hv d-
;t univrsal.Nvrthlss,during
ly avian palontologists and or_
t anstrystill might found in
l that similaritis btwn dino-
lpndntadaptations to similar
thir dissnsionwas th asn
:riutsof irds. Ths inludd
ula) and fathrs,as wll as th
ght to th anstors to birds'
l smdunlikly andidats for
)pter. Spurnd y thir on-
on th study of ths faturs. In
r dinosaus.Rmination of
thy had nt n prviously r-
linosaursfrom hina. Prsumaly
rrgulation.on spimn, a ur-
Diosurs 521
portd glidr, appars to hav had wings. Finally, thr was th disovry of
irlrptor, th smallst known dinosaur-rcheopter-sized and a
asal dromaosaur to oot. larly miniaturization had ourrd for th
volution of irds. Additional findings solidifying th argumnt for dino-
saurian anstry for irds inlud th disovry that gg mirostrutur of
nlnaviandinosaurs is mathd in birds, that thropods roodd thir ggs
I i k i r d s , a n d t h a t x t n s i v a v i n s p i r r o ya i r s a s y s t m s x i s t i n d i -
nosaur skltons. !(/ith ths latst findings, th dat ovr avian anstry
has all but ndd. Dinosaurs did not go tint at th K- oundary (th g-
ologial bordr sparating sdimnts from th sozoi ra, whn th di-
nosaurs ruld, from mor modrn nozoi ra sdimnts)'dstroyd by an
astroid or othr atalysmi hang. Thy in fat rsid fluttring about
outsid today.
Dinosars hav playd an important rol in th dvlopmnt of modrn
volutionary thory y providing on of th most lbratd ampls in sup-
port of volution-rhopterx and th volutionary stags lading to its
gnsis.But do ths animals hav anything to offr for pomoting volution-
ary undrstanding in th futur? I liv th answr is ys. Alng with x-
ampls of human and quin (hors) volution, studis of dinosaurs will
srv as a valual vhil y whih th puli an om vrsd in the si-
ntifi mthod and at th sam tim grasp th workings of volution. Th r-
beopter story is not th only xample that is workd out in intriat
dtail, although argualy it has sen mor xamination than any fossil vrt-
rat. In fat it is just on of many. Sintists now know how many tims di-
nosaurs rahd lphantin proportions, how thy did it through sltion
for variant dvlopmntal trajtoris, and how thy am dwarfd in is-
.s
land sitations. know how thy divrsifid with ky innovations suh as
dntal attris (th volution of hwing using multipl tth) in onjun-
tion with floral hangs. And w hav a good undrstanding of how limati
hangs' ontinntal drift, and th dmis of nonavian dinosaurs-plausily
at th tim of an astroid impat-ld to volutionry hang that shapd th
world.
Th aformntiond studis of volutionary vnts hav rivd onsid.
ral attntion in th worldwid mdia and hn in th publi y.
Through dinosaurs th volutionary mssag is gtting through. As suh, for
roadiy promoting voiutionary undrstanding' dinosaur rsearh is ar-
gualy on of th most important filds today. owvr, dinosaur rsarh's
influn in promoting sintifi undrstanding and its potntial for hlping
volution gain roadr aptan ould onsidraly gratr. }yIor of-
tn thn not our hildrn's first introdution to sin is through dinosaurs,
and thr is no ttr opportunity to tah th sintifi mthod than to
young' urious, opn minds. Thos who ask for data in support of what thy
ar told will ttr quippd to judg for thmslvs whthr what thy
ar told is a rality or not. It is for this rason that dinosaur rsarh is an
important sintifi pursuit, on that an failitat sintifi litray and
522 Dobzhnsh

provid th rquisit akground for undrstanding and iudging th lgiti- that fild and st its arl1
may of astrat onpts suh as volution. Tahing sien at an arly Getis d the rigm
ag should of primary importan for sintists intrstd in promoing a Was that th study of g
mor sintifi litrat soity. Dinosaurs an provid th modl ampls to duil to gntispr s
mak this a rality. th ntirty of volution.
gard h stipulatd y d
B|LIoGRAPHY gn frqunis'' (Doz
op, . D. \867 . An untof xtint rptilsthat aproahird,s.Procedigsof Th main prolm of
th cdem of Nturl Siecesof Phildelphi19: 234_25. ..parado
Gtics, is th
Darwin, . 1859. on the origin of Speles.London:John Murray. to hav variation prsn
nhantdLarning.Zoom Dinosaurs.[.!bsit for hildrn.]http://www
othr hand, th partiul
.nhantdlarning.om/sui ts/dinosaurs/.
ar vry likly maladapt
Ggnaur,. 1864. Untersuhge 7ur uergleichnd Atomider ..volutt
,1., mati fashion,
Wirblthir.o|. rpus und rsus.Lipzig:!7ilhlm nglmann.
Huxly,T. H. 1868.On th nimalswhih ar mostnarlyintrmdiat btwn lssly dar pri of ont
irdsnd rptils.Geologilgzi5:357_365. unfavorabl mutations''
ostrom, J. H' 1969. A nw thropoddinosaurfrom th Lowr rtaousof D o z h n s k y . so w n i n s
]Vlontana.Postill 128: T-17.
variation in natural (nor
I975 . T origin of irds.ul Reuit,c'
of rth d PltrSins3:
55-77 . in Russia. At that tim
Own, R. 1853. on theArheoptero| Von Myr, with a dsriptionof th btls,ut it was his stt
fossil rmainsof a long-taildspisfrom th lithogrphistonof Solnhofn. f|, Drosrlphil, that w
PhilosophiclTrsctionsof th Rol Soietof Lodo |53: 47. drtakn in th 1930s,
Univrsityo{ California N4usumof Palontology.www.ump.rkly.du. working with th Drosa
.Wagnr,
J. A. 1861. Ur in nus,anglihmit Voglfdrnvrshns Rptil aus
Dozhansky,s arly Wor
dm SolnhofnrlithgraphishnShifr.Brishkdmider
'Wissshft (unih) 2z 146-154. -G.'. 1981).
Drosophil at first s
thr smd to no int
gan's itmnt at finalL
Dobzhansky'Thodosius(|900-975)
i n h r i t n o u l d s t t
Thodosius Dozhansky is prhaps most famous for th slogan, ..Nothing in gratrnumrs in his la]
iology maks sns xpt in th light of volution,' (Dozhansk 1973|, ations wr artifats. D/
whih is ndlssly invokd y volutionists in thir attls with rationists. traspifi variation' and
But that is only half th attl that Dozhansky wgd on half of th im- volution. In trying t
portan of volutionary thinking. H had two targts in mind: rationists wild Drosop hil' Dozhl
and thos would- volutionists who naivly suppos that molular iol- who proposd that most
ogy is th ky to lif whn in fat nothing, not vn th prosss of molu- al diffrns only wh
lar biology' an undrstood pt in th light of volution. Ironilly, our mor oftn in la<
Dozhansky himslf was osssdin a rligious or at last a moralisti way as thr is signifianti
with th ..maning'' of vrything, volution inludd (Batty 1'987a, 1'9871 way, natllral population
Rus 2001; van dr eer 2007)' rnst ayr (196) may hav blivd that Dozhansky, originl
volutionary iology providd th ultimat planations of lif, whil mol- W a y t o o s r v t h r w i s
ular iologial explanations ar at st proximat' ut for Dozhansky, th rosop on variations l
ultimat understanding of lif was dpr still than volution (sth disus- that involv diffrns
sion latr in this ntry). gn diffrns as Wll.
To thos familiar with his work, Dozhansky is known not just as a flag f o v o l u t i o n a r y h n g
arr for volutionary thinking, ut in partiular for his ontriutions to Dozhansky at first part
population gntis.H did as muh o mor than anyon ls to stlish to p|in rhis varitit

andingand iudging tlr lgiti-
. T a h i n gs i n a t a n a l y
l t i s t si n t r s t di n p o m o t i n g a
providth modl ampls to
rt approah irds. Proedigs of
hi 79 234_235.
rn: John urray.
lr hildrn.] http://www
h,1tomi dr
'zig:Wilhlm r.rglrnann.
lst nearly intrmdi:rttwn
/-5b).
,m th Lowr rta<lusof
of ,trth d PItr Sis 3:
yr,with a dsription of th
lithographi ston of Solnhfn.
et of Ldo 153: 47.
. w.tvw.ump.rkly.du.
Voglfdrnvrshns Rptil aus
kdmi nn'
|rist-,|l
-..'.
/Jl
rous for th slogn,
v o l u t i o n ' '( D o z a n s k I 9 7 ) ,
t n t h i r a t t l sw i t h
rsky wagd on half of th im-
] W t r g t si n m i n d : r a t i o n i s t s
ly supposthat molular iol-
ot vnth prosssof molu-
h light of volution. Ironially,
,iousor at last a moralisti way
Linludd(Batty 1,987a,1987;
,r (1963\ may hav livd that
:planationsof lif, whil mol_
intt,ut for Dozhansky, th
ill than volution (sth disus-
Lnskyis known not just as a flag
t r t i u l a rf o h i s o n t r i
)r than anyon ls to stalish
Dolzhsk 52
that fild and st its arly agnda, spiallythrough his influntil ttook
Gnetics nd tb rigin of Spis.on of th urrnt thms of Gnetics
ws that th study of gn frquny hangesis an autonomous fild, not r-
duibl to gntispr s (i..,th gntisof individuals). And its domain is
th ntirty of volution. What Dbzhnsky ould not prov in th lattr r.
gard h stipulatd y dfinition: ..volution is [no rrrrthn] a hang in
gn frqunis''(Dozhansky 197, l t; s also susquntditions).
Th main prolm of population gntis' as Dozhansky l:rl.loratdin
Genetis, is th..parado of viailiry.'' On th on hand, ah spisnds
to hv vriation prsntin ordr to adapt to nvironmntal hangs.On th
otlrr hand' th partiular vriations that might pov adaptiv in th futur
r vry likly ml1aptivat prsnt.As h onludd in his typially dra-
ati fashion, ..vrlutionaryplastiity an puhasd onl at th ruth-
lssly dar pri of ontinuously srifiing som individuals to dath from
unfavoralmutations'' (Dozhansk 797, 126-127).
Dozhansky's own insights into th paradox wr asd on his studis of
variation in naturl (nonlortory) populations, whih gan in th l920s
in Russia. At tht tinr h ws studying intraspifi variation in ladird
btls,ut it was his studis of virriation in naturl populations of th fruit
f|, Drosophil, tat would prov most influntial. Ths studis Wr un.
drtakn in th 1930s, aftr h had movd to th Unitd Stats nd was
working with th Drlslpbil gntiist Thomas unt organ (onrning
Dozhnsky's rly w<lrk,s Btty |987a, 1'994 Lewontin 1981; Pror,in
o. r98r).
Drosophil at first smd vry diffrnt from ladyugs. Phnotypially,
thr smd to no intraspifi varition at all. on nd only rall Mor-
gan's itmntat finally disovringan osrvalmutnt whos pttrn of
inhritan ould studid. Susquntly,variations wr found it-tvr
..Nothing in
gratrnumrs in his l:rortory.But it was livdy rrranytha thsvari-
ations wr artifts. Drosophil in th wild smdto hav littl if ny in.
a t i o n i s t s . traspifi variation, and hn prplingly littl rrrrialfor futur adaptiv
volution. In trying to mak sns of th diffrn twn laoratory and
wi|d Drosophil' Dozhansky followd th suggstilnof Srgi htvrikov,
who proposd that rost mutatios wr rssivand rvor-rldsult in osrv-
al differnsorly whn dorrld up in th homozygous stat.Tlris would
our mor oftn in laoratory stoks than in natllr' bus in th formr
s thr is signifiantly mor inrding than in wild populations. In this
way, natural populatins sak up vaiations lik a spong.
Dozhnsky, oiginally in ollaoratiln with Alfrd Sturtvan' found a
Wy to osrv othrwis hiddn variatiorri Dros<lpilay fousing lris rni_
Osopon varitilnsin hromosom strutu1.g-.(inyg15i6n''yx1i;11i1;n5-
tlrat involv diffrns in gn arrangmnt and rroftn assoiatd with
gn diffrnsas wll. In oth rspts'ths verietions onstitut rntrial
for volutionay hang. But how ws th osrvd vriation minturind?
..shifting
t i o n st o Dbzhirnsky at first partly adoptd Srvall!7right's balan'' thory
to plain this virriation. H rirsond that tlr variations Wr adaptivly
524 Dobzbnsk

insignifiant and that thir frqunis wr drifting randomly. This might As muh as h laim
lad to a rdution in variation within ah population of a spis' ut dply tzz\ed and un
would lad to an inrasin variation twnpopulations and hn within prodd. H had four
th spis. t h p r s r v a t i o no f v i
Dozhansky was quit surprisd, thn, to find vidn that th invrsion that ronild th go
diffrns wr adaptivly signifiant aftr all, lading him to plor a rang gratr fitnss of htro
of variation-maintaining forms of natural sletion. H finally sttld on th tinu to orn in ah
ida that htrozygots ar adaptivly suprior to homozygots us hav- undrstood this situatlo
ing two gns for a ttalt, instad of two opis of th sam gn, allows mor fr will, whih givs us
adaptaility at th individual lvl. Sltionfor adaptaility at th individual Prhaps. But h also x1
lvl in turn prsrvsvolutionary adaptaility at th spis lvl. ntur'' tht so many ir
..alan''
Dozhansky's aount am to known as th thory of vo- of th group (Dozhan
lution. His main dtrator was rmann Mullr' whos viw of volution 1_1'22;van dr r
..lassial'' (old-fashiond)
was dud y Dozhansky th thory (on th Dozhansky drw p
lassial/alanontrovrsy, s Batty 1987 [and othr ssayson that is- sours in his invstigat
su] and Lwontin 1974). Aording to Mullr, thr was no parado of via- was a major rason for t
ility. Th vast majority of mutations ar dtrimntal for th individuals in txt. Dozhansky's und
whih thy our, and lso for th spis in whih thy aumulat. Thr intraspifi variation a
was, in ullr's opinion, only on spis-humans-in whih mutations a- dr Mr 2007), lthug
umulat to any grat tnt. Through th amnitis of ivilization, w hu- and many of th sam
mans hav managd to sap natural sltion,to our ultimat dtrimnt. also n raisd in th l
Only through a onsious ugnis program an W now sussfully rvrs in onvrsations with D.
th rising tid of dltriousmutations. Dozhansky, for his part, ritiizd
..th B|LIoGRAPHY
ugnial Jrmiahs'' who ovrlook th importan of volutionary
plastiity and who sm unawar that natural populations of many spis Adams, M. B., d. 1994..
possssonsidralrsrvsof varition (Dozhansk 1'97, |26). PrintonUnivrsityI
Btty,l. 1987a. Dozha
Dobzhansky and ullr wr also at th ntr of ontrovrsis from th
biology.In L. Kriigr
1950s through th ear| |970s onrning th impat of radiation-indud amridg,A: Har
mutation from atomi Wapons tsts. (Both mn rivd onsidral fund- |987..!ighint
ing from th Atomi nrgy ommission during this priod.) Adding yt Jourl of th istor
mor mutations to th human spis was straightforwardly ngativ from 1994.Dozhansk
signifian of gn
ullr's point of viw, but lss so from Dozhansky's viwpoint. Intrst.
ThodosiusDobzh
ingly' Dozhansky was strongly opposd to th tsting and us of atomi
Dozhansky,T. 1'937.C
Wapons' whil th on pro-Sovit ullr livdthat, up to a point, suh UnivrsityPrss.
Wapons had an important rol tl play in th <rntainmntof ommunism 1.962,Mnhidl
( B a t t y1 9 8 7 ) . avn' T: Yl Unrv
Dozhansky also usd his studis and prsptivson variation to suppot l 9 7 . N o t h i n i r r
mri Biolog T
liral dmorati idals. As h and Lsli Dunn argud in thir popular
Dunn' L. ., and T. Doz
Hredit, R, d Societ (Dunn and Dozhnsk 1'946,45), ..thsolut Pnguin.
uniqunss of vry human individual . . . translatd into mtaphysial and Lwontin'R' , 1974,h
politial trms is fundamntl for this nd dmoray.'' In his widly rad Harvard UnivrsityPl
kid uoluig, Dozhansky argud that, far from gnti divrsity un. I9 8 l . l n r r d u t
drmining qulity' it atually srvs as an llnt rational: ..quality of Lwontint al.,ds.'
N w Y o r k : o l u m h i
opportunity tnds to mak th oupational diffrntiation omport with th
Lwontin,R. . t al.,ds
gnral polymorphism of th population, nd would maninglss if all I-XLIII. Nw York: (
popl wr gntiallyidntia|" (|962, 244). N4ayr,. 1963.ausnd

drifting randomly. T'his might
r population of a spis, ut
l populationsand hn within
i n d v i d n r h a r t h i n v s i o n
, lading him to xpl<lra rang
:tion' H finally sttld on th
r to homozygots aus hav-
; of th sam gn' allows mor
r adaptailityat th individual
y at th spislvl.
..lan''thory
rs th of vo-
ullr' whos viw of volution
(old-fashiond)thory (on th
7 [and othr ssays on that is.
:r,thrWas no paradox of via-
jii'mf;..f, . r.tl * ;nci;.'ict r]s-;'
l whih thy aumulat. Thr
u m n s - i n w h i h m u t t i o n sa -
rmnitisof iviliztion, w hu-
: i o n ,t o o u r u l t i m a t d t i m n t .
an W now sussfullyrvrs in onvrsationswith Dozhansky; B21tt|994,207 and not 3).
lzhansky,for his part, ritiizd
h importan of volutionary
. l p o p u | a t i o n so f m a n y s p i s
o b z h a n s k yt 9 7 ' 1 2 6 ) .
ntrof ontrovrsis from th
himpat of radiation-indud
mn ril,donsidalfund-
Juring this riod.) Adding yt
;traightforwardly ngtiv from
lzhansky'sviwpoint. Intrst-
) th tsting and us of atomi
rclivdthat, up to a point, suh
: h o n r i n m n to f o m m u n i s m
;ptivson variation to support
Dunn argr.rdin thir ppular
: h a n s k y\ 9 4 6 , 4 5 | , . . t h s o l u t
anslatdinto mtphysial and
dmoray.'' In his widly rad
, far from gnti divrsit un-
llntrational:,.quality of
diffrntiationomport with th
rnd would mninglss if all
41
Dobzhsk 525
As muh as h laimd to undrstand volution, Dozhansky rmaind
dply puzz|ed and unomfortal aout th mans y whih evolution
prodd.H had found a Way to ronil th sltion of th fittst with
th prsrvation of vaiation for futur adaptiv volution, ut not in way
that ronild th good of th spiswith th wlfar of its mms.Th
gratr fitnss f htrozygotsmans that lssr-fithomozygots will on-
tinu to orn in ah gnration.It hs n suggstdthat Dozhansky
undrstood this situation rligiously, as a nssaryvil-along th lins of
fr will, whih givs us moral frdom' ut at th ost of many sad hois.
Prhaps. But h also pliitly rgardd it as an ..unplasingimprftion of
natur'' that so many individuals should suffr suh ..misry'' for th good
of th group (Dozhansk197,1271Dozansk 1962, 35; Rus 2001,
|00_\221van dr 2007).
Dozhansky drw produtivly on his Russian hritag and Russian
sours in his irrvstigationsand planations of intraspifi variation. This
was a major rason for th originality of Gtics in an Anglo-Amrian on-
txt. Dobzhnsky's undrstanding of th rligious/moralisti signifian of
intraspifi variation also rfltd his Russian hritag (s spilly van
dr Mr 2007), lthough this influn has n mor diffiult to ronstrut,
and many of th sanr onrns-suh as th vils of natural sltiln-had
also rrraisd in th Anglo_Amrian ontt (..g., y th athist organ,
|BLIoGRAPHY
Adams,M. B., d. 1994. Th ullutioof ThodosisDobzhsk.Printn,NJ:
PrintonUnivsityPrss.
Batty,J. 1987a.Dozhanskyand drift:Fats,valusand hanin volution;rry
iology.In L. Kriigr t al., ds.,The ProbilistiRuolutk-l,olr.2.
ambridg,A: avrrdUnivrsityPrss'
1987.!7ighingth isks:Stlmt in th lssil/alan
ontrovsy'
lourl of th istor of Biolog 2:289_|9.
\994. Dozhanskyand th iologvof dmoray:Th moral and politial
signifianof gntivariation.In . B. Adms, ed.,Tbe uolutio of
TheodosiusDobzhsk'Printon,NJ: PrintonUnivrsityPrss.
Dozhansky,. 1937, Geticsd the origi of Sples.Nw York: olumia
UnivrsityPrss.
|962. nkitl uoluig:Th uolution lf the um Speles.Nw
Havn' T: Yl UrrivrsityPrss.
1973.Nothingin iologymakssnsptill th lightof vllution.
triB iollgTrhr5 : 1 5-129.
Dunn,L. ., arrdT. Dozhnsk'1946.Hrdit,R,d Soiet1'. Nw Yok:
Pnguin.
Lwontin,R. . 1974. h Geti Bsisof uolutirlrhge.Camridg,MA:
Harvard UnivrsityPrss.
1981.Introdution: Th sintifiwork of ThdosiusDozhansky.In R. .
Lwontint al., ds.,Dobzhsk'sGetisof Nturl PopultiosI-XLIII.
Nw York: ColuriaUnivrsityPrss.
Lwontin,R. C. t al.' ds. "|981. Dobzhnsk'stisof Nturl Popltions
LXLLI]. Nw York: olrrmiaUnivsityPss.
ayr, . l963. irusand ittin iologv.Si134:1501_1506.
526 Drosophila
.v.
Provin, B. 1981.origins of th Gntisof Natural Populationssris.ln R. .
Lwontin t al., ds.,Dobzhsk'sGtisof Nturl PopltionsI_XLI.
Nw York: olumbia UnivrsityPrss.
Rus, . 2,|.ster of stris:Is uolutio Slilostrutio?
ambridg,A: Harvard UnivrsityPrss.
van dr Mr,J. 2007.ThodosiusDozhar-rsky: ..Nothingin volutionmakssns
xptin th light of rligion.''In N. Rupk, ed,,mint Lius i Tt,utith-
tur Sic d Religion. Frankfurt: Ptr Lang. -1.8.

DrosophiI
Drosophil is a gnus of flis in th family Drosophilida that ar widly
usd for volutionary rsarh. Th origin of this rsarh lis in Thomas
Hunt organ's Noel Priz-winning work on inhritanc in Drosopbil
mlogstr (s figur) starting around 1908. organ's intlltual
dsndants hav pandd th sop of work on D. melnogstr from
gntis into virtually vry aspt of iology, turning it into on of th
most usful modl organisms. volutionary rsarh has n a roust
part of this tradition from th bginning. urntly' th omination of
th as and spd with whih many spis an rard in th laora-
tory and th tnsiv akground knowldg of gntisand dvlopmnt
mak thm usful in addrssing a wid vrity of qustions in volutionary
gntis.
Drosophil adults ar usually only a fw millimtrs long' with prominnt
rd to rown ys, and ompl trnal mophology that has rmaind
fairly onsrvativ ovr volutionary tim. Drosophil sow holomtaolous
dvlopmnt, undrgoing a omplt transformation from th fding larva
to the adult form during a pupal stage. Thy ar ommonly alld fruit flis,
although most spis fd on miroorganisms that inhait rotting plant ma.
trial. Th gnus Drosopbil urrently inluds twn 1,150 and 1,500
spis, dpnding on whih groups ar inludd in th gnus, and has a
worldwid distriution. Phylognti rsarh has larly shown that th
genus is paraphylti, maning that othr gnra ar mor losly rlatd to
som of t'e Drosophil than thy ar to othr groups. In th road sns,
thn, Drosophil sou|d inlud many other gnra' inluding th spis-
ri Sptomz (approximtly 260 spis), Hirtodrosopbil (approi- Drosophil mlogs
matly 150 spis), and codrosophil (approimatly 120 spis). Th way in whih it wl
N{olular vidn sttggststhat th gnus is at last 50 million yars old, \994.
a onlusion supportd y drosophilid spimnsin ambr that ar around
this ag.
Around 1910, organ disovrd a singl D. mellgsler with whit
ys' th inheritan of whih turnd out to s linkd. This simpl obsr- undrgraduats Alfr
vation inspird Morgan and his laoratory studntsto isolate othf mutants, N4ullr, who all playr
with whih thy wr al to onfirm th hypothsis that gns rsid on Morgan, Bridgs, an
hromosoms. In thos yars, Morgan's tiny (16 ft 24 feet)..fly room'' whn thy all movd
at olumia Univrsity had ight dsks; thr wr oupid y olumbia whr thy rmaind
 Drosophila 527

aturalPopulations sris.In R. .
of NturI Popultios I-Llll.

SoiI ostrutio?

.Nothing in
vlution mks sns
d.,liet Lius i Ttuentietb-
:r Lang. -1.8.

Drosophilida that ar widly

of this rsrh lis in Thomas
: on inhritan in Drosophil
l 1908. Morgan's intlltual
rork on D, mlogstr from
ogy, turning it into on of th
y rsarh has n a roust
urrntIy, th omination of
s an rard in th labora-
g of gnetisand dvlopmnt
ty of qustions in volutionary

r i l l i m t r lso n g , w i t h p r o m i n n t
morphology that has rmaind
r osophil show holomtaolus
rmation from th fding larva
.ar ommonly alld fruit flis,
ns that inhabit rotting plant ma-
u d s t w n1 , 1 5 0 a n d 1 , 5 0 0
ludd in th gnus, and has
h has larly shown that th
|nraar mor losly rlatd to
'hr groups. In th broad sns,
)r gnra' inluding the spis-
ies\, Hirtodrosopbil (approxi- Drosophil mlogster'th fruit fly, th workhors of volutionarygenetis.
r (approximatel 720 spis). Th way in whih it was tailord for suh rsarhis disussdin Kohler
is at last 50 million yars old, 1994.
imns in amr that ar around

31eD. melogster with whit

s linkd. This simpl osr-
tudntsto isolat othr mutants'
hypothsisthat gns rsid on
' (\6 feet 24 feet) ..fly room''
r wr oupid y olumi
undrgraduats Alfrd Hnry sturtvant' alvin Bridgs' and Hrmann
ullr, who all playd pivotal rols in gnetis and volutionary iology.
Morgan, Bridgs, and sturtvant workd togthr at olumia until 1928
whn thy all movd to th alifornia Institt of Thnology (al Th)'
whr thy rmaind for th rst of thir arrs.
528 Drosophila

sturtvant produd th first gnti map asd on th raliztion that th lvls of natural variz
frquny of rossovr vnts ould b usd to dtmin th ordr and loa- whthr gnti varr
tion of gns. H also workd on a wid varity of topis with volutionary maintaind by natur:
impliations, inluding hromosomal invrsions, th gntis of spis dif- ontrasting Dzhan
frns, th gntis of dvlopmnt, and th phylogny of th gnus. with ullr's mpha
Bridgs was among th first to xploit th disovry that th salivry glands was a major fator in
of Drosophil ontaind polytn hromosoms onsisting of thousands of of Drosopbila to stu
DNA strands rsulting from rpatd rounds of rpliation without ll divi- quning of Dros<lpb
sion. Th rsult is hromosoms visil at low magnifiation with hartr- of volutionary patt
isti anding pattrns that allow homologous rgions to idntifid aross Th as with whi<
individuals and spis. Bridgs's dtaild dsriptions of th anding pat- in th study of volut
trns allowd linkag and physial maps to alignd. ltion xprimnts <
ullr is bst known for his Nol Priz_winning work on mutation. B- Knnth Mathr and
ginning in th mid-1920s, ullr was th first to ploit th fat that hro- Susquntly' sus
mosomal invrsions-hromosoms in whih a pi appars in rvrs of haratristis, no
th normal ordr-supprss romination whn in a htrozygos ondi. gg siz, dvlopmn
tion with an uninvrtd hromosom. Mullr found that h ould us an in. totais' gotais' m.
vrtd hromosom markd with a dominant phnotypi mark, trmd a (rats of rominat
.!7ith
alanr, to aptur and study th proprtis of hromosoms with th nor- th xption
rnal gn ordr. Initially h usd this thniqu to dtt hromosoms arry- vry trait studid. S
ing lthal mutations' whih an nvr produ a homozygous adult. Mullr rats an nvironm
masured th lthal frquny aftr induing mutation with -rays. Mullr's ara of ativ rsar
onpt of gnti load-mutational aggag that ullr saw as harmful- van in th study of
am from ths arly primnts. Balanr hromosoms wr sus- ing and mat hoi
quntly dvlopd for ah hromosom in D. mlogstr, and alanrs Th study of sp
rmain on of th ky advantags of doing gntis in th spis. Mullr, and Dozhl
Shortly aftr Morgan movd to al Th, h hird Russian sintist Thodo. sarh. Th gnus ir
sius Dozhansky as part of his tam. Dozhansky quikly ros to prominn numr of rossal
as a synthsizer writing Geetics d the rigi of Species (197), on f barrirs an idnt
th founding ooks of th volutionary synthsis,in whih h assrtdth pri dnts, and his stud
may of a population gnti viw of volution, basd initially on rmarkably rough stimats of
littl mpirial vidn.At aout th sam tim, inflund y Sturtvant and lopatri populations
his Russian mntor Srgi htvikov, Dozhansky took up th study of wild hav now n idnt
populations of Drosophil pseudoobscur. Populations of this ommon .s7hil
th study o
spis of th mountains of wstrn North Amria ar polymorphi for hr- melogster am
mosomal invrsions. Dozhansky usd this variation as th asis for a long s- asd on th dtail
ris of xprimnts doumnting th ffts of th thr major evolutionary Sturtvant's studnts
fors: sltion,gnti drift' and gn flw. In this, h was grtly aidd y an ffts arly in dv
.lfright
xtnsiv ollaoration with Swall during th lat 1940s. shus. Ths ffort
Modrn volutionary work on Drosophil urrntly has svrl major m- ways that ar ons
phass that grw out of this arlir work. Th study of variation t th mo- .Wishaus
and shar
lular lvl was pionrd y Rihard Lwontin, on of Dozhansky's akground inform
studnts, who, starting tn 1965, xploitd th disovry of protin l- lution of dvlopmt
trophorsis by Jak Huy and othrs to dmonstrttht a larg portion of idntify th spifi 1
gnti loi in Drosopbil spis wr polymorphi. This disovry of high ral and xprimnta

|asdon th ralization that th
o dtrmin th ordr and loa-
i yo f t o p i s w i t h v o I u t i o n a r y
ns' th gntis of spis dif-
th phylogny of th gnus.
iovry that th salivary glands
ms onsistingof thousands of
of rpliation without ll divi-
v magnifiation with haratr-
; rgions to idntifid aross
sriptionsof th banding pat-
alignd.
winning work on mutation. B-
st to ploit th fat that hro-
h a pi appars in revrs of
whn in a htrozygous ondi-
: f o u n d t h h o u l d u s n i n -
rt phnotypi markr, trmd a
i of hromosomswith th nor-
l to dtthromosoms arry-
a homozygous adult. N4ullr
mutationwith -rays. Mullr's
-'that Mullr saw as harmful-
er hromosoms w sus-
D. melogstr, and alanrs
)ntisin th spis.
: hird Russian sintist Thodo-
nsky quikly ros to pominn
rigi of Spcies (1937), on of
:sis,in whih h assrtdth pri-
'n, asdinitially on rmarkably
n, influndy Sturtvant and
ransky took up th study of wild
Populations of this ommon
rri ar polymorphi for hro-
riation as th asis for a long s-
of th thr major volutionary
n this,h was gratly aidd by an
rringth lat 1940s.
u r n t l yh s s v a lm a j o m -
r study of variation t th mo-
3wontin' on of Dozhansky's
th disovry of protin l-
lonstratthat a larg portion of
nrphi. This disovry of high
Drosophila 529
lvlsf natural variation in most spis ld to an arimonious dbat ovr
whthr gnti variation is gnrally sltivly nutral, dltrious, or
mintaind y natural sltion. Lwontin hlpd to cata|ze this dat by
ontrasting Dozhansky's viw that muh gnti vaiation was adaptiv
with Mullr's mphsis on th osts of mutation and variation. This dat
was a major ftor in th dvlopmnt of population gnti thory. Th us
of Drosopbil to study variation arris ovr to th DNA lvl, whr th s-
quning of Drosophila gnoms allows unprdntd prision in th study
of volutionarypattrns.
Th as with whih Drosophil an rard has ld to its inrasing us
in th study of volution in th laoratory. Som of th first sophistiatd s-
ltion xprimnts on ontinuous traits wr arrid out in Drosophil
KnnthMathr and Fors \/.Rortson in th lat 1940s and arly 1950s.
Susquntly,sussful artifiial sltion primnts took pla on many
haratristis'nompassing mophology and physiology (.g.' fundity,
gg siz' dvlopmntalrat, lif span, rsistanto thanol)' havior (pho-
totais, gotaxis, mat prfrn), and aspts of th gnti systm itslf
(ratsof romination, hang in varian rathr than man valu of traits).
lfith th ption of sx ratio and asymmtry, hrital variation xists for
vry trait studid. Studis of primntal volution, whr th invstigator
ratsan nvironmnt in whih natural sltion an take pla, is also an
area of ativ rsarh using Drosophil. This approah has prtiular rl-
van in th study of lif historis and th ultimat origin of prosss lik ag-
ing and mat hoi.
Th study of spis diffrns, pionrd tn Drlsophila y Sturtvant,
ullr, and Dozhansky, is anothr vry ativ ara of volutionary r-
sarh. Th gnus is wll suitd to th study of spiation aus of th
numr of rossal spis and th rlativ as with whih rprodutiv
arrirs an identifid and quantifid. Jrry oyn, on of Lwontin stu.
dnts, and his studnt H. Alln orr usd this aumulatd data to mak
rough stimats of th tim rquird for spition in sympatri and al-
lopatri populations. Th gns rsponsil for aspts of spis barrirs
hv now n idntifid in a numr of Drosophila spis pairs.
\/hil th study of Drosophil dvlopmnt dats ak to th 1920s, D.
melogstr m a prmir modl systm for th study of dvlopmnt
asd on th dtaild study of th ithorax ompl y d Lwis, on of
Sturtvant's studnts' and th systmati mutation srn for mutations with
.$i-
ffts arly in dvlopmnt y hristian Nusslin-Volhard and ri
shaus. Ths fforts ld to many of th fundamntal dvlopmntal path-
Ways that ar onsrvd widly among animls. Lwis, Nusslin-Volhard,
and \Wishaus shard a Nol Prize for this work tn 1'995. This tnsiv
akground information on dvlopmnt has cata|zed th study of th vo-
lution of dvlopmnt in Drosopbil. Rsarhrs hav now n al to
idntify th spifi gnSrsponsil for many volutionary hangs in natu-
rl and primntaIDrosophil populations.
s0 Drosophila

|LloGRAPY
Dozhansky'T. 1937. Geneticsd the rigin of Speles.Nw Yrk: Columia
UnivrsityPrss.
Kohlr' k. 7994. Lords of th F/y. hiago:Univrsityof hiago Prss.
Lwontin,R. . 1'974.The GeticBsis of uolutionrhge.Nw York:
E
olumi UnivrsityPrss.
Powll' I. R. |997. Progrssd Prospetsi uolutiorBiolog: e Drosophila
odl. Nw York: oxford UnivrsityPrss.
Ruin, G. ., and . B. Lwis. 2000. A rif historyo| Drosophil'sontriutions
to gnomrsarh. Si287:2216-2218.
Sturtvant,A. H. 1965. istor of Gtics.Nw York: arpr and Row.
-D.H. d B.H. E,cologicI Geetlcs
ologicl G entis, t|tr<
s t u d y i n g n a t u ls l t i o
..dmund B
btial ntry
studnts, and olltdth
tis, is dsndd from tI
don that natural sltl
...W.F. R..Wldon'
ntry
gntis ovr work lik \
gnti asis suh as th
mot|7P rt i d omiul .
logial gntiists to stu
sponse to sltion as l
hangs in trait valus r
ths disovriswr s
trm ologil glis
dmonstral gnti fo
trait variation in natur l
hr typs of invsr
thr ar dmonstrations
in th moth Biston betul
vstigations of gograph
snails, that doumnt ho
anothr hang th sl
valus to favord in d
adaptiv xplanatins fo
dmonstrations of adapt
unovrd th signatu
s n a i l s h l lp a t r r n sT. h i r
an alan on noth
tion. Ths studis rpr
Innumral thortial
tion gw from th initi
sgrgatdspiting
cologil Getis l
s t u d i s l i m i n da n y d
Spis.Nw York: lumia

rsity of hiago Prss.

tionr hg. Nw York:
E
'
utio lrB i olog : TDe Drosophila

tv of Drosophlla's orrtiutions

:w \rk: arpr and Row.

-D.H' d B.H. E,cologicIGnetics (. B. Ford)
olr>giclGnetis, trough its svrl ditions, prsntdth approah to
studyingnatural sltionin th fild pionrd y . B. Ford (sth alpha.
btial ntry ..drnund Briso Ford'' in this volum), his ollagus,and his
studnts,and olltd thir susss.This approah, alld ologial gn-
.!.
tis, is dsndd fom th arly studis of R. C. Punntt and F. R. .07l-
don that natural sltion an studid mpirially (s th alptial
ntry *W. F. R. Vldrr'' in this volum). Th advn offrd y ologial
gntisovr work lik !7ldon's was a fous on disrttraits with a simpl
gntiasis suh as th variation in th numr of spots on th wing of th
mot Pi| dominl. Knowing th gnti asis of variation allowd o-
logial gntiiststo study not only sltion' ut also th volutionary r-
spons to sltion :rs trait valus hangd ovr tim or spa. Baus
hangsin tait valus rfltdhangs in th frqunisof altrnatgns'
thsdisovrisw small-saldmonstrationsof volution in ation. Th
term ecologil geetis is drivd from its fous on trait variation with a
drnonstral gnti foundation (gntis) and its orintation to studying
trait variation in natu (ologial).
hr typs of invstigations donrirrt Ford ologil Gtis. First,
thrar dmonstrations of volution in ation; th saga of industrial mlnism
in th moth Bistott betulri is th st-known xarnpl. Sond, thr ar in-
vstigations of gographi variation in trait valus, suh as th shll olors of
snails, that doumnt how hangs in ologial ftors from on loation to
aothr hang th sltion prssus on orgnisms nd aus diffnt trait
valus to favord in diffrnt loations. Ths invstigations dmonstratd
adaptiv planatins for gographi variation in hatrs, omplmnting
dmonstrations of adaptiv hang in tim. In som ass, ths studis also
unovrd th signtur of random vnts as in th so-alld ara ffts on
snail shll pattrns.Third, thr ar studis of how diffrnt sltion prssurs
an lrrol1 i1l1othnd prsv trait variation within a singl popula-
tion. Ths studis rprsntda partiularly frtil ara of ologial gntis.
Inumral thortial paprs on th population gntis of alaning sl-
tion grw from th initial disovris that altrnat gns would ontinu to
sgrgatdspit ing atd upon y strong natural sltion.
ollgil Gntis was normously influntial. Its ompilation of as
studisliminatd any dout tlrt natural slionwas an ongoing for and

) . t,
52 ign

ould studid in ral tim. Th studis of gographi variation ondutd individual frquny.
by Ford and his ollagusomplmntd th work of othrs' suh as Gt .
of th quasi-spis,
Tursson and Jns lusn with plants (s th alphatial ntry ..Jns and dirtd voluti
lausn'' in this volum), F. B. Sumnr with mi, and Srgi htvrikov slf_rpliation.
and Thodosius Dozhansky with flis (s th alphatial ntris ..Srgi In addition to om
Srgvih htvrikov'' and ..Thodosius Dozhansky'' in this volum)' opration lading to t
who wr making similar disovris, oftn with haratrs lss amnal to svral quasi-spisr
gnti analysis. Th waning of th ook's influn gan with th disovry adjant unit. This
of iohmial vriation nd ontinud through th mrgn of a sophisti. in th snario and al
atd approah to studying ontinuously varying traits that ar mor rpr- fat that RNA squ
sntativ of th stuff of volution than disrt polymorphisms in visil lngth (about 100 nu
haratrs. Nonthlss, th qustions aout sltion and volution pr. would aumulat,d
sntd in th book' and th onvition advand that ologial and gnti ordf to od for prot
studisar nssaryto undrstandadaptiv volution, rmain drok influ- mad it possibl to 1
ns on modrn volutionary iology. spis without lngtl
funtional protins. I
|LIoGRAPHY
pndd on thir nl
Ford, . B. |975. ologilGnetis,4thd. London:hpmanand Hall. -J,' form in th rih pri
partmnts ould thn
ign,s thory wal
ign, anfrd (. t927)
hmistry podud l
anfrd Bign is a Grman iophysiist and formr dirtor of th a dant supply of amin,
Plank Institut for Biophysial hmistry in Gttingn. In 1967, toget.er srv as nrgy-rih l
with Ronald Gorg.!ryford Norrish and Gorg Portr' h won th Nol sis of short gnti t
Priz in hmistry for th study of xtrmly fast hmial rations. Bgin- slf-organization. Th
ning in th 1960s, ign's rsarhintrststurnd to iohmial qustions, and mpirial assum
and during th 1970s and 1980s h formulatd an laorat thory on th wr svrly ritiiz
mrgn of lif on arth foundd on mathmatial alulations and pri- to idas on th volu
mnts in viral populations. tion of th onpt
ign's fundamntal thortial and philosphial postulat was tht th prinipl of natu
natural sltion Was th organizing prinipl rsponsil not only for vo. ut on any physial r
lution ut also for th mrgn of lif. This laim was basd on nw ad- standing th rol of
vanmnts in th fild of molular iology. Starting in th 1960s, it lawful pross.
am possil to dsri volution asd on rpliation, mrrtation, and
|LIoGRAPH
omptition not only of organisms ut also in mixturs of slf-rpliating
moluls and nzyms atalyzing rpliation. ign's rsarh along ths ign, M. 1992.Stpst
lins gratly advand th work f iohmist Sol Spiglman and othrs in ign, ., lW. Gardin
of gntiinforma
th 1970s that dmonstrtd th volution in th tst tu of viral RNA s-
Fry, I. 2000. h mr
quns. Basd on ths xprimnts' ign dvlopd his modl for th Nw Brunswik,N
origin of lif.
ign formulatd a ky nw onpt in origin-of-lif study and virology
..quasi-spis.''
alld th Following th omptition for uilding loks in
a population of RNA squnsngagdin slf-pliation?a uniqu pattrn
of th whol population mrgd asd on th distriution of th various
mlrtants. This rsulting population, th quasi-spis,rfltdth onsn-
sus squn rprsnting an avrag of all th mutants aording to thir

og'faphivariation ondutd
) work of othrs, sh as Gt
l th alphatial ntry ..Jns
mi, and Srgi htvrikov
..Srgi
h lphatialntris
)ozhansky'' in this volum),
'ith hrtrslss amnbl to
ngnwith th disovry
gh th mrgnof a sophisti.
ging trits that ar mor rr-
:rtpolymorphisms in visil
t sltionand volution pr-
d that ologial and gnti
volution,rmain drok influ.
ion:hprnand all.
l a fomr dirtor of th Ma
n Gttirrgn.|n 1967, togthr
;orgPortr, h won th Nol
fast hmial rations. Bgin-
-rrndto iohmialqustions'
Ltdan laoratthory on th
matilalulationsand pri-
losophial postulat was that
[ rsponsilnot only for vo-
is laim was asd on nw ad-
ogy. Starting in th 1960s, it
l on pliation' mutation, and
in miturs of slf-rpliating
'n. ign'srsrh along ths
st Sol Spiglmanand othrs in
n th tst tu of vial RNA s-
n dvlopd his modl for th
rigin-of-lifstudy and virology
nptitionfor uilding loks in
lf-rpliation,a uniqu pattrn
t h d i s t r i u t i o no f t h v a r i o u s
si.spis,rfltd th onsn-
th mutants aording to thir
ign .'3J
individual frqunv' ign disovrd that, asd on th spifi dynami
of th quasi-spis,th mharrisrnlf sltion guarntd r-rlnrandom
and dirtd volution toward a population apl of aurat nd fast
slf-rpliation.
In addition to omptition, ign also postulatd stags of molular o-
opationlading to th mrgnof a ..hypryl,''an nsmlin whih
svrlquasi.spisunits partiiptd, ah naling th rliation of th
adjnt unit. This oopration was dmd rr-rialto ovro1 a cath-22
in th snario and allow furthr volution. Th diffiulty stmmd from th
ft that RNA squnsin th quasi-spishad t limitd to a rtain
lngth (about 100 Irulotids).Aov this thrshold, too many mutaions
would aumulat,dstroying th squns'idntitis.At th sam tim, in
ordr to od for potins,muh longr hains wr rquird.Tlr hypyl
md it possil to pull togthr th information ontnts of ah quasi-
siswithout lngthningindividul hins' thus naling th synthsisof
funtional protins' Howvr' th oopration of svral quasi-spisd-
pndd on thir nlosur within a ompartmnt, postulatd y ign to
-l.T. fom in th rih primodil ..soLlp.''optiion mong th divrs om-
partmntsould thn prmit furthr volution.
ign's thory was asd on th optimisti ssumption that prioti
hmistry produd in th primordial soup a far-from-quilirium, aun-
dant supply of anrino aids, nulotids, and thi polymrs. Ths ould
srv s ngy-rih uilding loks and primitiv talysts for th syrrth-
sis of short gnti squns irnd for th ruil prosss of iologial
slf-organization. This optimism s wll as svral of ign's rhortial
and mpirial assumptions rlating to th quasi-spisand th hypryl
wr svrlyritiizd y othr rsrhrs.Nonthlss' his ontriution
to idas on th volution of gneti information and to th latr formula-
tion of th onpt of th RNA world is indisputirl. So is his lim that
th prinipl of natural sltion Oprats not only on iologial systms
ut on any physial systms fulfilling rtain onditions and that, notwith-
standing th rol of random vnts' th mrgn of lif was a naturally
lawful pross.
BIBLIOGRAPHY
ign,. 1992.StpstlrdsLif. ford: Ofrd UrrivrsityPrss.
ign,., W. Gardinr,P. Shustr, rrndR. Winklr-Oswatitsh. 1981.h lrigin
of gntiinformtion.Sitifimri244, no.4:78_|18.
Fry, I' 2000. Th mrgof Lif t rth: lListorild Scintifit',arui.
Nw Brunswik,NJ: RutgrsUnivrsityPrss. -t.t.
54 dosmbioti rigi of uhrots

all tims. mrans

ndosymioti origin of ukaryots
utils, dfin th inl
whih r mad of it
PREDNs AND PRDEssoRs
this implis that th in
on ontntious' th thory is now aptdthat all ukaryots (organisms prtin, and lipid n<
omposd of lls with nuli and tru hromosoms),inluding th familiar mnt yond th outr
plant and animal organisms, volvd from symioti mrgrs of atria. th uiquitous ativ
lar gntividnonntsth plastids (grnphotosynthtiorganl1s) waxy surfa, shath,
of alg to fr-living photosynthti atria alld yanoatria. And - maintains idntity of li
aus alga and plants hav a distant ommon anstor' this vidn on-
nts th yanoatria to modrn plants. So, too, th oygn-using ..powr
ALL LIFE FROM
paks'' in lls, th mitohondria, hav n trad via gntis,physiology,
and morphology to fr-living rspiring tria.For th last 3 illin yars All lif forms on arth
or mor' through mutations and gn trading' on has th sam kind of gn- proteins. Protins, whi
ti nginring that gos on in iothnology firms. This aumulation of ar synthsizd on intr
DNA mutations and passing of sts of gns from on kind of atrium to osoms look lik tin1
vry diffrnt kinds is, and has n sin anint tims' th way of th a. tron mirosop. m
trial world. Yt mrgrs, whol.ody fusions, and prmannt allians as faturs of lif that att
ways of survival in th hardst of tims has sltdatrial assoiationsto lar anstors.
form nw individuals. Batria, y virtu of litral takovrs, hav also Batrial gns,thos
volvd into somthing ls. By fusion of diffrnt kinds of odis, rathr of DNA; th hmial
than y mutating and growing largr, atria volvd to om largr lls. rial hmistry, ar iik
Th largr lls, inluding th anstorsto all animals and plants, indd all last 3,000 diffrntg
lls with nuli (all ukaryots),volvd y symbioti mrgr. tins. Batrial lls gr
known as fission. Th
..
apparan than th
ARIAL ANsRY
in ukaryots. Known
Batria ar uiquitlus organisms that ar prsnt in th fossil rord long isms always involvs
for any animals or plants. Batria, not plants, volvd photosynthsis tuuls (sfigur on 1
and hmial synthsis,th prosssthat provid food to th rst of th io- moving, as sn throu[
sphr.All atria shar a ommon llular struturknown as prokaryoti. prokaryoti lls divi
Prokaryoti lls, unlik ukaryoti lls, do not hav an nlosd nulus mirotuul-asdmo
housing thir DNA, ut rathr an opn ra alld a nuloid in whih th disontinuity twn
DNA rsids. Typially thr ar no ntriols, spindls' or mirotubuls tinary gap in th livi
(indd thr is no mitosis as in ukaryots)' and prokaryoti lls lak Although larly w
mmran-bundd nzym sas within whih organi moluls ar oi- volvd, somhow, fro
dizd. Batria' whih ar all prokaryoti, shar mny ut not all traits in tria and ukaryotsa
ommon with th ukaryots. Lik th thr main groups of mor familiar, and th mmrs of th
largr ukaryots-animals, plants, and fungi (all yasts, molds, and mush- forms of lif ar om
rooms ar fungi)-atria r mad of lls that ar oundd y dynari nuli. For that raso
protin-rih lipid mmrans.All lls from th tinist atria to th largst Grk). All atri h
gg lls us th nzyms of thir ompl nrgy and aron mtaolism to and th tru hrom(
produ thir own nssaryoutr mmrans.Th mtaoli maintnan protins.
of th dynami protin-rih, phosphorus.rih, lipid-rih mmrans x- Batria' as a group
hang omponnt parts. lls us foodstuffs as raw matriis. Mmrans mals' and fungi. Wh
napsulatidntity. Th asolut nssityof ontinuity through spa and rath sulfat' aro
tim rquirs th prsnof tiv, intat mrranssurrounding all lls at thir small siz, mor

d that all ukaryots (organisms
nosoms),inluding th familiar
symioti mrgs of atria.
grnphotosynthti organlls )
Laalld yanoatria.And -
lon anstor' this vidn on-
..powr
)' too' th oygn-using
. tradvia gntis,physiology'
r i a .F o r t h l s t i l I i o n y a r s
4' on has th sam kind of gn-
,gy firms. This aunrulation of
; from on kind of batriumto
-rinttims, th way of th a-
lns, and prmannt allians as
l s s o i a t i o n st o
s | t d a t r i
of litral takovrs, hav also
liffrnt kinds of bodis, rathr
ia volvdto om largr lls.
'll nimals and plants, indd all
symiotimrgr.
prsntin th fossil rord long
plants' volvd photosynthsis
'ovidfood to th rst of th io_
struturknown as prokaryoti.
o not hav an nlosd nulus
a alld nuloid in whih th
'iols, spindls, or mirotubuls
ts), and prkaryoti lls lak
h i h o r g a n i m o l u l s r o i _
shar many ut not all traits in
main groups of mor familiar,
Lgi(a11yasts,molds, and mush-
ls that ar oundd y dynami
th tinistatriato th largst
nrgynd aron mtaolism to
n s .T h m t l i m a i n t n n
rih, lipid-rih mmranS x-
:fs as raw matrials. N4mrans
of ontinuity through spa and
mranssurrounding all lls at
ndosmbioti rigi of ukrotes 55
all tims. Mmrans, and oftn additional ovrings suh as 1lwalls and
utils, dfin th insid and th outsid ordrs of all organisms, all of
whih ar mad of ithr prokaryoti or ukaryoti lls. In hmil terms,
this implis that th insid of any organism is in a stat of Watr, salt' sugar,
protin, and lipid onntration vry diffrnt from its outsid. Th nviron-
mnt yond th outr mmran is vr-prsntand oftn hanging, but it is
th ubiquitous ativ mmran (whthr nakd or supplmntd y skin,
waxy surfa, shath, or othr additional outr layr) that stalishsand
maintinsidntity of living ings.
ALL L|F FRo H ARLIs LLs
All lif forms on arth mak protins insid thir lls, inluding mmran
protins.Protins, whih ar long hains of smallr units alld amino aids,
ar synthsizd on intrallular tiny sphrial struturs alld riosoms. Ri.
bosoms look lik tiny dots vn at high magnifiation as sn with an l.
tron mirosop. mbran iohmistry and protin synthsis are univrsal
fatursof lif that attst to th fat that ll lif volvd from ommon llu-
lar anstors.
Batrial gns,thos of humans as wll as all othr animals, ar omposd
of DNA; th hmial omposition of gns, lik many othr dtails of at-
rial hmistry, ar lik thos of th largr organisms. Most ateria hav at
last 3,000 diffrnt gns that dtrmin thousands of diffrnt nzym pro.
tins. Batrial lls grow and rprodu y rprodutiv division, whih is
known as fission. Th prokaryoti ll division pross is far lss omplx in
apparanthan th ..hromosomdan'' alignmnt and sparation pross
in ukaryots. Known as mitosis' th ll division pross in nlatd organ.
isms always involvs movmnt insid lls on littl traks alld miro-
tubuls (sfigur on pag 536). Th insid of a ukaryoti ll is inssantly
moving, as sn through a mirosop whn th ll is aliv. vn th largst
prokaryoti lls divid y fission; thy lak th mirotuls and th
mirotuul-asdmovmntstypial of nulatdlls. Som laim that th
disontinuity twn atria and all othr forms of lif is th largst volu.
tionary gap in th living wor]d.
Although larly w (i..,largr forms of lif omposd of nulatd lls)
volvd, somhow, from trial prdssors, th diffrns twn a-
tria and ukaryots ar signifiant. Th major diffrns twn atria
and th mmrs of th animal' plant, and fungal kingdoms is that all largr
forms of lif ar omposd of lls that haror doul mmran_oundd
nuli. For that rason thy ar ukaryots (u=tru, karyon=krnl, sd;
Grk). All atria hav DNA, ut thy lk oth th nular mmran
and th tru hromosoms whr th DNA is pakagd with spial
protins.
Batria, as a group' are far mor mtaolially divrs than plants' ani-
mals, and fungi. Whras all th lattr rath oygn' som atria an
rath sulfat' aron dioid, nitrat' or vn arsnat. Batria, dspit
thir small siz, mor anstrl lifstyl, and rputation as parasits and
536 dosmbioti rigin of ukarotes

fid with various lvls.

diffirrlt to prov, sym
swimming motility (s
ond, now provn dfin
ll, thus allowing th t
nt and tlrrfor th t
limitd to th anstors
our plant grn, ld t
quisition of photosynth
ukaryots, alga, and ;
But how did tli dfir
Th first nulatd mil
Th nuloytoskltalorganellar systmof ukaryots.h systm,y frnt kinds of atria
hypothesis,volvd from a symioti mrger of a motil uatrium(a sulfid- lakd oxygn). On t1
oxidizing spirohttht bamth undulipodium)with n arhaarrium hydrogn sulfid gas fr
(a sulfidognithrmoaidophilodsndantwith hermoplslzasp.).S volani origin. This or1
argulis t al. 2005. tion, hypothtially aq
y symiognsis.Th s
Sugars' atat' or orga
grms' ar in fat mor profiint and oadly skilld at many hmial tasks sulfidogn, was a whiz l
than ar largr lif forms. minut. Th nulusitst

ARLIs NULAD ELLs

How might our atrial anstors hav volvd to form th rlist nul-
atd lls? Th volutionary pross always involvs th apaity for rampant
population growth far yond th nvironmnt to sustain it (i.., ioti po-
..hks', growth
tntial),th gnrationof inhritd variation, and th on of
the population (inluding its variants) y th failur of th ioti potential to
rahd. The inhibition of maximal population growth y limination of
many' for whatvr rason' is alld natural sltion. Naturl sltion
maintains halthy rprodutiv populations f many typs of organisms' ut
it dos not produ thm in th first pl.
.!7l1,
thn, how is inhrited variation gnratd? Although random muta-
tion larly xists and has n prodigiously doumntd, inhritd variaion
is gnratd in many mor diffrnt and important ways than via random
mutation. Indd, random mutation almost rtainly did not produ nul-
atd lls from atia.Rathr, th maior mod of volutionry variarin in
atria that ld to th first nulatd lls is likly to hav n via a pross
alld symbiognesis. This pross is allid to symiosis' whih is th olog-
ial rlationship twn ings of diffrnt typs (diffrntly nanrd organ- -''!*\.

isms). Symbiognsisis simply a protratd,long-lastingphysial assoiation,

Th [9(2)+2] rnirturr
usually twn diffrnt spis.It is n evolutionry, Darwirrian pross. sn witlr eltron mirtl
Symiognesis rfers to th apparan of a nw trait, havior' tissu' or- of volutilnary importal
gan, organism, spis, or othr highr taxon as a sult of an idntifial ili. zoospt-lrsof masti
long-trm symiosis. Svral volution-hangingsymiosshav n idnti- pojtions suggsts its
 ndosmbiotic rigin of uhrotes 57

fidwith various lvls of assuran.Although th arlisthang is th most

diffiult to prov' symiognsis ld to th ukaryoti organism itslf, with
swimming motility (sfigur low) nd th apaility for mitosis; th s-
ond, now provn dfinitivly, ld to th prsn of mitohondria inside a
ll, thus allowing th ll to rath oxygn. Th last symiosis, th most r-
nt and thrfor th on aout whos history w ar most onfidnt' was
limitd to th anstors of alga and plants. This symiosis, whih rndrd
our plant grn, ld to th prsn of hloroplasts insid th lls. By a-
quisition of phtosynthti prokaryots (yanoatria), th photosynthti
ukaryots,alga, and plants appard on arth.
lliiliiilt But how did th dfining fatur of a ukaryot' th nulus itslf, volv?
loL
Th first nulatd miroorganisms likly volvd y th mrgr of two dif-
karyots.Th systm' y frntkinds of atria ut undr anoxia (i..,nvironmntal onditions that
I a motile uatrium(a sulfid. lakd oygn). On typ of trium, apal of frmntation, produed
l i u m )w i ha n a h a r r i u m hydrogn sulfid gas from amint lmntal sulfur glouls, proaly of
tt'T bermoplsm sp.). See volani oigin. This organism, alld a sulfidogn ausof its gas produ-
tion, hypothtially aquird an ntir gnom of anothr typ of atrium
by symignsis.Th sond organism also livd on organi foodstuffs (.g.,
sugars' atat' or organi aids). But th sond atrium, whih is not a
y skilld at many hmial tasks sulfidogn, was a whiz at loomotion: it swam mor than 100 miromtrs a
minut. Th nulus itslf volvd only aftr intgration, that is, aftr th first

llvd to form th arlist nul.

nvolvsth apaity for rampant
l n t t o s u s t e i ni t ( i . . 'b i o t i p o -
..hks'' on growth of
, and th
. failur of th ioti potntial to
rlation growth by limination of
'rl sltion. Natural sltion
f many typs of organisms, ut

ratd?Although random muta-

d o u m n t di'n h i t dv r i a t i o n
rportant ways than via random
rtainlydid not produ nul-
l o d o f v o l u t i o n a yv a r i a t i o n i n
likly to hav n via a pross
symiosis,whih is th olog-
typs (diffrntly namd organ- -&.t.
long-lstingphysial assoiation,
Th [9(2)+2]mirotuularra of all undulipodiain transvrsthin stionas
volutionary,Darwinian pross. snwith ltronmirosopy.Th osrvationthat this sme(multigenitrait
t n w t r a i t , h a v i o r ,t i s s u ,o r - of evolutionaryimportan)is onsrvdin sprmtails' vrtratsnsory
on as a rsult of an idntifial ilia, zoosporsof mstigotmolds, and myriad othr intrinsiallymotil ll
ging symbioss hav n idnti- projtionssuggstsits ommon anstry.
58 dosmbioti rigi of ukrotes

atrium (th sulfidogni arhabtrium) mrgd with th swinrming

atrium to fornr th symioti anstor of all nulatdlls.
Th arlist produts of th first proposd symiognti mrgr wr ami-
tohondriat protists, lls that did not possss mitohondria. Th group'
alld arhaprotists (argulis and Shwartz 1998), still survivs in airlss
mds and anoi inrstins.Many gnra, dsndaIrtsof ths mitohon-
drit lls, still liv today, ut thy rnust sought irr strng haitats .
aus thir rlativs with mitolrondria in this oygn-rih world gratly
outnumr thm. Indd, ntir familis suh as th Trihomonadida and
Calonymphida still thriv in organi-rih anoi haitats suh as th guts of
trmits and wood-atins roahs.

PRoo|s D|VRslY: KEY o ELL EvoLU|oN

Th grat group of rlit nulatd miroorganisms, th arlist nulatd

lls, and thir xtant dsndants is alld Prototista. This group, whih to-
day is stimatdto ontain som 250'000 spis,is onsidrdits own king-
dom in th fiv-kingdom lassifiationshm.Th Prototists inlud all th
alga, th so-lld protozoa' th slim molds, and many mo Osur
.Wos
groups. Th thr-domain molular-biologial lassifiationof arl
lumps plants' animals, and fungi togthr in a singl kingdom and th pro-
.W.os
totists are put with all nulatd organisms into Dornain ukary.
sparatsth prokaryots into two trial groups, sd mainly on diffr- z,
ns in th dtails of thir riosom's RNA. on is th Arha (= n..1'".- ,
;
(J
atria) and th oth is ubatria' ln th sh of nulatd ll origins
IE
n(
outlind aov, th mrgr was twn an arhaatrium (th sulfido-
gni arhan, similar to today's Thermoplsm idophil) and a ua-
trium (similar to tody's Spiroht,a genus of snaklik swimmrs).
ARAAR
Th prototists inlud all photosynthti ukayots that d not grow from
mryos, as plants do (s figur)' Familiar arnpls inlud diatoms; rd, L**--*
grn, and rown sawds (inluding klp); hrysophyts; phytoplankton;
and all othr alga. (Th blu-grnsar not inludd; thy ar not alga ut origin of ukaryotsfror
yanoatria Protozoa (an obsolt trm insofar as i suggstsanimals), ili- thory. Lft to right: Th
ats, slim nts, foaminifra, hytids, and mny othr vn mor osur uatrilto form rir
Intllularmotility,in
ta-xa ar in th grat kingdom of Prototista. It is in this unruly grouP of u-
of digestionof oxygn-b
karyoti miros and thir many nrultillular dsndantstht so many fa-
mid-Protrozoi.Somo
turs of nulatd organisms volvd: mitohondriat and plastidi lls, that om th ansto
mitosis and mioti s, animal ll juntions (sptatjtrntions,dsmosoms)'
and animal-styl multillularity. vn ys (in rtain dinoflagllats), ll
Pntrationdvis (in Apimplans, th group to whih th malarial para-
sit longs),and hunting and shll-making (in th prdatory and gglutinating sdimntsof suh wll-p
foraminifra' rsptivly)volvd in ths groups of prototists. son t l. 1999) or th s
ypothtially, th origin of nulatdlls (ukaryosis)ourrd y th mid- Pulishd vidn d
dl of th Protrozoi on. Th dats for volution of elny innovation must oigin f th nultd
st y osrvtions in tlr fossil rord. Tl.rrfor,it is likly that th symiog- tthrd in a puliar s
nti volutionar squndsidaov ourd prilr to th dposition in totists' alld th kar
 dosmbioti rigi lf ukrots 5J9

L) mrgd with th swimming

'll nulatdlls.
y m i o g n t im
r g rw r a m i -
;ss mitohondria. Th group' @
# @

r,1998\' still survivs in airlss

:sndantsof ths amitohon-
sought irr strang haitats -
:his oygn-rih world gratly
:h as th Trihomonadida and
i haitatssuh as th uts of
g
o
N

!E
,
t-
(}
LL VoLU|oN

ganisms, th arlist nulatd
ototista.This group, whih to-
is, is onsidrd its own king.
:. Th prototists inlud all th
llds, and many mor obsur
.s7os
:l lassifiation of arl
a singlkingdom and th pro-
; into Domain ukarya. \os
groups'basd mainly on diffr-
' on is th Arha (= arha-
;hmof nulatd ll origins
L arhatrium(th sulfido-
rcm cidophil) and a ua- t
Lsof snaklikswimmrs). o
lkaryotsthat do not grow from
AR}|AARlA s
UBAR lA
ampls inlud diatoms; rd,
; hrysophyts;phytoplankton;
inludd;thy ar not alga ut origin of ukaryotsfrom spifitriallinags:srialndosymioti
ofar as it suggstsanimals), ili- thory. Left to right:Thrmoplasma-like arhabatriamrg with spiroht
many othr vn mor osur uatrialto form amitohondriatprotists in th low Protrlzoion.
Intrallularmotility, inludingphagoytosis'potntiatsth ingstionut lak
It is in this unruly group of u-
of digstionof oygn-rthing atrialanstorsof mitohondriay th
Lrdsndantsthat so many fa-
mid-Protrozoi.Som of thsaroiprotistsingstand rtainyanoaria
lhondriat and plastidi lls, that omth anstorsto th plastidsof algaand plants.
lsptatjuntions, dsmosoms)'
(in rtain dinoflagllats)' ll
oup to whih th malarial para-
r thprdatrynd agglutinting sdimntsof suh wll.prsrvdmirofossils as Vdlospberidium (Samuls-
lups of prototists. son t al. 1999) or th spiny sphrsin th Doushantou.
ukaryosis)ourrd y th mid- Pulishd vidn doumnts that th first struturs that appard in th
ution of any innovation must b origin of th nulatd ll wr not ff nuli. Rathr, th nulus gan
r,it is likly that th symiog- tthrd in a puliar strutur' an organllar systm still insid many pro-
:urrdprior to th dposition in totists' alld th karyomastigont. Th karyomastigont is an xampl of
'40 ndlsmbioti rigi of krlts

D:rrwinian imprftiorrand oddiry from wlrih pth f history rrry . Sry' D. G. l987. Ph,vlog
onstrutd.Th signifianof th karyolasigont,n organllr systm in ukayoti nuloytpl
Netu Yorh dm of !
whih a nulus is mddd, was dsrid in th arlY yars of th twntith 'Wos,
.202. )n tlr vol
ntury (Janiki 1915). inimlly th karyornastigontonsistsof nulus,i.t Sicct'sUS 99: 8742-
nular otllltor (alld a rhizoplst in arly potozoologial litrtur)'irnd
an undulipodium (th 9+0 kintosom/ntriol and its shft, th 9+2 miro-
tuular aonm). (Som prf to writ this univrsal pattrn of ninfold
thology and th stu
symmtry of th mirottrulsin th shaft m()rauratlyas [9(2)+2].) Whn
th ntriol in mitosis dvlopsinto th kintosom of th motility organll, \/hil rlirny individuals l
w ar sing th lgay of th swimmr omponnt of th arlist ukaryots. Austrian Konrad Lornz
h strutur and lravior of prototists tht lak mitohorrdri undr (1907-1988) ar rgrd
anoi orditilnsand today's motility s,vmiossin many organisms with thology as a rognizds
swimming batria' sllh othrwis odditis and puliaritis, ar undrstood trvld to Lidn in 193
in th ontxt of thir volution. h nuleus itslf is most likly produt <lf ollaortors. L<rrnzwas
gnti-lvlintgation of batrizrlsymior-rts(arhtriLrm sulfidogn and his visionary br.rlth
plus swimming uatrium) tht gan as part of tlr karyomstigont from primntal appoah to th
whih it was rlasd. As wll as bing prat
This volutionary snario fo th origin of ukaryots is asd on th in- rvrot out its mthodol
valuai dsriptionsy Harvard profssoLmul R. Clvland (1892_|971I ogists slrould attmPt to a
and Harold Kiry Jr. (1900_1952),hairman of th Dpartmntof Zoo|og at vlop? Wht ar its imn
th Univrsity of alifornia at Brkly. lvland providd dtails of ll volv? Lrnz strssdt}
motility (movrnntsof th kayomstigontszrnd othr ll parts irr mitosis) having spontan()usly
and Kiry am to undrstnd :rlonymphid volutio y rognizing th ail- himslf shokd y th d
ity of karyomastigonts to rprodu indpndntly of th nuli in mitohon. in havioristi psyholog
drizrlll lingsthat proaly nvr aquird mitohondria. Prsrvationof vlopmnt of an thogra
th pulishd work of ths sholars (inluding th unprdrtd16--rm hvis-is th stating
blak and whit films y lvland) would of intrst to thos who wish to In 797, Lornz and T
dlv furthr into th details of ll evolution. Grman zoologist Karl vt
dislvisonrnig or
havior pattrns.''Von I
BIBLIOGRAPHY
languag of hony bs.\
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i<ltrihprd<li:rnd its ssoiatdmioorgnisrs. Prodittgs of th
onstrut thir phvlogny
RllSoietf L-ldo, SrisB 157:668_683.
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Jniki,. 1915.aminationsof parasitiflagllats.
Dues<lui, Prioni,Stphttn<lmph, lomph. on th p:rraasl morphology in ronstru
appartlls.)n th oIrstitutinof tlrnulr"rs
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Wisssbftlih Zoologie | 1Z:57_691' arly thologistsWr
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and Tinrgn mt at a W
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Mrgulis,L., . . Doln' and J. H. Whitsid.200.5...lmprftions dditis''
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in th originof th nulus.Pillbiolog1', 1: 175-191. rpatdly liitd s
L., and D. Sagan.|998. Wht ls Se?Nw York: Simonand Shustr.
l!,Iargulis' t i u l rp k i n g h e v i t '
2000.'V/htls Lif? Brkly:Univrsityof lifrniaPrss. simpl modls of th adul
argulis, L., rrdK. V. Shrvartz,1998.Fiu igdms: Illustrtduid trthc o l l p tt r i . r f i d r i n
Pbl of Life lrtb.3rd d.Nw Yrk:W. . Frmanand Company.
s i g n s t i l n u l ia n d f i s d t
Samulsson, .I.,P. R. Darvs,and G. Vidl. l999. Orgni-rvalld mirofossilsfrom
th ProtrozoiThul suprgrouP'Northwst Grnlan1. PrmbriRsrll undrstood in th on
96:1-23. volutionary ontts fat

h a path of history may r-
r i g o n t .a n r g a n l I a s s t m i n
th early yars of th twntith
astigontonsistsof a nulus, a
protozoologiallitratur),and
l and its shaft, th 9+2 miro-
; univrsal pattrn of ninfold
auratl1.
as L9(2)+2]').W.hn
lsom of th motility organll,
lnnt of th arlist ukaryots.
-hat lk mitohondria undr
ioss in many organisms with
rd puliaritis'ar undstood
tslf is most likely a produt of
s (arhaatriumsulfidogn
Lrt of th karyomastigont from
ukaryotsis asd on th in-
nul R. Clvland (\892_1971)
f th Dpartment of Zoo|og at
rland providd dtails of ll
rnd othr ll parts in mitosis)
olution y rognizing th ail.
lntly of th nuli in mitohon-
Lmitohondria. Prsrvation of
ing th unprdntd 16-mm
lf intrstto thos who wish to
in strutur of th flgllat
oorganisms.Prodigs of th
83.
s.Part II. Th gnr
ilompb.on th paraasal
rnd amitsis.Zitshrift fr
l'ites. Uiursit of |ifor
..Imprftior-rs
05. nd odditis''
7 5-19] .
w York: Simon and Shustr.
alifornia Prss.
Ioms: lllustrtd Guid tl tb
H. Frmnnd ompany.
trgani-walldmirofossils from
]rnland. P rembri Re serh
thrllg d the Stud of Bhuiorl uolutio 541
Sary,D. G. 1987. Phylogntind hnotypi rlationshipsbtwn th
ukaryoti nuloytoplasmand thrmophili Arhaatria.ls of the
Ne York Adm rlf Sies 5 168_\79.
Wos, . 2002. On th volution of lls. Prodigs of tb Ntill dem of
-L.^. d D.S.
SisUS 99: 8742_8747.
thology and th study of bhavioral volution
Whil many individuals hav ontriutd to th dvlopmnt of thology,
Austrian Konrad Lornz (1903_1989) and Duthman Niko Tinrgn
(1907*1988) ar rgardd as its ntral figurs' whos fforts stlishd
thology as a rognizdsubdisiplinof volutionary iology. \hn Lorenz
trvldto Lidn in 196, h mt Tinrgn and thy am frinds and
ollaortors.Lornz was known for his grat passion for wathing animals
and his visionary radth,whil Tinrgn rought a mor analytial and -
primntalapproah to th study of animal havior.
As wll as eing pratitionrs of thology, oth Lornz and Tinrgn
Wrot aout its mthodolgy. Tinrgn idntifid four qustions that thol-
ogistsshould attmpt to answr aout any spifihavior:How dos it d-
vlop? Nhat ar its immdiat auss? \/hat is its funtion? How did it
volv? Lornz strssdth importan of spnding hours wathing animals
havingspontanouslyfor formulating any hypothss'and h profssd
himslf shokd by th dp ignoran of animals that h thought h dttd
in bhavioristipsyhologistssuh s John B. Watson (1878_1958). Th d.
vlopmnt of an thogram for a spis-a standardizd list of osrval
haviors-is th starting point for thologial rsrh.
| 197, Lornz and Tingn shard th Nobl Priz for mdiin with
Grman zoologist Karl von Frish (1886-1982). Thy wr itd ..for thir
disovrislnrningorganization and liitation of individual and soial
havior pattrns.'' Von Frish is st known for his disovry of th dan
languagof hony s.Von F'ish'sstudnt,Martin Lindaur, ondutd a
omparativ study of th dans of svral losly rltd spis to r-
onstrutthir phylogny, similar to Lornz's study of havioral homologis
in t tide (waterfowl). Th ida that havior ould as rlil as
morphology in ronstrutingth rlationshipsamong spisws signifiant
for th aptan of thology as an important part of volutionary iology.
arly thologists wr intrstd in th onpt of instint; indd' Lornz
and Tinrgn mt at a workshop on this topi. Lornz is ssoiatdwith th
ida of a fid ation pttrn-a ompl havior that an rlialy and
rpatdlyliitd y a spifi sign stimulus. Tinrgn showd that a par-
tiular pking havior of hrring gull hiks ould liitd y rlativly
.Whil
simpl modls of th adult ak with a prominnt rd spot. th gnral
onpt of a fid ation pattrn applis aross numrous spis'partiular
sign stimuli and fid,ationpattrns ar highly spis-spifiand an only
b undrstood in th ontt of th partiular dvlopmntal,adaptiv, and
volutionary ontts fad y mrrs of th spis.Lornz's onpt of
s42 tholog d th Stud of Bebuiorl uolutiln

instint was strongly ritiizd in th 1950s y Amrian psyhlogistDanil ojts of study, lik a
.Wilson's
Lhrman for ing insuffiintly attund to dvlopmntal prosss.Tinr- ward o. xtn
gn's inlusion of dvlopmnt among his four qustions rprsntdhis at- insts to xplain huma
tmpt to aommodat Lhman's ritiqu. Susquntly' many yars of o8, a fild that also is t
fruitlss ..natuf vrsus nurtur'' dats nd onflits ovr dfinitions Anothr important of
hav mad urrnt thologists lry of th onpt of instint. Nvrth- funtion known as nut
lss, th study of inhritd havioal adaptations is still n lmportant part partiular haviorl tas
of thology. th startl rspons of fi
Aras of partiular urrnt intrst and ativity within thology inlud tional haring of owls,
prdator-pry intrations and th various systems of antiprdatory alarm mols.
alls found in a wid rang of spis, th soial dominan struturs that ..ognitivthol<
Also,
also vary widly aross diffrnt spis,ompartiv studis of soial play, to dsri th rintrodl
th mmory dmands of food ahing nd rtrival, tool manuftur and volutionary study of i
us, and th xistn of ulturally transmittd traits. onduting arful phys
thology an undrstood y ontrast to altrnativ approahs to th Nxt, h turnd his att
.!ithin
study of havioral volution in iology and psyholgy. iology, - M a n y d i s a g r w i t h h i s
havioral olgy applis population-lvl modls of volution to th undr- and anthropomorphi.
standing of animal havior. Bhavioral ologists frly orrow modls in human psyhology v
from onomis and gam thory, whih rprsnt individul organisms as instrumntal in gtting
rational optimizrs and allow analysis of intrations twn nvironmnt tions aot th volutr
and havior on rprodutiv fitnssand th distriution of bhavioral traits. ranging from ognitiv
Unlik traditional thology, havioral ology has n lss onrnd with primats.
immdiat auss of th havior of individual nimals or its dvlopmnt.
|LIoGRAPHY
hanisms rsponsilfor gnratingoptimal havior ar oftn simply as-
sumd and not plaind. Nvrthlss,thr hs n signifiant rossovr Bkoff,., . Alln,and
twn th two sudisiplins, with many of th originators of havioral mpiril nd Theor
N4ITPrss.
ology having first traind as thologists.
Burkhardt, k. 2005.Px
omparativ psyhology also taks an xpliitly volutionary approah to Fondig of thobg^
animal havior. Howvr, unlik iologists, who tnd to intrstd in dif- hny'D. L., and R. M.
frnsamong individuals and among spis,psyhologists ar traind to Soil id. hiag
lok for gnral pattrns of havior and prinipls of larning' somtims Frish'K. von. 1.967.h
.!7hr hadt'ik,trans.t
rfrrd to as laws of larning. omparativ psyhologists do s dif-
G r i f f i n ,D . R . 2 0 I . n m
frns twn spis, thy often rgard ths as rvaling stps on an vo-
Univrsityof hiag
lutionary laddr to full-lown human ognition, and thy ar oftn drwn to Lhrmn,D. S. 1953.A
invstigatingtasks to mark thrsholds of ognitiv sphistiation.Thus, for Qurtrl Ruituof
ampl, thr has n muh intrst among omparativ psyhologistsin Lindaur,. 1961.om
finding out whih spis an rogniz thmslvs in mirrors, an pass a UnivrsityPrss.
Lornz' K. 1981.The Fu
fals lif attriution task, or an larn a human languag.any thologists
Tinrgn'N. 1963.on t
prfr to osrv animals in natural or naturalisti haitats. Som sintists, ir1lshll ogi 20: l
to found on oth sids of this divid, ar pluralists who livthat th in- .Watson,
J. B. 1913.Psy
tgrativstudy of animal havior rquirs oth field and laoratory invsti- 158-177.
gation; othrs ar lss umnial. W i l s o n , . o . l 9 7 5 . S o
thology has had svral important offshoots. Both Lornz nd Tinbrgn
xtndd thir approahs to th study of humn havior: Lornz in his
study of aggrssin, and Tinrgn (with his wif lisath) in studying
autisti hildrn. Th willingnss of tholoists to trat humans as sintifi
 thobg nd tb Std of Bhuklrl uolution 54

y Amrian psyhologist Danil ojtsof study, lik any othr animal, also importantly forshdowd d-
vlopmntalprosss.Tinr- ward o. \/ilson'stnsion of volutionary modls of th soial havior of
ur qustionsrprsntdhis at. insts to plain human soiality, thus giving ris to th fild of soioiol-
. Susquntly,many yars of ogy, a fild that also is tightly onntd to havioral ology.
and onflits ovr dfinitions Anothr important offshoot of thology is th omparativ study of nural
onpt of instint. Nvrth- funtion known as nrrrothology. Spifi adaptations of nural systms to
ations is still an important part partiular havioral tasks hav n dsrid in a varity of spis, suh as
th startl rspons of fish, th oordinatd voalizations of frogs, th dir-
'tivity within thology inlud tional haring of owls' or th unusual snsory mhanisms of star-nosd
jystms of antiprdatory alarm mols.
oial dominan struturs that Also, ..ognitivthology'' is th lal oind y Donald Griffin (1915200)
nparativstudis of soial play' to dsrith rintrodution of issusof ognition and onsiousnssto th
rtrival,tool manufatur and volutionary study of animal havior. Griffin mad his arly rputation
:d traits. onduting arful physial analyss of th holoation apabilitis of ats.
o altrnativapproahs to th Nt' h turnd his attntion o qustions of animal mind and awarnss.
psyhology.\ithin iology, - any disagrwith his approah us thy find it xssivlyandotal
ldls of volution to th undr- and anthropomorphi. Nvrthlss, th ognitiv rvolution that took pla
:ologists frly orrow modls in human psyhology was slow in oming to th thology, nd Griffin was
p r s n ti n d i v i d u a l o r g n i s m s a s instrumntal in gtting othr sintists to tak sriously th ida that qus-
:rtionsbtwn nvironmnt tions aout th volution of mind ould addrssd y studying topis
distriutionof havioral traits. ranging from ognitiv maps in honys to intntional ommuniation in
gy has n lss onrnd with prlmats.
ual animals or its dvlopmnt.
B|LIoGRAPHY
rl havior ar oftn simply as-
. has n signifiantrossovr Bkoff,., . Alln, and G. . Burghardt,ds.2002. Th ogitiuil:
mpiricl d TheortilPrs1letiuslnimI ogitio.amridg,A:
of th originators of havioral
I Prss.
Burkhardt'R. 2005. Pttrsof Bhuior:Konrd Lornz, Niko Tibrg,d the
lliitly volutionary approah to Fodigof thrllog,Chiag<r: Univrsityof hiago Prss.
who tnd to intrstd in dif- Chny,D. L.' and R. M. Syfart.2007. Bbootphsis: T uolutionof
is' psyhologists ar traind to Soil id. hiago:Univrsityof hiago Prss.
r i n i p l so f l a r n i n g ,s o m t i m s Frish,K. on. 1967. The De Lgugnd rittioof Bs.L. .
hadwik,trans.amridg,MA: HrvardUnivrsityPrss.
arativ psyhologists do s dif-
Griffin, D. R. 2001. iml inds: Bod ogitio to osiosess.hiago:
tsas rvalingstpson an vo- Univrsityof hiago Prss.
ion, and thy ar oftn drawn to Lhrmn,D. S. 1953.A ritiquof Konrad Lornz'sthoryof instint.e
gnitiv sophistiation. Thus, fr Qurterl Ruieluof Biolog 28: 337_363.
1g omparativpsyhologists in Lindaur,\4. 1961.ommuitiomogSoilBs.amridg,A: Harvard
UnivrsityPrss.
mslvsin mirrors, an ass a
Lornz,K. 1981. b Foudtionsof thology.Nw York: Sringr-Vrlag.
man languag.Many thologists Tinrgn,N. 1963. on th aims and mthodsof thology.Zitshriftfiir
l i s t i h i t t s .S o m s i n t i s t s , Tierpsologie 2: 414 63,
.\X/atson,
'luralistswho livthat th in- J. B. 1913. Psyhologyas th havioristviws it. PshologilRuie2:
, o r h6 l d a n d | o r a t o yi n v s t i - t58-177.
.Wilson,
. o. |975. Soiobiolog.amridg,A: Harvard Univrsityo,.,':-..o.
rots' Both Lorenz and Tinrgn
human havior: Lornz in his
his wif lisath) in studying
ists to trat humans as sintifi
544 ugis

tr and tmpramnt \,
ugnis
matrial (th natur of
ugnis is fundmntally th grrralization tO humans of th pratis of at irth. H lso assum
plant nd animal rdrs'who popagtth st and ull th worst of thei mnts, and that thos l
stoks. Th ida that suh mthods might applid to humans has a long produing at a shokil
history, raing ak at last to th anint Grks. Thus, in Plato's Repub- don to Prvnt it, ivil
li (1945,157-158), Sorats tlls Glauon that..anything lik unrgulatd tak harg of thir ow
unions would a profanation in a stat whos itizns lad th good life''' Th ida that huma:
and Glauln knowldgs ht h is arful to 1i1tonly th st of his nW-nor wr dat
sporting dogs and gm irds so as to ilvoid th othrwis rapid dtrioration losphr Jhn Stuart l
of his stok. man natur has takn
Apart from a fw fforts to systmatially shap human hrdity, s in th plial. | Priipls o,
program adoptd by th utopian ommunity at onida, Nw York' in th ..of all th vulgar mod
1860s, ugnis rmained largly onptual until th turn of th twntith moral influns on th
ntury. It ws only thn tht <lrganizationsxprssly dvotd to this aim th divrsitis of ond.
wr stalishdin th Unitd Stats,Britain, and Grmany. Thr ar multi- ill was swimming a6
pl rasols why th nd to implmnt ugnis on a larg sal am to laim that natur trurr
sm urgnt around this tim, ut prhaps th nrost important Was a nw al studis of inhritan
worry aout th possiility of iologial dgnration. Darwin, who wrot in
In his oz the origi of Spcis (1859), harls Darwin argud that im- agr with Franis Ga
provmnt in plants and animals rsultd from a fir omptitiv struggl, due only a small ff
.What
a pross napsulatdin th famous xprssion ..th survival of th fittst,'' tis ar innt.''
oind by Hrrt Spnr in 1866 nd latr adoptd y Darwin' Although anity in whih it ap;
Darwin did not disuss his own spisin the rigin' many of his ontmpo- popl that th Pros
raris wondd aout its impliations for humans. ln partiular, thy frttd that th mentally and :
ovr th possiility that a rlaxation of th struggl mant that human But whil Glton so
progrss would slowd, haltd, or vn rvrsd. Darwin himslf was with th risis, and th
dply disturd y this prospt' arrd in he Dscet of (1871,168) h than disouragingth
pulilv xprssdhis onrns out th ountrsltivffts<lfharity, ..p
what Galton alld
vaintions gainst smallpo, th uilding of asylums for th sik and in- any squamishnsso
san' and othr aoutrmnts of ivilizd Soity' noting that as a sult' middl-lass popl rr
..th
wk mms of ivilisd soitispropagat thir kind. No on who hrd to th prvailin1
has attnddto th rdingof dmsti animals will dout that this must th stat should kr
highly injurious to th ra of mn.'' govrnmnt into th il
Darwin's ousin, Franis Galton, ws among th first to xplor th soial om an antistatistwo
impliations of th thory of volution y natual sltion.Galton notd in am a mattr for lg
his mmois that h had n inspird y rading t rigi to pursu a th inrasing apta
long-standing intrst in th topis of hrdity ..and th possil improv- whih smd to ols
mnt of th Human Ra.'' In ..Hrditary Talnt nd haratr'' (1865), prvious to th nviro
pandd to th ook Hereditr Geius (1'869), Galton showd tht mn lin in p<lpulationqu
who had ahivd distintion in sin. litratur. and th law wr mor unfit for srvi in th
likly tlran mmrs of th puli at larg to hav had mal rlativs who Unitd Stats, th low
wr thmslvsminnt. In a raft of latr works, Glton tndd this r- and astrn urop ol
sarh and dvlopd what h saw as its orollris. Galton thought that his I smd to onfirm f
inquiris provd byond dout that all human ailitis and traits of hara- rs who wr flm

to humans of th pratis of
st and ull th worst of thir
applid to humans has a long
]rks.Thus, in Plato's Repb-
hat ..anything lik unrgulatd
ls itizns ld th good lif,''
Ll to mat only th st of his
o t h W i s ra p i d d t r i o r a t i o n
hap human hrdity, as in the
'at onida, Nw York, in th
until th turn of th twntith
prssly dvotd to this aim
and Grmany. Thr ar multi-
nis on a larg sal am to
h most important was a nw
nration.
harls Darwin argud that im-
m fir omptitiv struggle'
ion ..th survival of the fittst.''
adoptd by Darwin. Although
rigin, many of his ontmpo-
nans. In partiular' thy frttd
struggl mant that human
rvrsd. Darwin himslf was
D e s e to f ( 1 8 7 1 , 1 6 8 ) h
lntrsltivffts f harity,
lf asylums for th sik and in-
oity, noting that as a rsult,
Pagatthir kind. No on who
ralswill dout that this must
1gth first to xplor th soial
:ural sltion. Galton notd in
ading the rigin to pursu a
ity ..and th possil improv-
Ialnt and haratr,' (1865),
869), Galton showd that mn
ratur' and th law wr mor
hav had mal rlativs who
rorks,Galton xtndd this r-
llaris.Galton thought that his
n abilitisand traits of hara-
ugics 545
tr and tmpramnt Wr transmittd fom parnt to hild in th hrditary
matrial (th natur of whih was thn unknown) and thus wre largly fixd
at irth. H also assumd tht popl varid gratly in thir hrditary ndow.
mnts, and that thos with th fwst talnts and th worst hratrs Wr r-
produing at a shokingly rapid rat. It followd that if somthing wr not
don to prvnt it, ivilization would ollaps. Th solution was for humans to
tak harg of thir own volution' a pross h trmd eugeics in 1883.
Th ida that human diffrns Wr innat and immutal was nothing
..ivilisation'' (1835), th phi-
nw-nor wr datsaout its validity. In
losophr John Stuart Mill argud that th study of history tahs that hu-
man natur has takn an infinit varity of forms and must thus highly
plial.Io Priipls of Politicl coom (1848, 319), h famously assrtd:
..of all th vulgar mods of saping from th onsidration of th soial and
moral influnes on th human mind, th most vulgar is that of attriuting
th divrsitis of ondut and haratr to inhnt natural diffrns.', But
ill was swimming against th tid. Thus Galton's originality lay not in th
laim that natur trumps nurtur, ut in th ffort to prov through statisti.
al studisof inhritanthat th laim was tru. On of thos onvind was
..inlind
Darwin, who wrot in his autobiography (1958, 43) that h was to
agr with Franis Galton in living that duation and nvironmnt pro-
du only a small fft on th mind of any on, and that most of our quali-
tis ar innat.'' !hat gav Galton's work its signifian was th limat of
anxity in whih it appard: thr was widsprad far among middl-lass
popl that th pross of sltionhad n haltd in ivilizd soitis,and
that th mntally and morally Worst Wr now swamping th st.
But while Galton soundd an alarm, h had fw pratial proposals to dal
with th risis, and ths nouragd th giftd to hav mor hildrn, rathr
than disouraging th stupid and rklss from having any. Th mphasis on
..ngativ'' ugnis did not rflt
what Galton alld ..positiv'' rathr than
any squamishnss on his part aout th lattr. In Galton's day, thos
middl-lass popl most attratd to ugni prinipls also gnrally ad.
hrd to th prvailing Vhig idology, arding to whih th funtions of
th stat should kpt to a minimum. Proposals to xpand th rah of th
govrnmnt into th intimat sphr of family lif would thus hv to ovr-
om an antistatist worldviw. That is a ntral rason why ugnis only b-
am a mattf for lgislation in th twntith ntury. Anothr rason Was
.Wismnn's ..hard''
th inrasing aptan of August thory of hrdiry,
whih smd to olstr Galton's viw that th hrditary matrial was im-
prvious to th nvironmnt. A third rason was apparnt vidn of a d-
lin in population quality. In Britain, th larg numr of rruits dmd
unfit for srvi in th Bor War (1899_1902) gnratdmuh alarm. In th
Unitd Stats' th low sors ahivd y rnt immigrants from Southrn
.!7orld .!far
and astrn urop on IQ tsts administrd y th army during
I smd to onfirm fars that th ountry was ing inundatd y nwom-
rs who wr flmindd as wll as fund.
546 ugnis

In Britain, th land f its irth, ugnisrmind mostly a mattr of prop- |869. Hrditri,
aganda. Howvr, in North Amrir, Sandinavia' Grmany, astrn urop, L l n d r l n :N l . l l l l i l ] l l l .
and Japn, among othr plas, laws wr passd prmitting th ompulsory \98. rtlricst
Gillhm' N. !. 2001' l
strilization of th flmindd, riminals, sual dviats, and othrs
Birth of ugis. >
dmd unfit to prorat. |n th 1927 as of Bh u. Bll, te U.S. Suprm K v [ s ,D . l . 1 , 9 9 5 . l t h
Court uphld th onstitutionality of ompulsory strilization in dision rdit. Rprin <J
.!7ndll
signd y hif Justi olivr olms' a politial progrssiv. fa- Kli, $7. 201,.Btliltlig
mously Wrot: ,.It is ttr for all th world if, instad of waiting to xut rn of th tttttr1,,
Prss.
dgnratoffspring for rim, or to lt thm starv for thir imility, soi-
M i l l ' J . s . [ 1 8 3 5 ]1 9 7 7 . i
ty an prvnt thos who ar manifstly unfit from ontinuing thir kind.'' In Sturt ill. Vll. 18' l
th Unitd Stats, ugni argumnts wr also invokd in support of rstrit- Llnivrsity of Tlrtlttt
ing immigration frorn Southrn and astrn uop, argumnts that playd a [ | 8 4 8 1 1 9 6 5 .I '
rol (aithough how larg a on is datd)in passagof th national_origins pplictkls to St
quota systm adoptd in 1924. But ugni zal rhd its apex (or nadir) in Joh Sturt il/. Vols
Paul, D. B. 2003. Dlvin,
Gmany, whr troitis gainst nrntal patints' Jws, and othrs wr jus-
..raial hygin.'' Radik, eds., Th l
tifid y th ostnsibl nd for a program of ambridg Univrsiti
As th as of olms suggsts'sllpport for ugnison appald to di- Plttl. l 945. h RpttllI
vrsonstitunisand was invokd in suppt of disparatagnds:rh d- Univrsit1.Pss.
..flov'' as wll as traditional marriag, Trrrda, .' and P. J. V/in
fns of paifisrrras wll aS War'
RiI Ntioli rr
and ass to irtlr ontrol information and dvis as wll as thir rprs-
ntral uopn Un
sion. In its hyday in th 1920s and 1930s, ugnis garnrd support not
just from soial and politial onsrvtivsut also from Faian nd vn
arxin soialists and rformrs of many strips. In tinr' ugnism to
vo|ution: Th
quatd with its worst instntiations'and aov all, with Nazism. As a r-
sult, th word is today usually rsrvd for pratis that th spakr or Julian uly had
writr dtsts.Thus ritis of prnatal diagnosis dm it ugniswhil its ad- ollat vast amounts of
voats strnuously rsist th lal. And lgislation barring fist-ousin mar- lutil: Th ldern S\,r
riags is not rgardd as ugnis,notwithstanding its ompulsory hartr. rat n nylopdir
Sltivrding is still pratid, and in som forms it is widly pprovd, ing his survy togtl-r
ut on lgay of ugnis'assoiationwith atroitisis a distrtd disours Chishd among ths
in whih w shrink from alling ths pratis y thir rightful nam. strssing sltion'sirn
spially in Britain' as
ILIoGRAPY lution.
Brorg,G., and N. Roll-Hansn,ds.2005.ugisdth WIfreStte: Tlr rrylopdiap
SteriliztioPoli in Dmrk, Sdn,Norta,ltd Finlnd.Rv. paprak votd svralhaptrst
d. ast Lansing:ihignStatLlnivrsityPrss. m a i n t a i n v r i a t i o r - rI.I
,1'927.
Buk u. Bll. 274 U.S. 200. gnom lvl' suh as p
arlson, . A. 2001. The Ufit:A Histor of Bd lde, old SpringHaror, NY:
ogy' Huly ppriat
old SpringHaror LaboratoryPrss.
his for.rson organisrrrs
Darwin, . [1859]|964. th rigi of Spis.Introdutiorr y . Mayr.
amridg,A: HrvardUnivsityPrss. inftlrration aorrtlo
'h
[1871]198l. Dseto|-n,nd Seletil
i Rltioro S' stttturdhis suv:v.
Introdutionby J' T. Bonnr and R' M. May. Printor.r, NJ: Printon I i o n .H s r r s s dt i
UnivrsityPrss. ation. lins wr his s;
.With
1958. Th Autobiogrphof hrlsDnui, 1809_1882: rigil
a dad for. Ths
missionsRestord.N. Barlow' d. Nw York: arourt Bra.
Gltorr,F. 1865.Hrditar,v tlntnd hratr.
mill'slgzi|2: v y s , r v i t h H u [ y t 1i r
1 5 7 _ 1 6 6' 1 8 - 2 7 . Th hif virtu of ths

aind mostly a mattr of prop-
Lavia,Grmany' strn urop,
rssdprmitting th ompulsory
s, sual dviats' and othrs
f Buk u. Bll, te U.S. Suprm
rlsoy strilization in a dision
s,a politial progrssiv.H fa-
if, instad of waiting to xut
starvfor thir imility, soi-
t from ontinuing thir kind.', In
lo invokd in support of rstrit-
iurop, argumnts that playd a
l passagof th national-origins
al rahdits ap (or nadir) in
i n t s J, w s , n d o t h sw
..raialhygin.''
lr ugnison appald to di-
ort of disparatagndas:th d-
a s w ] l a s t r a d i t i o n Im r r i a g ,
dvisas wll as thir rprs-
ugenisgarnrd support not
ut also from Faian and vn
rips.In tim, ugnisam to
aovall, with Nazism. As a r-
r pratis that th spakr or
sis dmit ugniswhil its ad-
; l a t i o n a r r i n gf i s t - o u s i nm a -
nding its ompulsory harater.
m forms it is widly approvd,
ttroitisis distortd disours
s y thir rightful nam.
icsd tb WelfrStt:
,tu,nd Fild' Rv. paprk
'rss.
d lde. o|d SpringHaror, NY:
s.Introdutiony . ayr.
'lliuin RltilIlS,
Printon'NJ: Printon
i' 1809_1882:With riginl
:k:HarourtBra.
milln.sgzi|2:
vlution: Th Modrn Svnthsis 547
|869. Hrditr Gis: Iquir itltl lts Ltt,s d osqs'
Llndon: Marillr-r.
1'908. mor is of 1' Lif. Londt.ln Mthun.
Gillham, N. W.200l . l-if of Sir Fris Ito: Frlm fri 1llortio to th
Birth lf ugzis. oford: oford Univrsity Prss.
Kvls, D. J. ] 995' ] th Nm of gis: Glttis d the Uss of Hum
Hrdit. Rprint d. amridg, MA: Harvard Univrsity Prss.
Klin, W. 200,l'. Bildig Btter R: Gder, Sulit, d ugis from tb
Tur lf th tttr to th Bb Bool. Brkly: Univrsity of alifornia
Prss.
Mill, J. S. l183.5l 1977. ti|tsation'In .|.M. Roson, d', olleted'V/orks of lohn
Sturt ill. Vl. 1 8, sss rl Politis d Soit1,,Pt' I, 117-148. Toronto:
Univrsity of Toronto Prss.
[1848l 1965. Prii1llsof Pllitil oom, ith Som of Thir
pplitills to Srlil Phillso1l.ln J. . Roson, d., olltd Works of
Joh Sturt ll/. Vols. 2_3. Toronto: Univrsity of Toronto Prss.
Paul, D. B. 2003. Drwin, soial Darwinism, and ugnis.In J. odg and G.
j u s -
Radik, ds., h mbrirlg om1lio tl Dri,214_29. anrridg:
amridg Univrsit Prss.
Plato. 1945. Th l|publi of Plto. F. . ornford, trns. London: Oford
Univrsity Prss.
,,Blood
Turda, ., and P. J. \7indling' ds. 2007. d Homld'': ugertisnd
Ril Ntiolism i trl nd Soutrn L|rop, 1900_1940. Budapst:
Cntral uropan Univrsity Prss. -D,B.P.
uolutio: Th odr Sthsis (Julian Huley)
Julian Huly had ptional skills as a ommrial writr. knw how to
ollat vast amounts of informtion and how to xtrat ssntialpoints. o-
lutil: Th odern Sthesis (1942) is prft ampl. Huly didd to
rat an nylopdiaof rnt dvlopmntsin volutionary studis.Hold.
ing his survy togthr Wr a small numbr of arfully hosn thrads.
hrishd among ths was natural sltion. Although h mad othr points'
strssingsltion'simportan ws Huxly's ntral aim. This ook srvd,
spially in Britain, as a training manual for th ..synthti''thory f vo-
lution.
Th nylopdiaapproah in uolutiol? movd systmatially.Huxly d-
votd svralhaptrs to gntiphnomna,spiallythos that rat and
maintain variation. H thn shiftd to dsri auss of variation at the
.!(iith
gnom lvl, suh as polyploidy. a kground in dvlopmntaliol-
ogy, Huxly appriatd th importan of ths phnomna. Shifting from
his fous on organisms and ologial phnomna, uly nt onsidrd
information aout 1oalpopulations, varitis'and suspis.Thr thms
struturdhis survy:vriation, prosssof divrgn'and prosssof iso-
lation. strssdth inrportn of polytypi spis and ggraphial vari-
ation. lins wr his spial intrst,a phnomna h had hlpd populariz
a dad for. Ths stions of uolutirln read simply as litratur sur-
vys' With Huly trying to Ompil informtion s is don in a txtook.
Th hif virtu of thsstionsis thir radth.Hul mad tnsivus
548 volution:Th odrn Synthsis

of knowldg gland from his ontats' who spannd oth urop and th IBLIoGRAP|
nglish-spakingworld.
uly, J. |\942' 196
uolutiln Was no mr survey. Huxly plad natural sltion and adap.
Alln and Unwin.
tation solidly at th ntr of his study. Sltion dominats all volutionary
prosss'h insistd;it was th xplanation of first rsort. To sur, Hu-
ly idntifid a wid rang of oth iologial prosss(.g.,drift, isolation, volutionary ol
hyridization, and hanging mutation rats). frquntly gav thm ky volutionay ompu
rols in rtain onfind topis. Nvrthlss,sltionrmaind his most po- from iologial vo
tnt volutionary mhanism. For Huly' adapting to wid rang of sl- Taylor 1999). an
tion prssurs was lif's most frqunt pattrn. of possiilitis for
Huly onsidrd himslf a philosophr as muh as a iologist. Th final human programmr!
haptrs of uoltio onsidr volutionary trnds and progrss. uxly pr- Biologil voluti
sntd svral spifi (and ojtiv, h thought) ritria to masur progrss: diffiult prolms.
gratr ontrol ovr th nvironment' gratr indpendn from th nviron- normous numr
mnt, and gratr ffiiny. Th volutionary advantags of ths qualitis surviv and to rpro
smd ovious to Huxly. Natural sltion drov toward ths qualitis. Th origins of v
This onntion was ky. uly prsntd his approah as an improvmnt ( 1 9 5 0 ) s u g g s t i o no f
ovr his prdssors,who rquird suprnatural drivrs and guiding fors. volution.'' This os
His shm did not. Ovrall, humans srvd as Huly's pinnal of progrss, solutions to prol
with our intllign (onsisting of tru sph, onptual thought, and om- 1950s, svral grou
plx motions) apping volution's ours. Huly hld out for furthr approah.
progrss' too' suggstinghumans had a moral imprativ to ful that pross. In th lat 1960s,
uolutio nds with uly onsidring ways humans might at ithr to has- dpndntly introdu
tn or to str thir own volutionary futur. gnti algorithms (l
For svralgnrationsof iologists,uolutio providd an importnt his- and volutionary pr(
torial narrativ for volutionary studis.HuIy prsntdharls Darwin of suh approahs i
as a gnius who disovrd in natural sltion on of lif's ntrl prinipls. volutinary algoritl
This gnius was lipsd in th yars |880_1920, supposdly owing to Dar- Th simplst vrsi
win's failur to undrstand inhritan. Th ris of gntis solved th pro- odd solutions to a
lm. Its rnt growth, and th dvlopmnt of othr spialitis sin, vastly al vlu to ah in
improvd our undrstanding of partiula pis to vlution's puzzl. This th population to r(
narrativ mad Huly's rol sm natural. Th tim was rip, h said, for rossovr' and muta
somon to ollat th many sparatndavorsinto a omprhnsiv'syn- gnti algorithm it
thti whol ' uolutio was uxly's attmpt at that synthsis. uxly's ria is mt.
ovrall mssag was that Darwinism had rturnd in improvd form, a volutionary om
phonix rising from turn.of-th-ntury ashs. ntifi appliations,
uoltion dlivrd no rilliant nw insight. Instad, it providd a omp- root havior, mor
tnt survy of rnt dvlopmnts in volutionary studis. Within that r- suprsoni jt dsign
sarh ommunity' Huxly promotd natural sltion as an organizing An intrsting vri
thm. Latr gnrations followd th sam path and pointd to uolutil mig,was introdu
for its nthusiastisupport in that dirtion. viwd as a gnti
Huxly pulishd a sond dition of ullutio in 196. ollagus pu. truly utal ol
lishd a third dition, with a sond nw introdution, in \974.In ah, th dmonstratdin a v:
main haptrs rmaind ssntialIyunhangd. Rnt trnds in .
manuftur of phy

spanndoth urop and th
d natural sltionand adap.
o n d o m i n sa l l v o | u t i o n a r y
f first rsrt. To sur, Hux-
prosss(.g.,drift, isolation,
H frquntly gv thm ky
ltionrmaind his most po.
,ptingto a wid rang of sl-
muh as a iologist. Th final
:ndsand progrss.Huxly pr-
rt)ritria to masur progrss:
r d p n d n fr o m t h n v i r o n -
. advantagsof ths qualitis
drov toward ths qualitis.
s approah as an improvmnt
ral drivrs and guiding fors.
Huly'spinnal of progrss,
onptualthought' and om-
Huly hld out for furthr
imprativto ful that proSs.
.lumansmight at ithr to has-
io provided an important his-
dy prsntdharls Darwin
on of lif's ntral prinipls.
)20,supposdlyowing to Dar-
s of gntissolvd th pro-
f othr spialitissin, vastly
s to volution's puzzl. This
'h
tim was rip, h said, for
rs into a omprhnsiv,syn-
pt at that synthsis. Huly's
turnd in improvd form, a
. Instad,it providd a omp-
onary studis.Within that r-
a l s l t i o na s a n o r g a n i z i n g
)ath and pointd to uolutio
iio tn 196. ollagus pu-
ldution' in \974. In ah, th
L.
uolutior ompting d Gnti lgorithms 549
|BLIoGRAPY
uxly,J. [1942,96311974.uolutil:h oder Sthsis'3rd d. London:
Alln and Unwin. -I..
volutionary omputing and gntialgorithms
volutionary omputing modis a olltion of thniqus that us idas
from iologial volution to solv omputational prolms (lvIithll and
Taylor 1999). any suh prolms rquir sarhing through a hug spa
of pssiilitis for omplx solutions. Ths solutions ar usually diffiult for
human programmrs to dvis.
Biologial volution is an appaling sour of inspiration for addrssing
diffiult prolms. volution is, in fft, a mthod for sarhing among an
..solutions''
normous numr of possiilitis for that allow organisms to
surviv and to rprodu in thir nvironmnts.
Th origins of volutionary omputing an trad to Alan Turing's
(1950) suggstionof an ..ovious onntion twn mhin larning and
volution.'' This osrvation itd popl's imagination toward volving
solutions to prolms using iologial volution as a mtaphor. In th lat
1950s, svral groups indpndntly dmonstratd th possibilitis of this
approah.
In th lat 1960s, John olland, Ingo Rhnrg,and Lawrn Fogl in-
dpndntly introdud fundamntal volutionry ompuing pproahs:
gnti algorithms (Holland 1992), volution stratgis (Rhnrg 197),
and volutionary programming (Fogl 1998|, rsptivly. Th gnral trm
of suh approahs ts euolutionr lgorithms. Th most widly usd form of
volutionary algorithms ar gnti algorithms.
Th simplst vrsion of a gnti algorithm onsists of a population of n-
odd solutions to a givn prolm, a fitnss funtion that assigns a numri-
al valu to ah individual, and a st f gnti oprators t applid to
th population to rat a nw population. Ths typially inlud sltion,
rossovr' and mutation. Starting with a randomly gnratd population, a
gnti algorithm itrativly transforms th population until som stop rit-
ria is mt.
volutionary omputing is finding us in a varity of ommrial and si-
entifi appliations' inluding intgratd iruit dsign, fatory shduling,
root hvior' mophology dsign. omputr animtion. imag prossing.
suprsoni jt dsign, finanial markt prdition, and drug dsign.
An intrsting vaiant of volutionary omputing, trmd gti prgrm-
mig, was introdud y John R. Koza (1992). Gnti programming may
viwd as a gnti algorithm that volvs not just paramtrs' ut rathr
truly utal omputr programs. This mthod has n sussfully
dmonstratd in a varity of appliations, inluding optimal foraging.
Rnt trnds in volutionary omputing inlud th automatd dsign and
manufatur of physial roots. For ampl, Lipson and Pollak (2000)
.'50 uolutilnr rlgrss

hav omind simulation of havior nd n.rorphologyto automatially flt y many palonto

volv and onstrut atual roots. Furth work y Murat and Yamaguhi (Darwin 1859, 36.1).
(2005) mploys gnti algorithms to slf-onfiguroots that an assml In volutionary dis
and rpair thmslvsin simpl situations. many volutionists, u
Th parallls btwn iologial volution' sarhing through a spa of philosophially and s
gnotyps' nd omputr volurion' sarhing through a spa of omputr lutionary progrss har
pograms o othr data struturs,hav rystallizdto ratth sintifin- tndny to rad into
trpris of volutionary omputing. This fild has provn usful in a varity w think w s in hu
of ommrial optimization prolms, showing promis for many ompl two indpndnt orr
sintifiand nginringprolms. laim that lng_tmd
trnd has n for th
|LIoGRAPHY of a long-trm trnd is
Fgl, D. B. 1998' olutior otputtiltt:be Fossil Rord. Nw York: I rasing. But what, x
Prss. progrss in what sns
olland, J. H. 1992. dpttio i Nturl nd rtifiil Sstems: A Itrodutor disours has nald
lsis t't''ithpplitios to Bilog, otrrll d rtil Itlligece. Disssions of vol
amridg, A: IT Prss'
gists, palontologists
Koza, J. R. 1992. Ceti Progrmig: the Progrmmig lf omputrs b
s of Ntri Sltio. Cmridg, A: lT Prss. l y ( 1 8 9 3 ) ,H r r t S
Lipson, R., nd J. B. Plllk.2000. Automati dsign and manufturof rooti (1'961"), Ldyard Stb
lif forms. Ntur 406:974-978. son (1949),Stphn
ithll' ., and . . aylor 1999. volutionary omputtion: An ovrvlw. a|en (1973)' Grat
Aul Ruitu of olog d Sstmtis 30: 59-61'6. (|996}, and othrs.Ir
Muatir, T., nd . Ymilguhi. 2005. Nighoring rossovrto improv GA.sd
has n rlgatdto
Q-larning for multi.lggdroot ontrol. Ppr prsntdat.Washington,
th Prodingsof
th 2005 nfrn on Gnti volutionry omputation, DC, has rmaind ntral
ltne 25-29. vating fator hind
Rhnrg' I, 1973. t,r>ltilltsstri'1tgi:
optimirugechnishr Sstm lmd th twntith ntur)
Prizipi dr Biologish n ullutil. Stuttgart: Frommann-Holzoog. disussion of this valr
Turing, A. 1950. omputing mahinry and intlligne.id 59 43-460.
-..v. d..T.
PRoGRss IN

Thr ar two Ways iI

volutionry prgrss
vlu. Th taturo
odrn organisms sm learly mor advand than anint ons. ompar tionary pross' so to
an lphant with a modrn atrium. (Th omparison is apt in that rtain standpoint th first a]
modrn btria a praly vry simila to th first fossil orgnisms tural (ligious,soi
known, frm roks 3..5illion yars old') Th notion of an ordring among Indd, thy ar di
organisms dats at last to Aristotl, who arrangd living forms on a linar ality in volution w(
sal asd On dg of prftion, also alld the scl lturlor grat fatr w valu, su
hain of ing' For mor than 2'000 yars aftr Aristotl, th hain Was un- ours' w might t
drstood to stati' maning that organisms and thir rnkings did not rathr than trnds g
hang. But in th rly nintnthntury Jan-Baptistd Lamark addd vord in volution l
a volutionary omponnt' so that orgnisms rovd up th hain as thy trnd' and that w vz
volved. A half ntury latr th Darwinian viw hallngd th ordring it- hr too th valuat
self. Drwin sw volution aS a pross of ranhing and divrgnrthr as original motivati
thirn linar asnt. But Darwin nvrthlssrognizd progrss' witing fa- a trnd in fatursvi
mously in on th rigi of Spcies of ..that vagu yt ill-dfind sntimnt' Th sond altrn

d morphology to automatially
work y urata and Yamaguhi
'nfigurroots that an assml
'n' sarhing through a sp of
ng through a spa of omputr
tallizd to rat th sintifi en-
l d h a s p r o v n u s f u Ii n a v a r i t y
ving promis for many omplx
,he
FossilReord.Nw York: I
ArtificiISstms:A Itrodutor
-roId rtificil Itllig.
Progrmmigof omputrs b
\: MIT Prss.
signand manufaturof roboti
ry omputation:An ovrvrw.
30:593-616.
ing rossovrto improv GA-asd
aprprsntdat th Prodingsof
ry omputation,Washington,DC,
ierugThnisherSstmund
gart:Frommann-Holzoog.
llign.
id 59:4-460.
-E'.,.v.d.'T.
ld than anint ons. ompar
omparison is apt in that rtain
ar to th fist fossil orgnisms
.h
notionof n ordring among
trrangd living forms on a linar
:alld th sl tLrr or gteat
rftr Arisrotl, th hain was un-
isms and thir rankings did not
Jan-Baptist d Lamark addd
sms movd up th hain as thy
i viw hallngd th ordring it-
ranhing and divrgn rathr
; rognizd progrss' writing fa-
.t vagu yt ill-dfind sntimnt'
uoltionr Progrss 551
flty many palontologists'that organisation on th whol has progressd''
( D a r w i n1 8 5 9 , 3 6 3 ) .
In volutionary disours sin Darwin' progrss has n aknowldgd y
many volutionists, ut th ida hs also om to sn as troulsom, oth
philosophially and sintifially.It is widly rognizd that intuitions on vo-
.W
lutionary progrss hav n ontaminatd y ultural influns. hav a
tndnyto rad into th history of lif th soial and thnologial progrss
w think we s in human affairs. Also' progrss is now rognizd as having
rw indpendnt omponnts that ar frquntly onflatd, a valu-nutral
laim that long-trm dirtional hang has ourrd and a valu laim that th
trndhas n for th ttr.Among ontmporary volutionists th istn
of a long-trm trnd is widly aptd.Most would agr that somthing is in-
rasing.But what, atly? And inrasing in what sns?Is it progrss? If so,
progrssin what sns? Ths issus hv not n rslvd, ut th modrn
disourshas nald us at least to formulat thm fairly larly.
Disussions of volutionary progrss an found in th works of iolo-
gists,palontolgists, philosophrs' and historians, inluding Thomas Hu-
ly (1893)' Hrrt Spnr (1857), Julian Huxly (1942)' Thoms Goudg
(1961)'Ldyard Stins (1,969),rnst ayr (1988), Gorg Gaylord Simp-
son (1949), Stphn Jay Gould (1988), Franiso a\a (\974), Ligh Van
a|en (1973)' Grat Vrmij (1987), Rort Rihards (1'992),ihal Rus
(1'996|,and othrs. Intrstingly' among th sintists most of the disussion
has n rlgatd to popular works. Rus argus that th notion of progrss
has rmaind ntral in volutionary studis, oftn its proof th hif moti-
vating fator hind th work that popl do, ut that in th sond half of
th twntith ntury th inrasing profssionalization of th fild ford
disussion of this valu.ladn suit out of th thnial litratur.
PRoGRss lN WHA sNs?
Thr ar two ways in whih a trnd in som fatur of organisms might hav
valu.Th fatur ould valal to us, or it ould valud y th volu-
tionary pross' so to spak, in th sns of ing psrvd. From a sintifi
standpointth first altrnativis problmati. In sinour prsonal or ul-
tural (rligious,soial) valus shold not rlvant or at last not ntral.
Indd,thy ar a distration. For xampl, if th atual dfining dirtion-
ality in evolution wr inrasing nrgy usag' an infatuation with som
faturw valu, suh as intllign' might aus us to ovrlook it. of
ours' w might intrstd in th volutionay sours of our valus
rathrthan trnds gnrally. For xampl, it might that intllignis fa-
vord in volution by natural sltion, that w ar th produt of that
trnd,and that w valu intllignon aount of its advantagsfor us. But
hr too th valuativ omponnt of our intrst is sid th point, pt
as original motivation. Thus it is hard to s how progrss' if it is dfind as
a trnd in fatursvalual to us, ould a sintifially usful notion.
Th sond altrnativ is to intrprt valual to man that whih is pr.
552 uolutior rogress

srvd' or' aus W ar intrstd in trnds, that whih inrases ovr

tim. Progrss thn oms purly sientifi trm, ut now douts aris
aout th aptness f th word. It might e apt if th volutionary pross pr. )
srvs faturs that strik us as positiv, lik intllign, fitnss, or ompl-
ity. But suppos w disovr that th main long-trm trnd is an inras in a
fatur aot whih w ar amivalnt' lik nrgy usag' or a fatur that
sms downright ad, lik fragility (lading to an inrasing proaility of
tintion). It would sound odd to all ithr of ths progrss.
I think that ths prolms are fatal, and w should aandon talk of volu-
advanem
tionary progrss and instad dvot our nrgisto th study of trnds. But th
notion of progrss has grat ultural momntum and sms likly to prsist' at
last outsid th thnial litratur, for som tim to om. In what follows, I
ad to th follis of my tim and us th word pro?rss, ut only in th s-
ond sns,to man that whih is prsrvd.For onvnin,I will also us th
phras degr of duement, aS though th varials involvd in progrss
wer ithr known or masural. (As will sn' at prsnt th ar not.)

A RND IN WHA sNs?

Disussion of progrss has n muddld, vn rntly' y onfusion about

(A) A hypothtiallong
how trnds in groups work. In urrnt trnd thory it is not nough to say
gins with a singl sp
that som varial inrass.To lar' on must spify whih of sval
vertial lak lin sgm
group statistis is inrsing-man' mximum, or minimum-aus aris,and som spis
trnds in th diffrnt statistis hav diffrnt intrprtations. A trnd in th evry nw spis.Hr
mean is an inras in th avrag dgr of advanmnt among all spis strongly drivn. (B) Sa
in xistn at a givn tim. In th figur, A shows a hypothtial long-trm th maximum and mini
trnd. Lif gins with a singl spis' a singl linag' with sm low lvl prdominate'ut a nu
of advanmnt.As tim passs,nw splsris, and sm spisom whih inrassand d
(th vrtil dottd lin
xtint' with hange ourring in the origin of vry new spis. Th man
disussion.
at an givn tim sli is th avrag lvl of advanmnt of all spis in x-
istn at that tim. A trnd is an inras in that avrag' that is, movmnt
of th avrag to th right. A laim that a trnd has ourd, without any Disussion of progr
qualifir, usually mans an inrasin th man. trnd' prhaps aus
A trnd in th maimum is a ris in th dgr of advanmnt ahivd by masur it). Rathr, it
th most advand spis at a givn tim, in othr words, a ris in th highst mhanism. In figur ,
lvl of advanmnt ahivd. (In th figur, B is th sam as A, ut annotatd that all hangs ar i
to show th trnd in th maimum' th dottd lin on th right.) aima ar ras. Figur is th
of spial intrst aus humans ar gnrally thought to rprsntth most maning that sltion
progrssiv spis in xistn and, if so, would rprsnt th mimum tn a th volution of rtair
.W.
tim slie that inludd th prsnt. would th last spison th right at advand spis prs
th vry top of th graph. Finally, a trnd in th minimum is an inras in th minimum stays rough
dgr of advnmnt of th last advand spis (th dottd lin on th lft driu, or mor pris
in figur B). inima are of spial intrst ausapparntly th lowst lvl Figur D is diffrnt in
of advanmnt has not hangd. Batri hav istd, and indd hav dom- Inrass and das
inatd th iosph, throughout th history of lif. (In othr words, it is lar symmtrially' oth u1
that figurs A and B ar not a good rprsntationof th history of lif.) limit on dgr of adv

in trnds, that whih inrss ovr
sintifi trm, ut now doubts aris
; b apt if th volutionary pross pr-
, lik itllign,fitnss, or omplx-
n a i n l o g - t mt n d i s a n i n
;rt' lik nrgy usag' or a fatur that
ading to an inrasing proability of
l ithrof thsProgrss.
, and w should aandon talk of volu-
nrgisto th study of trnds. But th
lomntumnd smslikly to prsist, at
'r som tim to om. In what follows, I
th word progrss, ut only in th s-
vd.Fo onvnin,I will also us th
lugh th varials involvd in progrss
will b sn' at prSntthy ar not.)
dld, vn rntly' by onfusion out
nt trnd thory it is not nough to say
lar, o must spify whih of svral
n. maximttm. r minimum-aus
diffrnt intrprtations. A trnd in th
:grof advanmnt among all spis
gur, A shows a hypothtial long-term
s,a singl linag,with som low lvl
'spisaris' and som spis om
: origin of vry nw spis.Th mn
vl of:rdvnnrntof all spis in -
ras in that avrag,that is, movmnt
that a trnd has orrd' without any
n th man'
l th dgrof advanmnt ahivd y
im,irr othr words, a ris in th highst
: figur,B is th sam as A, ut annotatd
h dottd lin on th right.) Maima ar
gnrallythought to rpsnt th most
f s o , w o u l d p r s ntth m i m u m i n a
would b th last spison th right at
.ndin th minimum is an inras in th
vandspis(th dottd lin on th lft
:rstausapparntly th lowst lvl
|triahav istd, and indd hav dom-
histry of lif. (In othr words, it is lar
)prsntationof th history of lif.)
uolutionr Progrss
(l)
Strongly Weakly
a s i n
driven driven
adVanement
? ma
B
(A) A hypothtiallong-trmtrend in advanmnt'that is, prgrss.Lif
bginswith a singlspis,with som lw lvl o{ advanmni(thsigl
vrtialblak lin sgmntat th ottom lft).As tim passs,nw spis
arise,and som spisomtint,with hangouringin th origin f
vrynW spis.Hr all hngsar inrss,so th trnd is said to
stronglydrivn. (B) Sam as A, with dotted lines addd to show th trndsin
th maximum and minimum. () A wkl drivn trnd in whih inrass
pdominat,ut a numr of dasslso our. (D) A passivtrnd in
whih inrssnd drassar qually frqunr,ut lowr limit on hng
(thvrtialdottd lin) forslineagsto diffusto th ight.Stet for
d i s us s i o n .
Disussion of progrss has not n onrnd with th istn of a
trnd, prhaps aus:1trnd is takn for grantd (dspitou inability to
masurit). Rathr, it has n aout th pattrn of hang, th undrlying
mhanism. In figur , natural seltion strongly favors dvanemnr' so
that all hangs ar inrass.Th man, maimum, and minimum all in-
ras' Figur is th sam pt that th nt sltiv advantag is wak,
mning that sltion Somtims favors drass.(h lassi ampls ar
th volution of rtain rdud forms, lik parsits.)Th sult is that lss
advand spis prsist or ar rplaed y drassfrom aov' and th
minimum stays roughly th sam. h trnds in A and ar sid to
driun, or mor prisly,A is strngly divn, whras is wakly drivn.
Figur D is diffrnt in that thr is no ovrall sltiv advantag to inras.
Inrassand drassour qually oftn' so th only ndnyis t sprad
symmtrially, oth up and down. Howevr' thr is a oundary, a lowr
limit n dgr of advanmnt,that loks th sprad of th group on th
5 54 uolutionr rlgress

lft. This oundary an thought of s th lowst lvl of advanmnton- shold mo fit thl
..in tJ
sistnt with bing liv. Th rsult is that tir mn and maximum inras Darwin's trrns,
whil th minimurn stays th sam (as in th wkly drivn as).Th trnd is also knw, natul sl
said to e pssiu. on gologial timsal
Th urrnt ontrovrsy ovr mhanism gan in 1996 with ook on du just onstant h;
progrss y StphnJay Gould in whih h raisd th passiv mhanism to nss. On th othr
th lvl of plausiility. At issu is th rol of sltion in progrss, that is, adaptations, suiting tl
whth th (prsumd)trnd in th history of lif is wakly divn y sl- vord ovr thos mor
tion or mrly passiv.(A strong|ydivn mhanism is ruld out austh offer advantags on ti
tru minimum is thought to hav rmaind stal: atria prsist.) Gould Thr ar som iolo
h:rs bn widly nrisintrptd as laiming that volution is random, ut th suh as rtain llr"rl
irpliation of th passiv mhanisris not randomnss.Within ah volv- pathwys. But w do
ing linag sltion dos oprat, and organisms ar adapting. Rathr, th rigorous way, or how
implition is that volving linagsas a group r govrnd y many diffr- that is lar now is thz
nt sltiv fors, omplly onfigurd' so that sltion favors inrass ar possibl in prinip
and drassqually oftn. Th pross an undrstood s purly dtr- Th most widly r
ministi, lik th diffusion of a gs. As a possil asis ior
Unfotunatly,as will sn,thr ar hardly ny data availal,so from onntion with progr
a sintifi prsptiv th dat aout mhnism is taking plir in a nar usd in volutionay st
vauum. Still, thory has produd som insights.Fo ampl, it is now lar for progrss, suprfii
that in th dbat ovr mhanism two ommon osrvations ar irrelvant. sutly onnoting advar
Th first is th prsistnof nonprogrssiv spis,th stal minimrrm. This nition maks this usag
osrvation is not hlpful aus tlr minimum is prditd to rmain un- on an say anythinga
hangd in oth ass, passiv and wakly drivrr.Likwis, th is in th tain a thnial dfinit
maximum and th urrnt istn lf som highly advand spis,lik hu- srssof omplity. l
n-tans,ar not hlpful. A rising mimum is ptd in all ass. typs, numr of physi
dvlopmnt (sfigur
don, and with on -
A RND IN WA?
n dmntd.Nor
any andid:rtsfor a long-trnrtrnd hav n proposd: (1) fitnss;(2) in hang in ths vri
omplity; (3) aility to sns' ontrol' or rspond to th nvironmnt; (4) som vidn within
ody siz; (5) intllign;(6) vrsatility or volvability' maning th aility hs bn a trnd in nur
ttl hang in voltion; (7) th rvrs of volvaility, lso alld dgr of Th xption is <
ntrnhmnt;(8) nrgy intnsivnsso rat ofnrgy usag;and (9) dpth nstd, to som t
and sophistiationof mhanismsof inhritan'For th most pat advoats within whols' ovr
of all ths andidats hav takn a as-uilding pproah, with th rsult rs of lvls hav v
tlrat th litratur offrs a numr of lvr and somtims-powrfulargu- atria. Aout 2 bili
mnts in favo of ah. Unfortntly, som of th most intrsting-lik ioti rlationship to
fitnss, ompliry, nrgy intnsivnss, and aility to ontrol th protozoans suh as l
nvironmnt-ar diffiult to oprationaliz. Thus thr r hardl any lls, ut us th
data on th sal of lif as a whol, for any trrrd statisti, for ny of ths nxt lvl up. Thn
andidat varials. joind to produ th
For som of ths variabls, many wuld want to ount a long.trm trnd, ga), th anstor of
if on ould doumntd, as progrss.on varil is fitnss,ndrstood lias to muskrats. Fin
as aility to srviv and rprodu. [t might sm that Darwin,s prinipl of individuals joind to
natural sltion virtually guarants a trnd in fitnss. Latr organisms alld a ryozoirn. V

,Wstlvl of advanmnt on-
i m a na n d m a i m u r n i n r
,aklydrivn as).Th trnd is
gan in 1996 wit. a ook on
isdth pssiv mhnism to
lf sltion in progrss, that is,
lf lif is wakly drivn y sl-
:hanismis ruld out arrsth
stal:atria prsist.) Gould
lat volutin is random, but th
.s7ithin
andomnss. ah volv-
risms ar adapting. Rathr, th
lp ar govrnd y many diffr-
l that sltion favors inrass
undrstood as purly dtr-
rdly any data availal, so from
ranism is taking pla in a nar
hts. For ampl, it is now lar
non osrvations ar irrlvant.
is'th stal minimum. This
l u m i s p d i t dt o r m i n u n -
lrivn. Likwis, th ris in th
righly advand spis' lik hu-
ptdin all ass.
: bn proposd: (1) fitnss; (2)
:spondto th nvironmnt; (4)
:volvaility, maning th aility
olvaility, also alld dgr of
of nrgyusag;and (9) dpth
n.For th most part advoats
ilding approah, with th rsult
and somtims-powrfulargu-
r of th most intrsting-lik
and aility to ontrol th
z. Thus thr ar hardly any
trnd statisti,for any of ths
vant to ount a long-trm trnd,
r variabl is fitnss, undrstood
smthat Darwin's prinipl of
:nd in fitnss. Latr organisms
uolutior Progrss t-t i
should mor fit than arlir ons' on th whol, having atn thm, in
s Darwin's trms'..in th ra for lif'' (Darwin 1859,363). But as Darwin
also knw, ntural sltion produs adaptation to loal nvironmnts, and
on gologialtimsalsthshang dramatially.Thus sltionmight po-
du just onstant hang with no nt improvmnt' no nt inras in fit-
nss. On th othr hand, on long timsals organisms with gnral
daptations, suiting thm to a widr varity of nvironmnts' ould fa-
vord ovr thos mor narrowly adaptd. Ar thr gnral adaptations that
offr advantagson timsalsof hundrds of millions or illions of yars?
Thr ar som iologial mhanisms that may hav prsistd that long,
suh as rtain llular transport mhanisms and parts of rtain mtoli
pathways. But w do not know how to idntify gnral adaptations in any
rigorous way' or how to tst whthr thy hav bom mor pvalnt. All
that is lar now is that oth altrnativs, a long-trm trnd and its asene,
ar possil in prinipl, and thrforthis is an mpirial issu.
Th most widly rognizd andidt for a long-trm trnd is omplxity.
As a possibl asis for progrss it is rally a stalth andidat aus a dirt
onntion with progrss is rarly aknowldgd. But th way omplxity is
usd in volutionary studis suggststhat it funtions as a kind of od word
for progrss, suprfiially valu fr and thrfor sintifi sounding ut still
sutly onnoting advanmnt. Th asn of a widly known thnial dfi-
nition maks this usag problmati. Noody knows what omplity is, so
on an say anything at all about it. Th litratur of rnt dads dos on-
tain a thnial dfinition' or rathr svral dfinitions for ah of th svral
snssof omplxity. Ths inlud omplity in th sns of numr of part
typs,numbr of physiologial or havioral prosss'and numr of stps in
dvlopmnt (sfigur on pag 556). Howvr' littl mpirial work has n
don, and with on xption' no trnd that spans th history of lif has yt
bn doumentd. Nor do w know vry muh aout th mhanisms involvd
in hang in ths varials, passiv vrsus drivn. (On a smallr sal thr is
som vidn within th animals, ovr th past 500 million yars' tht thr
has bn a trnd in numr of part typs and that it has n passiv.)
Th ption is omplity in th sns of nstdnss.organisms ar
nstd, to som xtnt' lik hins os, with multipl lvls of parts
within whols. Ovr th history of lif, rganisms with vr gratr num'
rs of lvls hav volvd. Th first living things 3.5 billion yars ago wr
atria. Aout 2 illion yars ago som atria joind togthr in a sym-
bioti riationship to produ th first ukaryoti lls, similr to modrn
protozoans suh as mob' Thy w suprfiially similar to batrial
lls, ut aus thy volvd as olonis of atria, thy oupid th
nxt lvl up. Thn aout 600 million yars ago lons of ukaryoti lls
joind to produ th first multillular individual (proaly a kind of al-
ga),th anstor of all modrn multiIlulrs' from mushrooms to magno-
lias to muskrats. Finally' aout 480 million yars ago lons of multillular
individuals joind to form th first olonial animl' a oral-lik animal
alld a ryozoan. odrn olonial organisms inlud ryozoans, orals'
5.'6 uoltionr Progrss

Myr. . 1988.Tord,
u lluti li st.rn
M S h a 'D . W . 1 9 9 1 .
d hiloslph6, n
1 9 9 8 .P o s s i lla
hypothss... u
RihrdsR . . J . | 9 9 2 . h (
d ldologil R
hiago Prss.
R u s ' . | 9 9 6 . , , n u 7' ,
ambridg,A:
Simpson,G. G. 1949.Th
Prss.
S p n rH' . l 8 5 7 . P r o g r
Stins'G. L. |969, h
North arolina Prs
Van Valn,L. 1973.A n
Vrmij,G. J. 1987.u
A lassi ampl of why dfinitions of progressin trms of movs to Prss.
omplity ar prolmati.Th uppr skltonis of a lion and th lower
skltonis of a whale. By any measurmntth akonof th whale is th
simpler' and yt adaptivly the whl's akon is what is nddfor a martn
volutionary psy
mammal, just s th lion's akon is what is ndedfor a trrstrialprdator.
Whals dsndedfrom land ttrapods(four-lggdanimls)sO thir S o m h v i o r |s i
bkonsmark a sltion-drivn move from omplityto simpliity.(From study of human hav
MShea 199|. |5.\ In this road sns,v
q u i r y a r d f i n dn o t l
ut y th kinds of qu
fr from paradigms' W
th soial insts, and many vrtrats,from fish to humans. ! do not
ologis with whih th;
know if vn highr lvls-olonis of olonis-hav n rahd. Thr
a fild of inquiry, volu
ar no lar-t ass. Humans sem to ssoiat at many lvls' ut only
I i t i o no f h D s
wakly at th highr ons. Intrstingly,if humans hav not tuly rahd
i n d Aimls in
th olony-of-olonis lvl, nd if no othr organism has ithr, this raiss
t i u l t d p a a d i g m s
a novl possiility: th only varial in whih w an doumnt a trnd,
d i s t i n t t h o r i s o u t
omplxity in th sns of nstdnss,in fat dos show a trnd, ut that
human havior and m
trnd might hav ndd, prhps 480 million yars go. Prhaps th ag of
Th first of ths pa
progrss is ovr.
th 1970s. Th or id
BIBLIOGRAPHY n a t u r a Ia n d s u |s I t
many spis hoos m.
Ayala,F. J,1974. Th onptof iologialprogrss. In F. J. Ayala and
T. Dobzhansky,ds',Studisi th Philostlphof Biolog, 39_354.London: ml ourtship display
\4amillan. sltion will tailor m
Goudg,T . ^. 1961. Th Asent of Lif. Toronto:Univrsityof Toronto Prss. w a . s o i o i o I o g i s ras1
Gould' S. J. 1988.On rpiaingth idaof progrsswith an oprafionlnotionof rsn in a populati
dirtionality.In . H. Nitki, ed.,uolutilr
Progrss,319-38.hiago: thory. Aordingiy' hr
Univrsityof hiago Prss.
nations of how huma
1996. Full Hose: Th Sprdlf IlefrlmPIto to Dri'
Nw York: Hrmony Books. anothr-to plain hr
Huxly'I. S.1942. uoltio:Th odrn Snthesis. London:Alln nd Unwin. h i s , f o r m a n i f s t i n gl
uiy,T. . 1893.uolutilnd tbis.London:amillan. th ntrl thorti

in trmsof movs to
is of a lion and th low
akbonof th whl is the
i s w h t i s n d dt o
reddfor a trrstrialprdato.
4gdanimals)so thir
omplitvto simpliity.(From
lm fish to humans. !7 do not
ris-hav n rahd. Thr
iat at many lvls, ut only
tumans hav not truly rahd
o r g a n i s mh a s i t h r ,t h i s r a i s s
ih w an doumnt a trnd,
t dos show a trnd, ut that
1 yars ago. Prhaps th ag of
:ss.In F. J. Ayala and
, of Biollg, 39-54. London:
Univrsityof Toronto Prss.
ss with n opfational notion of
r Progrss,319_338. Chiago:
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n;amillan'
uolutior Psholog 557
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.Wht
Sh, D. !. 199 [. omplity and volution: vryody knows. illog
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-he
Rihards, R. J. 1992. etlllig of ullutio: Th lrpholtlgi! ostrutio
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Prss. -D.W.Nr.
m a r i n volutionary psyhology
Som havioral sintists dfine volutionary psyhology as simply th
study of human havior and psyhology from an volutionary prsptiv.
In this oad sns,volutionary psyhology is fild of inquiry. Filds of in-
quiry ar dfind not y speifi thoris aout th phnomna thy study,
but y th kinds of qustionsthy pos aout thm. Filds of inquiry thus dif-
fr fronr paradigms, whih r dfind y th spifi thoris and nrthod-
ologis with whih thy answr th qustions that dfin a fild of inquiry. As
a fild of inquiry, volutionaypsyhology gan with Charls Darwin's pu-
liation of Tbe Desct of in ] 871 and Tb pressilll rlf the motions
in d ils tn 7872.It was not until a ntury li1tthat larly ar-
tiulatd pradigms gan to emrg in volutionay psyhology, ah with
distint thoris about' and spifi mthods for studying, th volution.of
humn hvior urndmntillity.
Th first of thes paradigms was human soioiology, whih mgd in
th 1970s. Th or ida of soioiology was that havior has volvd undr
natural and sul sltionIusts org:rniform has. For ampl, fmalsof
many spishoos mats sdon th quality of mal or-rtship displays. If
mal ourtship displays vary in quality, and that variation is hritabl,sual
sltion will tailor mal otltship havior to fmal prfrn.In this
way, soioiologistsargud' a form of havior an an ildaptation' whos
prsnin a population is to plaind y th prinipls f volutionary
thory. Ardingly, humn soioiology sought to offr volutionary pla-
nations of how humns ar havioraily adptd to soial lif with on
anothr-to xplzrin human havioral adapttions for dominan hirar.
h i s , f o r m n i s t i n ga r t d d a l i n g w i t h a g g r s s i o n , n d o m a t i n g . l n d d ,
th ntrirl thortial prolr-rof humirn soioiology Was to plain th
5.'8 uolutitr Psholog

volution of altruism, th prforman of ats tht nfitothrs at a fitnss psyhologial adaptat

ost to th tor. thn worn.
In th fild of volutionarypsyhology today, lrumn soioiologyhas n Th othr notal ;
suprsddy svralaltrnativparadigms' two of whih ar partiularly no- m a n h v i o r a | o | o
tal. On of ths onfusingly gos y th nam ..vllutionrypsyhology.'' h v i o r i s a d p t i v l yr
o avoid onfusion, and to distinguislrth fild f in1iryfrom th pradigm, nvironmnts. Th fur
I will rfr to this par:rdigmas ..volutirnaryPs,vhllrlgy'' (apitlizd). tion has dsigndarri
volutionary Psyhology is a marig of ognitiv psyhology with vo- tsks tat ar ssnt
lutionary iology. In ontrst with soioiology, volutionarv Psyholo- food, apturing pry'
gists argu tht, whn z-rhavior hs volvd undr sltion, it is not th havioral ologistsvi'
havio tht is n ad:rpttion, ut rathr th psyh<lltrgialrrrlranism in spons to nvirn
that auss that havior. Th goirl of volutionary Psyhology is thus to of suvival and rprod
disovr nd dsri th information-prossing strutur of our psyho- tions aout animal b
logial adaptations. Baus adaptation is a slw pross' volutinary t h o r t i m o d l s .T h .
Psyhologists liv th.tt ou psyhlllgialadaptiltions vlvd dr"rringth survivl or rprodut
l i s t o n ( 1 . 8 m i l l i o n t o 1 0 , 0 0 0 , v a r s g o ) t r l s o l v t h p l I m so f s u - m a l s a r l p u r s u i n p l
vival and rprodution fad y our hLlntr-gathrranstors' orlvr, to ah of th stratgi
volutionary Psyhologists argu that a distint psyhologial mhanism b h a v i o r ls t t g yo P
volvd for ah distint prolm fad y our httntr-gathrrnstors' mals thy ar studying
Thy liv that th hrtnrilnmind ontains hundr{sor thor.rsrrds of ths imls will pusu thl
mhanisms, olltivly onstitutillg a univsal human n2ltur. Baus osvd havior.
th nvironmnts inhaitd y humns hav hangd dramatially and rap- ths thniqus to th(
idly sin th Pliston, howvr, volutionary Psyhologists argu that o i l l u s t r t , o n s i
our ..Ston Ag'' psyhologial mh:rnismsoftn produ maldaptiv - has mor than on hr'
havio in th modrn nvironmnts for whilr thy ar not dsignd. Thus, t i si n t h t h n g a p
volutionary Psyhologists laim, in ordr to disovr th volvd dsign of tiv fo mals rn lig|
th mind' w must ..rvrsnginr'' th mind from th vantag of our Psyhology's laim ab
volutionary past' figurirlg out th prolms our llstors fd and thn But humn havior
hypothsizing th psylrologial mhirnisms that v<llvdto solv thm' p o l y a n d r o t l sm a r r i e
volutionary Psyhologists thn ondut stanr]ard psyhologial pri- polyndry to th o-hu
mnts to dtrmin whthr popl hiit th avior or prfrnsthat offspring to whih th
suh mhanisms would produ. ours rnong brothr
To illustrt, onsidr :r fundmntl asymmtrv btwn th ss. A dividd into parls th
Woman's liftim rpodutiv otltpltt is limit]lry th nurr of prgnan- drous marriags oul
is sh n arry to trm' and hr maimum rprodutiv olltput an r o t h s l u l t i v a r i n gt I
ahivd with a singl mat. In ontrast, a man's liftim rprodutiv outpllt Und ths oIog
is limitd only y th nrnr of ggs lr an frtiliz' and thus his maimum polyandrous marriag
rprodutiv oLltputan hivd only with multipl mats.Amlng ans- r i , a t h rt h n m
trl hunrans, thn' having multipl ats Would av ntaild signifiantr- onditions for whih
produtiv nfitsfor mals' ut not for fma|s.volutionary Psyhologists rothshivgrat
argu that this would hav ratd sltion prssur for mn to volv a of th farm' working
gratr dsir for sxual vrity than rvomn, making lln mor il-rtrstd in joint offspring than th
polygamous mating tlran wotntr'voluriona1'Psvologistshilv tstdftlr unran havioralt
this s diffrnin dsir,and th1,hav found tlrat, on vrag'mn want ky rspts.First, w|
narly fiv tims mor sual partnrs ovr th ours of a liftim than womn. h o l o g i la d p t a t i o n
vlutionary Psyhologists onlud tlrat ml dsir for sxu:rl r,arity is a ogy studis how hum
 uolutionr Psholog 5.'9
ats that nfit othrs at a fitnss psyhologial adaptation, whih motivats mn to
sek mor s partnrs
than womn.
ody,human soioiology has n Th othr notal paradigm that grw out of human soioiology
is hu-
ts,two of whih ar partiularly no- man havioral ology. Bhavioral ology is th study
of how animal -
..volutionarypsyhology.'' havior is adaptivly rsponsiv to onditions in animals; physial
nam and soial
fild lf inquiry from th paradigm, nvironmnts. Th fundamntal prmis of havioral
ology is that sl-
ary Psyhology''(apitlizd). tion has dsignd animals to maximally ffiint in prform"ing
a varity of
"foraging
of ognitiv psyhology with vo- tasks that ar ssntial to survival and rprodution,
suh as for
oiology, volutionary Psyholo- food, apturing pry, luding prdators, or wooing
mats. Aordingly, -
olvd under sltion, it is not th havioral ologistsviw animals as apal of flily
altring thir havior
hr th psyholgial mhnism in rspons to nvironmntal onditions in ordr to maximiz
thir hans
ivolutionary Psyhology is thus to of survival and rprodutiv suss.Bhavioral ologists
tst ths ssump-
rossingstrutur of our psyho- tions aout animal havior with optimality modls
and volutiory gam
r is a siow pross' volutionary thortimodels. Ths modls spify a partiular tsk
that is ssntialto
al adaptationsvolvd during th survival or rprodution, postulat a rang of havioral
stratgis that ani-
ago) to sllvth prolms of sur- mals an pursu in prforming that task, and assign fitnss
ostJand bnefits
ltr-gathrranstors. orovr, to ah of th stratgis.This nals havioral ologists
to alulat whih
distint psyhologial mhanism bhavioral stratgy optimizs th vrag ratio of nfits
to osts to th ni.
y our huntr-gthrranstors. mals thy ar studying. Bhavioral ologists thn prdit
that th studid an-
ins hundrds or thousands of ths imals will pursu that optimal stratgy' tsting thir
prdition gainst
univrsal human ntur. Beaus osrvd havior. uman havioral ology is simply
th appliation of
av hngddramatially and rap- ths thniqus to th study of human havior.
lutionary Psyholgists rgu that To illustrat, onsidr polyandry, a marital systm in
whih on woman
;ms oftn produ maladaptiv - has mor than on husand and whih is pratid in
four of th 849 soi-
vhih thy ar not dsignd. Thus' tis in th thnographi rod. At first glan, polyandry
appars maladap-
r to disvrth volvd dsign of tiv for mals in light of th volutio.'a.y logi.
volutionaiy
:h mind from th vantag of our Psyhology,s laim aout an volvd ,.* diff....,. "".,tying
in dsir for polygamy.
lms our anstors fad nd thn But human havioral ologists hav disovrd that
th o-husands in
-rismsthat volvd to solv thm. polyandrous marriags ar typilly rothrs. This hlps
to offst th osts of
t standard psyhologial pi- polyandry to th o-husands,austhir rsours
ar poold to rar only
it th havior or prfrns that offspring to whih thy ar all gntially rlatd. olyandy
also typially
ours among rothrs who hav inhritd farmland that
is too small to b
asymmtry twn th ss. A dividd into parls that ould ah sustain a family. Finally,
whre polyan-
limitd y th numr of prgnan- drous marriags our, altrnativ sours of inom ar
not availabl to th
<imumrprodutive output an b rothrs; ultivating th family farm is th only vial
mans of subsistn.
l m a n ' sl i f t i m r p r l d u t i v o u t p u t Undr thes ologial onditions, human havioral
ologists argu'
: n f r t i I i z 'n d t h u s h i s m a x i m u m polyandrous marriag is atually an adaptiv hoi
for th .oth.. who n-
with multipl mats.Among ans- tr it' rathr than a maladaptiv y-produt of volvd
dsirs nountrlng
l would hav ntaild signifiant r- onditions for whih thy wr not dsignd. Undr
ths onditions' th
fmals.volutionary Psyhologists brothrs ahiv greatr rprodutiv suss y mintaining joint
possssion
;tion prssur for mn to volv a of th farm, working it togthr, marrying on woman,
and raring thir
nn,making mn mor intrstd in joint offspring than thy would y marrying
monogamously.
ionary Psyhologistshv tstd for Human havioral eology diffrs from volutionary Psyhology
in svral
found that, on avrag' mn want ky rspts. First, whras volutionary Psyhology
strives ,o di,.uu.. p,y-
th oursof a liftim than womn. hologial adaptations to Plistonnvironmnts,human
haviorai ol'
m r l d s i o r s u a l v a i t y i s a ogy studis how human bhavior is adaptiv to nvironmntal
onditions.
 ,,uolutio
560 i deli Popultils''

Sond, volutionary Psyholgy livsthat hrn havior is oftn mal.

daptiv in ologial onditions that do not smlthos of our hunter-
gathrr anstors) ut humn havioral ology livs humans hav
adaptivly in a wid rang f nvironmnts.Third, whil volutionary Psy-
hology tris to rvl.snginrth volvd natr of th hunran mind from
th vantag of our Plistonpast' human havitlralology applis volu-
tionary prinipls to osrvalhuman bhavior.Finally, whras volution.
ary Psyhology strivs to disovr a uniform and univrsl human natur'
human bhavioal ology strivs to dtermin how nvironmntal differ-
ns twn individuals afft havioral diffrns twn thm.

ILloGRAPHY
Barrtt'L.' R. Dunr, and J. Lytt.2002. Hum uoltionrPsholog.
Printon'NJ: PrintonUnivrsityPrss.
Bullr, D. J. 2005. Adptig ids: uolutiorPschologd th Prsistt
Qustior tlltNtur.mridg,N{A: IT Pss.
-h
Buss' D. |\^.2007. uolutior,Psholog: Nu.lSiof th id.3rd d.
Boston:Allyn and Bon.
Laland, K. N., and c. R. Brown. 2002.Ssd Noss:uolutionr
Prsptius lIItBhuiour.oxford: oxford UnivsityPrss.
.!ilson, Adaptiv landsps.Tl
. o. \978. um Ntre. amridg,]vlA:Harvard UnivrsityPrss.
-D.I.B. th tops of hills f adap
volutionary mt;rphor<
pssiblthat Wiglrta
..volution in N1ndlianPopulations'' (Swall Wright) aldan, ..A thmat
tstablPopuIation
Th lassi papr ..volution in ndlin Popultions'' y Amrian popu- 28 (191),17-,|,42'w|
fitnss,drawn y tlr g
l a t i o n g n t i i s tS w a l l W r i g t w a s p u l i s h d t n G e t i si n 1 9 3 1 . I n i t , a n d
Provin points out that t
in ..Th Rols of Mutation, Inrding,rossrdingand Sltionin vo- information aout th
lution,'' a shortr papr givn the following ya at an intrntional gntis ultimatly,th landsap
..daptivland-
ongss'in whih h introdud his famous mtaphor of an
..shifting
sap'' (s figur), \/rightprsntdis alan thory of volu-
tion.'' His fllow population gntiist aross th Atlanti' R. A. Fishr
(1930), hzrd just givn his no-Darwinin aount of volutionary hang, would working agai
..gntidrift''' lso at
on that saw suh hang oling s th rsult of trlutatinsinto a larg pop-
ulation, and sltiln thn liminating all ut th st. Wright had workd that rnathmatis sho
for a numr of yars at th U.S. Dpartmnt of Agriultur. From studisof may wll ountr th r
th rding of attl, h was orrvind that th st way to gt hang is by without (or vn dsp
isolating small goups, working within thm to gt th st of all possil slvs of grat valu
haratristis, and thn finally rintroduing th highst quality to th whol lutionary hang.
hrd (Provine 1986). his is a vry non-D
.Wright's undr th influn of
shiftirrg alan thory followd th sam pattrn: isolation of
groups' improvmnt within th group, and thn mingling with th whol. In natural sltion. (Do
his ongrsspaper, !(right rprsntdth prossas somthingourring on Species, first pulishd
a landsap,with th st (th fittst)ing at th tops of pks nd th worst tins gv mo rdn
down in th vallys. H saw volution as ing a mttr of moving from on was doing on hromt
pak to nothr highr pk. Th hallng was how to gt down th sid of 1999.) In part th infl
th first pak so that on ould start liming up th othr. Bussltion omplaind that natur
 ,,uolution
i ndeli Pooltios'' 561

human havior is oftn mal-

'smlthos of our hunfr_
logy blivshumans hav
hird, whil volutionary Psy-
tur of th human mind from
avioral ology applis volu-
'r. Finally, whras volution.
and univrsal human natur'
t how nvironmntal diffr_
.rnstwnthm.

uoltiorPsholog.

shologd tb Persistent
[T Prss.
of tbind.3rd d.
Silt,e

(oss: uolutior
lford UnivrsityPrss.
'A:rvaro
U"'*.YJT.;
;'' (SwllWright)
rulatiorrs''y Amrian popu-
in Getisin 1931. In it, and
rdingand Sltion in vo-
r at an intnationag l ntis
rtaphorof an ..adaptiv land-
[ting alan thory of volu-
ss th Atlanti, R. A. Fishr
:ount of volutionary hang,
of mutations into a larg pop-
th st. \7right had wokd
lf Agriultu.From studies of
r bst wy to gt hng is y without (or vn dspit) natllral sltioIr.Tlrs nw fatus may thm-
to gt th st of all possil
h highst qulity to th whol
:h sam pattrn: isolation of
n mingling with th whol. In
jssas solthing ouing on Speies,first pulished irl 197, gav strong mphasis to \/right.Latr di-
,htops of paks and th worst
] a mattr of moving from on
ls how to gt down th sid of
up th othr. Baus sltion omplaind that natural sltionnnot yild innovations. Br.rta f ar greater
daptiv landsaps.This pitur, showing how th fittr organisms lim to
thtops of hills of adptivfitnss'illustratsprhapsth most famous
volutionarymtphorof th twntithntuy(from lfright 1932).It rs
possilthat Wrigt am to tlr mtaphorfrm rdingan artily B. S.
J.
Hldn,..A athem:rtialThory lf Artifiial and Ntural Sltion'Prt VIII.
tstalPopulations,'' Proedigsof th mbridg Philosophil Soit
28 (I9|' \7_142, whih ontainsa pitorial reprsntation of paksof
fitnss,drawn y th gntiist. H. !7addington. Wright,s iograph.!7illiam
Provinpoints out tht th landsapis odnsinga hug amount of
infomationaout te fftsof diffrntgnsnd that onsquntlv,
ultirnatly,
th landspmy not ntirlolrrnt.SProvin l98o.
would working against this' !right introdud th ontrovrsial notion of
..gnti
dri{t,'' also at tims known as th ..Swall Wright fft.'' H argud
that mathmatisshows that, in small groups, th han fftsof rding
may wll ountr th ffts of sltion, so nw faturs an l.l produd
slvs of grat valu, spiallyin th whol, and an ld to frth vo.
lutionary hang.
This is a vry non-Darwinin thory, although latr vrsions, partiularly
undr th influn of Thodosius Dozhansky' gav a muh largr rol to
natural sltion. (Dozhansky's mastrwork, Getics d tb rigi of
tions gave mor rdnto natural sltion,thanks tl studis Dozhansky
was doing on lrromosomal variations of fuit flis in ntur. S Rus
1999.)|n part th influn on Wright m from nri Brgson, who had
562 uolutio of Lguge

influn was Flrrt Spnr (partiularly as passd on y !right's tahr' opn-ndd varity oi r
L. J. Hendrson). Populations r sn in a kind of balan, homognous; r o f m a n i n g , l spsh o
when somtlrigdisrupts th balirn,ausing frgmntationand htrogn- nit st of maningful
iry, th whol strivs to rturn to homognity' t a highr lvl' This is a f l x i v n s s i,n w h i |
lassi as of Spnriandynami quilirium. gua8'nd prvarieti
In rnt yasl th shifting aln thory has n vry svrly riti- sags. Although langu
izd, although rrstvolutionistsstill think that thr is som rol for gn- Kanzi, show subtl sig
ti drift (sCon, Barton nd Tr'rrlli1'997).At th molulr lvl, nath t h i r s i g n do o n l P u t
th ffts of natural sltion, drift is thought to b vry importnt (Kimura muniations rmain
1983) and is th asis of th major wy of alulating signifiantdats in vo- By th tim humal.r}
..gnti ..Molular volu. on hr parnts and ask
lution (through loks''). (S th main ssay
tiorr'' y Franiso J. Ayala in this volum.) y a r - o l d h i m p a n z ,
munirionon th orr
B|LIoGRAPHY tl.rirlanguag skills to
Brgson'H. 1911.rtiuuollltil.Nw York:Holt. produing th full rang
oyn,J. A., N. H. Barton'and . Turlli.1997.Prsptiv: A ritiquof Swali hominins volv lngu
.Wright's
shiftinglanthoryof volution.uolutkl51' no. 3: 64-671. from th anstorsof tl
)ozhansky, T. [|97] 1951,Getisd th rigi of Spis'rd d. Nw York:
Shllirrshav rrt
olumiaUnivrsityPrss.
Fish'R. A. 1930' he GticIThor of Nturl Sltion.oford: Oford win's tim. Nonthls
UnivrsityPss. m o u s l y a n n d a I I t - ' r
Hndrson,L' J. 1917.Th rdr lfNtre.mridg,MA: HarvardUnivrsity languag ausof its
Prss. nams that hav bn g
,b -rlr1'
Kimura, . 1983. Nutrl of oleulr ullutio.amridg: sis, for xampl, sl-t
amridgUnivrsityPss. ..moo,
sounds suh as
Provin,!. B. 1986.SuIIWrightttltdurllutilrl,Birllog.hiago:Univrsity
lf hiago Prss. attiuts th nrg,
Rus, . 1996.ond to : Tb oeptof Progrssi uolutilrBiolog. atd with physial l;ro
Camridg,MA: Harvard UnivrsityPrss. originatd from moti
1999.ster of stris:Is uoltion SliIostrutil?mridg' h y p o t h s sd o n o t l n d
A: Harvd UrrivrsityPrss.
Notions out wh
Snr,L7.1862, FirstPriipls.London:Willimsand Nrgat.
Wright,s. 193l. volutionin \,lnd|ian populatins'Gntis16:97-159. today as thy wr a
192.T|.lrolsof mutation,inrding,rossding and sltionin millions of yars old;
volution.Prodigsof th Sith ltrtiologressrlf Gtis1: Among th lattr. so
356-366. . -.R. arisn bfor th
..ultu
was hratrizdy
als, prsonal :rdom
volution of languag Uppr Palolithi tooln
Although many animals ommuniat using voalizations' only humans us to ston' and thir too
ompl languag, asd on a disrt numr of aritrary its of Sounds mor spilizd for p
and gsturs' to omprhnd and produ a smingly infinit numr of agr that Uppr Palo
novl mssags.Listrringto sph nd rading r snsory aspts of ln- 40,000 yars ago' thy
guag' whil talking, writing, and ody languag ar motor aspts.In th o l o g i s t s l i v t h l r t t h
|960s, hrls Hoktt listd 15 univrsal dsign faturs that distinguish sumption of linguisti
.W
human languag from ommuniation systms in othr animals. oktt n plad on th
viwd thr fattrrsas partiularly importnt for lruman languag:(1) dis- std, that numrol1s
plzrmnt,or bing al to ommIriatabout mattrs that ar rmot in points, vidn of tr
tirn and sp; (2) produtivity, whih is tlr ability to ommuniat an o u r r dm u h r l i r '

passdon y \/right'stahr,
.ind of alan, homognous;
fragmntation and htrogn-
ity, at a highr lvl' This is a
1.
y has n vry svrly riti.
h t t h r i s s o m r o l f o g n -
At th molular lvl, nath
t to vry important (Kimur
u l a r i n gs i g n i f i a n d
t tsin vo-
ain ssay
..olular volu-
Holt.
Prsptiv: A ritiqu f Swll
u o l u t i o 5 1 ' n o . 3 : 6 4 _ 6 7 \ .
,igi of Spcis. 3rd d. Nw York:
rl Slectio' Oxford: oxford
r b r i . l g . A : e r r . n r dI I n i v
' namsthat hav r-r givn to various hypothss.Th ..ow-wow'' hypth-
r u,,Itil,C.lrrlhridg:
r Bi,,log.Chig<-l:
Univrsiy
'rogrssi ulltior Biolog.
Soil ostrution! Carrrridg,
ms nd Norgar.
n s .G l i s | 9 7 _ | 5 9 .
: r o s s r d i ngn d s I t i o ni n
l ogrssof Getis'-,.*.
voalizations,only humanS us
r of arbitrzrryits of sonds
a smingly irrfinit nmr of
ling ar snsory aspts of lan-
]uagar motor aspts. In th
dsign faturs that distinguish
ms in othr animals. Hoktt
nt for human langug: (1) dis-
) o u t m t t st h a t a m o t i n
t h i I i t vt o o m m u n i a t n ourrdmuh rlir (prhapsas long ago.1s 10,000 yars) in th Afrian
uolutio of Lgug 56
opn-nddvarity of novl mssags;and (3) duality, in whih a small num-
rof meaninglssphonologial lmntsmay omind to rat an infi-
nit st of maningful uttrans.Two othr intstingdsign faturs ar
flivnss,in whih on an us languag to olmuniate aout ln-
guag,and prvariation,why languagan sd to dlivr flsms.
sags.Although langug_traindaps suh as th wll.known onoo,
Kanzi, show sutl signs of utilizing som of Hoktt's dsign fturs in
thir signd or Omputr-failitatd ommunitions with popl, thir om-
munitions rmain xtmly simpl dspit dads of intns shooling.
By th tim a human hild is four years old, sh is likly to fastn a ady y
on hr pants nd ask, ..\/hrdid I om fronr?'' Th highly traind four-
yar-oldhimpanz,on th othr hnd, is likly to sign a spontanousom-
nruitionon th ordr of ..ooki,ooki, hurry.'' Aps, in ft, nvr us
thirlanguagskills to ask qustions.Furrhr, thir larynxs ar inapl of
produingth full rang of human sphsounds. How and whn, thn, did
hominins volv languag during th 5 to 7 rillion yars sin thy divrgd
from th anstorsof thir losstgntiousins, th himpanzs?
Sholars hav n fasinatd by this qustion sin t last harls Dar-
win's tim. Nonthlss,in 1866 th influntial Linguisti Soity of Paris fa-
mously annd all onrmuniationsdaling witlr tlr origin and volution of
s i t y languagausof its spulativnatur, whili is valdin th amusing
sis, for ampl, suggStsthat languag gan as imitations of natural
sounds suh as ..rnoo''' ,,ozz,'' nd ..rash'';th ..yo-hav-ho''hypothsis
attriuts th mrgn of languag to rhythmi hants nd gunts assoi-
atdwith physial laor; and th ..pooh-pooh'' hypothsis suggstslanguag
originatdfrom motional ris or lamations. Although intrsting,ths
hypothssdo not lnd tmslvsto sintifitsting.
Notions aout whn nd lrow languag first aros r as ontrovrsial
tody as thy wr il ntury ago. Sonl rsrhrsthink tht larrguagis
millions of yars old; othrs liv it originatd a mr 40,000 yars ago.
Among th lattr, s<rmarhologists think that languag ould not hav
arisn for th ..ulturl plosion'' of th Uppr Palolithi in urop that
was haratrized av paintings,portal art, living sits,dliraturi-
ls, prsonal adornmnts' hunting and fishing thnology, and signs of trad.
Uppr Palolithi toolnrkrs usd and ngravd ivory and bon in addition
to ston' and thir tools wr ttr mad than rlirons and frquntly
mor spilized ior prtiuiar tasks. Althor'rghmost anthropologists wold
agrthat Uppr Palolithi innovations indiat that languaghad zrrisny
40'000 yars ago' thy disagraout whn it iniially mrgd.Som arh-
ologistsliv that th prior matrial rord is too simpl to justify an as-
sumption of linguisti haviors. Othrs think that too muh mphasis has
'Wstrn
n plad on th uropan Uppr Palolithi; thy suggst, in-
stad' that numrous potntial signs of languag inluding ard spar
points, vidn of trad ntworks, and th r'rsof rd pigmnt (h)
564 ultil<lfI'gg

rord of th Middl Ston Ag. Rathr thar-rsing |anguag as having vrrtuallyinvdd lang
mrgd suddnly and rlativly rntly,thy livit volvd mor gradu- t r g to n a t u r l 5 l t
ally ovr muh longr priods of tim. x p l i i t l y r j h n o t i
othr sintistshypotlrsizthat langug lrigirrrdstill urrlir.Skltal all systrns,lthor-rglrt
iologists who study th volution of th parts of th skull ald nk that f- . . t o p - d t l w n ' 's h o l o f t
ilitat sph livthat th laryn and throat had volvd to thir modrn mans volvd th ailit1
forms y 150,000 yars go' thus alllwing th rapid vlalizationsthat har- infinit numr oi rul-
aterize modrn languags.Thy lso:rknowldg that sph rrdnot ntail nuity hy;lothsis,''on th
th full rang of ontmporry voalizations nd that rudirrllrtarylanguag of inrmntal volutior
may hav istd at lst a million yars ago. and ss its roots rflt
Palonurologists,who study th volution of th rain, liv that lan. wild primats hav in t
gug mrgd vn arlir, around or for 2 rnillion yilrs ago. Thv man) xtnd th disuss
s t u d y a s t s o f t h i r r s i d so f f o s s i l s k u l l s ( n d o a s t s t) h t r p r o d u d t a i l s alls twn rnothr t
from th outsid part of th rain (or rral ort) that rvr imprintd Nl disussion of lan1
on th walls of th rains whn hominins (or othr animals) wr aliv. ing th wll-known gs
Thy also lrart th inras in rni:rl apaity (as a sur<;etfor rairr on th dp psyholtlg
siz) that ourrd durir-rgth ours of hon-rinin vOlLltion. Altlrtlugh guag' iniudir-rgmoth
spaking dpnds on ativity in vrirlus parts of th rain, I}roa'ssph til, and gsturl omp
aa on th lft sid of th frontal los is spially important for oordi- w also us flailing han
nating th intrit movmnts of th tongu' mouth) and larynx tht r physil ontt rr,ithp
nssary for sph. Broa's ara is assoiatd with a spifi pattn of mats also tlmirrsu
onvolutions that dos not appr in th rins of our losst gnti siorrLrllyolfatoy signa
ousins' th grat aps. This prt of th brain is somtims rprodud on diffrnis that, in ad
ndoasts. ints rvald on ndoilsts from f<lssilhominins suggst th:rt primilt produs voa
th frontal lo sph ar:rmay alrady hav gurrdvlopirrgy 1.9 mil- abov. Alvotslf g
lion yas gl. Studis f ston tools from th sam tim lso suggstthat v r l t u a l l yt l I l g . u gi
hominins wr in th pross of oming right-handd, whih aps as a 6nd narrowly (.g.,mz
population 21rnot. It is important to not, howevr, that rains had not yt tht rodu visuzrl,
nlargd to nywhr n:lr thir molrn sizs, and that th vi]nfronr M 1 ' o w n i d s. r r i n
ndoasts rgarding Broa's ara is diffiult to inrrprt. Th suggstion of livthat th oots o{
rudimntary languag t this arly point in tim should thrfor viwd rsnts a kind f sign
as intrsting ut spul:rtiv. mrgn of th first 1
Languag and populatior-r-lvl righr-hnddnssoth dpnd lravilyon th himprrzmothrs us
lft sid of th rain. Nonhuman pimatshv r:rinsthat a lso asynrnrtri. lnothrs dlivr e ons
al (or latralizd), to som dgr, ut thy do not gin to pproah human ..mothrs,''or tnusi
lvls of nurologial asmmty. In ddition to lirnguag funtions, th lft fatttrshlp aisr
hmisphrs of humans r diffrntilly invlvd irr skilld movmnts' suh gugs.himpanz m
as thos of th right har-rd, as wll as analytial,tilrr-squning prosss.Th tir infants, who rm
right hmisphr,n th othr hand, lsat mor gloal purstlits,suh as vi- unaidd' to thir hairy
suospatial and mntal imaging, ognizing fas, appriating humor, and voltltion of walking u
haring and produing mr.rsi.It also ontrols th lft hand. Although it is th thy do rtain a grasp
ltt sid f th rain that spks, it is tlr rntrsilright hmisphrthat pro- t h i h l p l s si n f l l l l t ' i l
vids th ton of voi, or prosody, lvhih olors our uttrans.In this sns, Un]rths irumsta
languag dpnds on th right as wll as th lft hrnisphr. adaptiv for assurin
Som sholars' howvr, think that prosody hd nothing to do with th movd from thir rnot
mrgnoi lnguagal-rd,instad,dvlopds ll kir-rdof uditor1'frill tht kirrd lf dismodr

tan sing languag as having
r iivit volvd more gradu-
originatdstill arlir. Skltal
s of th skull and nk that fa-
lt had vllvdto thir modrn
rapid voalizationsthat har-
dgthat sphnd not ntail
and that rudimntary languag
l o f t h r a i n , l i v t h a t l a n -
r 2 millin yars ago. Thy
coasts)that rprodu dtails
'al ort) that wr imprintd
; ( o r o t h r a n i m a l s )w r a l i v .
lity (s a surrogat for ain
hominin volution. Although
ts of th rain, Broa's sph
spiallyimportant for oordi-
l' mouth' and laryn that ar
rtd with spifi pattrn of
r i l l s o o u r | o s s tg n i sionally lfatory signals, in thir blnd of multinrodal ommuniation. Th
in is somtimsrprodud n
n f o s s i l h o m i n i n s s u g g s tt h t
b g u nd v l o p i n g y 1 . 9 m i l -
h sam tim lso suggst that
right-handd,whih aps as a
owvr'tht brains had not yt
s, and tht th vidn from
to intrprt.Th suggstion of
im should thrfor viwd
lnssoth dpnd havily on th
- , r a i n sr h a t r l s o s y m m r i -
o not ginto approah human
to languag funtions, th lft
lvd in skilld movmnts, suh
, tim-squningprosss.Th
moglobal pursuits' suh as vi.
fas, appriating humor, and
th lft hnd. Although it is th
sial right hmisphr that pro-
J 1 5o L l rU t t r a n sl n. t h i s s n s ,
ft hmisphr.
dy had nothing to do with th
d as a kind of uditry frill that
uolutil of Lgug .'6.'
vntually invadd lnguag.Thy also assrt that musial skills wr not a
targt of naturl sltionin th sme way that linguisti ailitis w, and
pliitly rjtth notion that languagould hav volvd from primatlik
all systms' although thy aknowldg that ton of voi likly did. This
..top-down''
shool of thought thrfor fouss on undrstnding how hu.
mans volvd th aility to link symoli rfrntialsounds into a potntially
infinit numr of rul-govrndmaningful sntns.An opposing ..onti-
nuity hypothsis,''on th othr hand, posits that languagmrgds a rsult
of inrmntalvolution from th primat all systmsof our erly anestors
and ssits roots rfltdin th ominations of prosodi alls that various
wild primats hav in thir rprtoirs.Som ontinr-ritists(.g.,John Nw-
mn) tnd th disussion vn furthr into th past y fousing on ontat
alls btwn mothr nd infant mammals.
No disussion f languagvolution would mplt without mntion.
ing th wll-known gstural.originshypothsis,whih rognizsand uilds
on th dp psyhollgiallinks twn thought, sph,and gstur.Lan-
guag' inluding mothr-infant intrations' nompasssvoal, visual, ta-
til, and gstual omponnts. Humans do not just ..talk'' with our vois;
w also us flailing hands, faial prssions,ody postur' and (somtims)
physial ontat with popl or ojts to onvy maning. Nonhuman pri-
mats also omin suh gsturswith motional voaliztions, and oa.
diffrn is that, in addition t motionl alls and inttlnations, th human
primat produs voliztions that hav th linguisti proprtis outlind
aov. Advoats of gstural origins think that th natural sltion that ld
vntuallyto languaginitially targtdgsturalommuniations, thy d-
find narowly (..g., manual gsturs)or mor roadly (rtiulatory gsturs
that produ visual, aousti,voal, or tatil signals).
My own idas ar in kping with th gstural hypothsis,writ roadly. I
livtht th roltsof languagar tmlyanint and that prosody rp-
rsnts kind of signatur that is lft ovr from vnts that prdd th
mrgnof th first languag.Although th ody langugthat human and
himpnz mothrs us with thir infants is virtually idntial, only human
mothrs dlivr a onstant stram of ay talk, whih is vry prosodi. Suh
..mothrs'''rrrusial
sph,is univrsal among humans, and its eousti
fturshlp ais round th world larn th strutursof thir nativ lan-
guags.himpanz mothrs dirt fw, if any, spial voalizations toward
thir infants, who rmain physially atthd to thir mothrs y linging'
unaidd, to thir hairy hsts.Bausof dvlopmntalfftsrlatd to th
volution f walking upright, human infants hav lost this aility, although
thy do rtain a grasping rfl. arly hominin mothrs must hav arrid
thir hlplss infants in thir arms' at last until ay slings wr invntd.
Undr thsirumstans'soothing nratrnalvoalizations would hav n
adaptiv for rassuringnary infants tht wr, of nssity,priodially r-
movd from thir mothrs' mras.Ton of voi may thus hav volvd as
a kind of dismdid tnsion of mothrs' radling arms. Taking a u
566 uolutio of Lgg

from how modrn infants gin to aquir languag' it is rasonal to sur-

mis that natural sletion sulptd inrmntal hangs in mothr-infant
volution of lif h
prosodi' tatil' visual, and gstural intrations,whih failitatdth mr- W h y d o s o m p l a n t s ''
gn of linguisti faturs and, vntually (prhaps far down th lin)' th yars aftr grminatin
first languag. Ndlss to say' lik th othr idas aout languag origins, as th nnuals in your
this ..putting th ay down'' hypothsis is ontrovrsial. d o s o m n i m a l s ,s u h
Baus of th wid disparity in urrnt njturs rgarding th volu- omprativly fw t
tion of languag,n might tmptd to onlud that rsarhon this su- dwlling krill' prdu
.W.hat
jt has not improvd mh sin th days of th ow-wow, yo-hav-ho, and ar th onsqu
pooh-pooh hypothss.This is not tru' howvr. Advans in mdial imag- tion of ths organism
ing thnolgy and th nurosins ar urrntly prmitting luidation of ralm of lif-history v
th nurologial sustratsof languag (.8.,th rol of mirror nurons);th A n o r g n i s m ' sl i f h
innat natur of languag is oming larr from fasinating rsarh on t h k y s t a g s n d t r r
daf individuals; and gns that hav a rol in th dvlopmnt of normal p o d u t i o n .L i f - h i s
spokn languag (.g.,FoP2) ar bginning to idntifid. Languag is p o d U t i o n la g a n d s i ;
what sts us apart frm th othr primats, and today w hav mor sophis- spring;and th g tll
tiatd idas aout how it volvd than w did a hundrd yars ago. Bause oftn vary sustnti
of th information xplosion that is urrntly undr way. w should soon triguing ways. Th goa
know mor. in th pattrns of gror

|BLIoGRAPY 8 n r a t h i s v a r i t i n
this variation. Th stud
Armstrong,D. F.' !. . Stko,nd S. . Wilox. 1995. Gstured the Nture of quirs intgrating on<
Lguge.amridg:amridgUnivrsityPrss.
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Falk, D. 24a,Brinde:Nt'uDiscrlurisbout Hum rigis d Bri
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2004. Priinguistivolutionin arly hominins:Whnmothrs? n l y s i si n h u m e n s .l n
Bhuiorld Bri Sis 27:491-541. t n so f h u m n m o r t e
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Hoktt,, 1966.Th prlmf univrsals in langug.In J. Grnrg,
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Uniuerslslf Lguge,1-29. amridg,MA: MIT rss.
Holdn, . 1998.No lastword on languagorigins'Si282: t455-I458. futur ptd rpo
-h 'Word: haratristis suh as
Knnally,. 27. First Th Serchfor th origis of Lgug.Nw
Y o r k :V i k i n g . varials ar dirtly rt
Laitman,J. T.' J. S. Ridnrg, and . J. Gannon.1992.Fossilskullsand hominid n r p o d u t i v r t )r
voal trats:Nw approahsto hartingth volutionof human sph.In
individuals that produr
J. !ind t al', ds.,I,ngugrigi: ultidisiplirpprlh,395_407'
Dordrht,Th Nthrlands: Kluwr admiPulishrs. th dirt Iink tw
asataka,N. 2003. he onset of Lnguge.amridg:amridgUnivrsity shaps th divsity of
Prss. | s r l s o n t r a il
l\4Brarty'S., and A. S. Brooks 2000. Th rvolutiontht wasn't:A nw ropi of importn r
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o d r n l i f - h i s t o r yl
Nwman,J. D. 2004. othrsy any othrnam:othr-infantommuniation
in non-homininmammals.Behuiorlnd Brin Sics27: 51'9_520. o n t o l i n t h 1 9 5 0 s .]
.William nithologist who infus
Pinkr' S. 1994. b Lgglstint.Nw York: orrow and ompany.
Savag-Rumaugh, S.' S. G. Shankr,nd T. J. Taylor.2\.Apes,Lnguge,d H i s m o s t i n f l u n t i a l a
the Hum il. Nw York: oxford Univrsity Prss tht irds ly in a nst
maximizs th torl o
known as th ..Lak l

ag'it is rasonal to sur-
ll hangs in moth.infant
;, whih failitatd th mr-
haps far down th lin), th
las aout languag origins,
rovrsial.
turs garding th volu-
rd that rsearh on this su-
bow-wow, yo-hav-ho, and
. A d v n si n m d i a l i m a g -
rtly prmitting luidation of
: rol of mirror nurons): th
:rom fasinating rsarh on
th dvlopmnt of normal
o r d n i f i dL. a n g u a g i s
tody rv hav mor sophis-
r hundrdyars ago. Baus
undr way. w should soon
)95. Gstred t Nture of
ss.
Humttorigis nd Brin
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-rins:.Whnmothrs?
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: IT Prss.
Sci2-82z1'455_|458.
th Origis of Lguge.New
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uolutiolt of Lif Histlris 567
vlution of lif historis
Why do som plants, suh as rdwood trs' gin to rprodu only nrany
yars aftr grminating nd liv for hundrds of yars, whras othrs' suh
aS th annuals in your gardn' mtur youllg nd liv only on sason?.Why
do som ar-rimais,suh as humans, produ on offspring at a tim and rar
mparativly fw throughout lif, whras othrs, suh as th oan-
dlvlling krill, produ thousands of young, most of whih annot surviv?
Wht are t onsqrrsof thsdiffrnsfor th ology and th volu-
tin of ths organisn-rs? Ths, and a varity of rlatd qustions' ar in th
ralm of lif-history volution.
An organisrn'slif history is its pattrn of survival and rprodution' plus
th ky stagsand transitions in th lif yl that influn its survivl and
rprodution. Lif-history traits inlud growth and dvlopmnt for r-
prodution;ag and siz t sual maturit; th numr, siz, and se of off-
spring;and th ag of dath. Spis,zlnd vn p<rpulationswithin spis,
oftn vary sustantially in lif-history hratristisin prdital and in.
triguing ways. Th goal of lif-history studis is to undrstand this variation
in th pattrns of growth and rprodution, th ologial onditions that
gnrattlris variation, and th logil and volutionary onsqunsof
this variation. Th study of lif historis, whthr mpirial or thortial, r-
quirs intgrating onpts of gntis nd dvlopmnt with ology and
with an urrdrstandingo h organism's lit yl.
Atuarial studis y insuran ompanis ar an xampl of lif-history
nalysis in humans. Itrsuranatuaris us lif tals to dsri th pat-
trns of human mlrtirlit1,at spifi a8s to ssss lrow ag influns th
risk of dath. ologial lif tl analysis is asd on ag-spifiirth nd
dathrats, from whih sintistsan alulat a vaity of usful varials
sulras th nt rprodutiv rat (th total numr of offspring produd),
futur ptd rprodutiorr (alld rprodutiv valu), nd mrgnt
haratristissuh as th rlationship of rprodution to mortality. Ths
variablsr dirtly rlatd to Iitnss,a fur-rtionof total rprodution (th
nt rprodutiv rt) nd th shdu]of prodr.rtion; fitnss is higirr fo
individuals that produ rlativly mor offspring mor quikly. Baus of
th dirt link twn lif-history traits and fitnss, natural sltion
shapsth divrsity of lif-histlrYstratgiesobsrvd in natur. Ths vari-
ls ar lso ntrlto how ologists prdit population flutuations, a
topi of importan to th managmnt of harvstd or prottd wild
spis.
odrn lif-history nalysis tras to Dvid Lak in th 1940s and Lm-
ont ol in th 1950s. David Lak (1947) was a population iologist and or-
nithologistwho infusd Darwinian thinking into th study of lif histories.
is most influntilarly lif.history hypohsiswas tlrt th numr of ggs
-D.F. that irds l:ly in a nsting attnlpt (thir ..luth siz'') should that whih
maximizs th total offspring fldgd from that luth. This has om
known as th ..Lak lrrth Siz.''To mximiz th podution of fldglings,
568 uolutio of Life istoris

parnts must alan th numr f hathlingswith thir aility to rar thm funtion annot .
to indpndn. If parnts attmpt to ar mor offspring than thy an sup- offs, and how hav
port' som or all of th young may di. Likwis, prnts prduing fwr yl influn survil
offspring than thy an rar will fldg fwr than thy ould, and will lik. th ntral qustion i
wis hav lowr rprodutiv suss than if thy had produd a largr sptivs of Lak anl
luth. lif historis that hs
primntal studis on many spisof irds, in whih luths ar rtifi- ativ today.
ially nlargd,hav found that luth sizs produd y irds ar gnrally Modern lif-histor
smallr than th Lak luth Siz (.g.,Vandr.Wrf 1992). This disrpany th nvironmnt' d
twn th Lak and th osrvd luth sizs stimulted a grat divrsity of strain lif-history val
xprimntal and thortial study that in turn gnratd many adaptiv hy- an volv rapidly w
pothss' mainly drivd from optimization analyss (.g.,..optimlity mod- dution of prdator
ls',; Orzak and Sor 1994) that am ommonpla y th 1970s and onto nw host sp
1980s. For ampl, modls prdit and xprimntal studis hv shown 2007). Seltiv prd
that urrnt yar rprodutiv ffort ffts futur rprodutiv suss of has ld to th voluti
parnts; that is, thr is a trad-off twn urrnt and futur rprodutiv tns of gnrations(I
ffort (s rviws in Godfray et a|. 1991 and Rlff 1992\. \/hn this and tion favors rprldu
othr trad-offs (suh as that btwn luth siz and gg siz; Smith and shard volutionary l
Frtwll 1'974) are inorporatd into modls of luth siz, w n plain isms, gnti varitio
muh of th srvd dviation twn rl luth sizs and Lak's pr. volution of lif-hist
ditd luth sizs. tionary biologists to
Following on Lak's introdutin of Darwinian thlnking into th study of ation, dvlopmnta
lif-historyvariation, Lamont ol (1954) introdud a mathmatialfram. so that aus-and-f
work to th study of lif.history ology y dmonstrating hlw hangs in d- Th Darwinian pr
mographi varials influn th rat of inras of a population (Roff and othr filds of
7992). ost famously, h raisd a simpl mathmatial onundrum known as n and lif span in
ol's parado: an rgnism that livs forvr, produing offspring pr flund y an organi
yar, lavs no mor dsndants' all ls ing equal, than on that livs for rprodution. In mos
just on yar (an annual organism),ut produsz + 1 offspring.Th rason is lins with ag. Thur
tht nly on offspring is rquird to rpla th parnt in our annual organ. lss than dos morta
ism, aftr whih oth th immortal and nnul organism produ th sam ar undr wakr sl
numr of offspring. Givn th impossiility of irnmlrtlity, why do not all this dlin in th int
organisms volv toward th shortst, simplst lif history? any iologists, Snsn.oupld
.Williams,
inluding Willim D. Hamilton, Gorg and ri harnov traits that fvor su
(harnov and Shffr |97), hav takld this onundrum, and its solution ost to ultimat long
ws sminal to th formalization of Darwinian thinking in modrn olgy. gns hav n idn
Th solution to ol's parado rquirs rognition that ah lif-history d- only on th old an r
ision auss a shift in th proility f aruing rtain nfits,and rtain mulation of dltrio
osts, in trms of survival and rprdution.For xampl' if rprodutivsu- N4or powrful mat
ss inrass with ag (.g., du to nfits of parntal prin) and th mor insightful pri
risk of motality twn rprodutiv priods is low, thn sltion may fa- lif.history variation l
vor rprodution for multipl sasons. In <lntrast,whn th risk of mortality in genrating nW spe
is high, seltionmay favor alloatingmor rsoursto urrnt rprodution. latd into th applid
Alloation trad-offs suh as ths onstrain how lif historis volv. R- volution influns tl
sours' inluding tim, nrgy, and ssntial nutrints' must dividd tainability of agriult
among growth, maintnan,and rprodution. Rsours alloatd to on and disasrlatsto

with thir aility to r2rthm
' offspringthan thy an sup.
vis'parnts pr<lduirrgfw
han thy ould, and will lik-
tlry had produd a largr
ls, irrwhih luths ar artifi-
'odudy birds ar gnrally
Wrf 1992).This disrpany
stimultda grat divrsity of
r gnrtdmany adaptiv hy-
..optimlity mod-
ralyss(.g.,
|mmonplay th 1970s and
rimntlstudis lrav shown
uturrprodutiv suss of
rrntand futr-rrrprodutiv
,i Ro{{ .'991-\1.\l/h..--t. r:.d
l siz and gg siz; Smith and
of lutlr siz, w an xplain
ll luth sizs and Lak's pr-
nin thinking into th study of
rodud 21mathmatil fram_
monstratinghow hangs in d-
nrasf a population (Rff
nmatialonundrum known as
r'r' prtlduing iz offsping p
rg qual,than on that livs fr
s/7+l ffsprng'h ason is
th parnt in our annual organ-
ual organism produ th sam
f immortality, why do not all
st lif history? Many iologists,
Williars, and ri Charnov
his onundrum, and its solution
n thinking in modrn ology.
nitionthat ah lif-history d-
-ringrtain nfits, an] rtain
jor
x a m p l 'i f p r o d u t i v s u -
of pantrlxpin)and th
Js is low, thn sltion may fa-
trst, whn th isk of mortality
:sours tO urrnt rproduti<rn.
,in how lif historis volv. R-
rial nutrints, must dividd
ion. Rsoursalloatd to on
uolutiolt of Life Histories 569
funtion annot lloatd to othr funtions. Undrstanding ths trad-
offs, nd how havioral or dvlopmntal disions at on stag of th lif
ylinflunsurvival and rprodution at othr stags(and totl fitnss),is
th ntral qustion in th study of lif-history volution. Th Darwinian pr-
sptivsof Lk nd ol ouglrt nw lvav of thinking into th study of
lif historis that has ld to myriad insights and a fild of inquiry that rs vry
ativtoday.
odrn lif-history analyssaim to undrstand how volutionary history,
th nvironmnt, dvlopmnt, and inhritan intrat to rat and on-
strain lif-history v:rriatior:r.An irnportant rnt insight is that lif histois
an volv rapidly whn nvironmnts hang (.g.,th xprimntal intro-
dution of prdtos, olmrizll xploitation y humans, and pansion
onto nw hst spis y parasits of plants and animals) (arroll t al.
2007). Sltiv prdatior-ror lrarvsting of lrg individuals, as in fishris,
has ld to th volution of sustantially rdud ag and size at maturity ovr
tnsof gnrirtions(Huthings nd Frasr 2008); this lrpns ussl-
tion favors rprodution for animals ar atn r harvstd. Thus, whil
shardvolutionary lristor},tndsto rat similritis among rltd organ-
isms, gnti variation for lif-history trits is aundant and allows th rapid
voiution of lif-history stratgis'Suh ongoing dptation allows volu-
tionary iologists to wigh th rlative influnsof phylogny' gnti vari.
ation, dvlopmntal systms, and th nvironmnt ir-rshaping lif historis,
so that us-nd-fftrlationslripsom larr.
Th Darwinian persptivon lif historis an ring insights to mdiin
and othr filds of iology, inluding publi halth onrns suh as sns-
nand lif span in aging human soitis.Th volution of lif span is in-
f]undv arr oganism'srprodutivvalu, th masur of futur ptd
rprodution. In most organisms on rding starts rprodutiv valu d-
lins with ag. Thus mrtality lat in lif affts total rprodution muh
lss than dos mortality arly in lif, so taits ffting mortlity lat in lif
ar undr wkr sltionthan ar traits afftingmortality arly in lif. It is
this dlin irr the irrtnsityof sltionwith ag that llows th volution of
snsn. oupld with th sltivdvantag of short gnration tims'
tritsthat fvo sussfulrprodution ariy in iif will fvordvn at a
ost to ultimate longvity (an ampl of antagonisti pliotropy); indd,
gnshav n idntifid tht hav ths ffts. Traits with ngtiv ffts
only on th old an volv in populations through gntidrift (i..,th au-
multion of dltriousmutirtions).
or powrful mathmatial tools, nw thnologis (.g.,gnomis), and
moinsightfulxprimnti.ttion pmittingiologiststo ttrundrstand
lif-history variation within and mong spis and th rol of suh variation
in gnratingnw spis.Th rsulting insights ar inrasinglying trans-
latd into tlr applid sins whr iologists ar studying how lif.history
volution influns th onsrvation of ioti rsours,th yild and th sus-
tainailityof agriultur,and how th volutilnof ontogny, rprodution,
and disasrlats to human halth.
570 uolution of Se

lLIoGRAPY
s-slnd to hirv
aroll,s. P.' A. P. Hndry,D. N. Rznik,and . W. Fo.
2007.volutronon lonirl fmal trans
ologialtim-sals.Functiolcolog 21: 87-9.
harnov, . L., and W. A4. Shaffer. 1973.LIi-historyonsquns mle trnsmits onl1
of natural
sltion:ol's rsultrvisitd.Amri Ntrlist|oi| hlf of hr parnt
79]..-79.
ol, L. . |954. Th populatiollonsquns of lif historyp.no.". Qurtrl mal' Thus s arri
Ruicuof Biolog29: |0_137. hug thortial dis
Godfra,H. . J., L. Partridge,and P. H. Harvy.1991.luth
siz.l Ruie erl alld ..th p;
of olog d Sstmtics22: 409429,
Huthings,J. A.' nd D. J. Frasr.2008. Th naturf fishris- spurrd svral lad
and farming-
indudvolution.olulr colog 17:294-13. highligliting th imp
Lak, D. 1947.Tsignifian of lut..i,.. Ibis89: 3o9-52. lutionary iology'' (
ozak, S. H.' and . So.1994.optimlitymodlsand fits of sxuality' it rv
th tstof 1aptatlon.
meriNturlist143:361-380.
twi as fit, on avr
Roff' D' ^. |992. h Fuolutio of Lif Historis:Thor
lnlsis.Nw Yok: quird to Ovrom1
hapmnand Hall.
Smith,C. ., and S. D. Frtwll, 1974.Th optimal alan viduals an havthi
twnth srzand
numrof offspring.Ameri Nturlist 108: 499-506. in a world that hal
Vndr.!rf, . 1992.Lk,sluth siz hypothsis:An aminatiorrof ally, s an out()
th vidn
usingmta-ana|ysis. olog 73:1699_1705. -s.P.' d .\Y.F. (1) kills a lrg rl
ft o podution
dividuals wit]r rtir
vlution of s () hangs rirdially
Hr,you se,it taksall th runningyou n do, to kp tys ar killd and l
in th sanr tion for s, th ph
pla.
_h Rd world, ffting mosl
Qun,in Lwis aroll's
Throughth Looking Glss(1872) ? In rnt dd
disas (Hamilton 20
Th ida tht inf
of th many manings of th word se*' th most fundamntal mak s dvntg
rfrs to th
prodution of nw gnoms by th romination th spts of an org
of pristing gnoms.
any batria and a fw viruss ar apabl of oasional. th thorists' dsrip
limid s' ut
th-most thoroughly sxual raturesar th ukryots, Bs parsirs an
whih iniud ani-
mals' plants' and othr organisms with nuli in quikly ompard to
thir lls. Humans ar
fairly typial ukaryots in ing oligatly sul: gnotyps. on striki
th only wy to mk
nw human is to split your gnom in half and to gt tions should ourin-
anothr lrali from som
othr uman who has don th same. Dspit th ar ommon in on
ntral rl that sx plays
in organismal lif yis, iologists do not undrstand Qun hypothsishas
what its funtion is.
Th first biologist to suggst a plausil hypothsis is of hosts and para
for tlr dvantag of
sxuality was August.V7ismann(1ss9). H proposd fls, ut sintistsd
that s inrasdth
vaiability within a population and thus inrasd th to .
fftivnssof natural
sltion. This is an unusual sort of advantag; typially, Th othr phnon-r
natural sltion
promots traits that onfr an advantag to th individual ful mutation. With hi
.!(i'ismann' that ars th trait.
howvr, saw sx as a fatur of a populatin that ronrbirrant offsprin
onfrrd an
advantag to that population. .sismann'seplanation This is orriy tru if rat'
was influntiai nd
wnt largly unhallngd until th 1960s, whn Gorg individul arryig <lr
. Williams o-
jtd to its rlian on th ruttion thory
8roup sltin and insistd tht sx should hv
short-trm, individual-lvl dvantg lik ny other harmful mutations [
trait. On th onrrarv.
msur' t stilnat

\W.Fo.2007. volution on
87-39.
'ory onsqunsof ntural
urIist107l791_793.
.ifhistory phnomna. Qurterl
l991. luth stze.AnI Reuitu
f fishris-and farming-
-3 13.
8 9 :3 0 9 - 3 5 2 .
ils and th tst of adaptation.
bor d lsis. Nw York:
Ll alan twn th siz and
i:499-506.
;:An xamination of th vidn
-s.P.C. d ''\Y.F.
tn do,to kpin th sm
.dQun,in Lwis rroll's
h thLookig Glss (|872|
most fundamntal rfrs to th
nation of pristing gnoms.
of oasional, limitd sx, but
:ukaryots,whih inlud ani-
li in thir lls. Humans ar
sual:th only way to mak a
rd to gt anothr half from som
t the ntral ro1that sx plays
n d r s t n dw h t i t s f u n t i o n i s .
hypothsisfo th advantag of
propsd that sx inrasd th
asdth fftivnssof natural
tag; typially, natural sltion
h individual that ars th trait.
f a population that onfrrd an
planation was influntial and
\/illiams o-
, whn Gorg .
1 insistd that sx should hav a
1ny othr trait. On th ontrary,
uolutio of S .t71
ssmdto hav an normous short-trm,individual-lvldisadvantag.A
lonal fmal transmits all of hr gns to all of hr offspring. A sual f-
mal transmits only half s many gns to th nt gnrtin, and wsts
half of hr parntal invstmnt propagating th gns of som unrlatd
mal.Thus s arris an normous (twlfold)ost. Th ontrast twnthis
hugthortialdisadvantagof sx and its pvalnin th ral world has
..th prdox
bnalld of s.'' In th 1970s, rognition of this parado
spurrdsvral lading volutionary thorists to Writ hand-wringing ooks
highlightingth importan and diffiulty of this ..quenof prolms in vo.
lutionaryiology'' (Bll 1982). Though it was sy ro imgin possil n-
fitsof suality,it ws hard to s how sual individuals ould mor than
twi s fit, on avag,as asxual individuals, whih is what smsto r.
quird to ovrom th twofold ost of s. Thorists found tht sxual indi-
viduals an hav this magnitud of advantag Ovr asxual individuals only
in a world that hangs radilly from on gnratin to th nxt. spifi-
ally, s an outompt asuality only if thr is som phnomnon that
(1)kills a larg proportion of ah population, or has an quivalntly dir f-
fton rprodution; (2) is snsitivto variation twngnoms'killing in-
dividualswith rtain gnotypsand sparing thos with othr gnotyps;nd
(3)hangsradially frm lngnrationtO th nt in trms of whih gno-
typsar killd and whih ar spard. Furthr, to provid a gneral eplana-
tion for sx' th phnomnon would hv to uiquitous in th living
world, afftingmost multillular organisms.What ould this phnomnon
?In rnt dads,two major andidatshav n rognizd:inftious
disas(Hmilton 2001) and harmful muttions (Kondrashov 2001).
Th ida that inftious disass,ausd y parasitsand pathogns,ould
mak s advantagousis oftn known as th Rd Qun hypothsis.of all
th asptsof an organism'snvironmnt' its parasitsand pathogns st fit
th thorists'dsription of th phnomenon that givs an advantag to s.
Baus parasits and pathogns hav larg populations and rprodu
quikly ompard to thir hosts, thy an volv to attak partiular host
gnotyps.one striking prdition of th hyplthsis is that strong flutua-
tions should our in th frqunisof alllsafftingimmunity; alllsthat
ar ommon in on gnration should om rar in th nt. Th Rd
Qun hypothsishas n sussfulin plaining th vlutionary dynam-
is of hosts and parasits in fw wll-studid spis of snails and watr
flas,ut sintistsdo not yt know how roadly applial it will turn out
to D.
Th othr phnomnon with th potntial to mak s nssaryis harm-
ful mutation. \X/ithhigh mutation rats' s an nefiial aus som
rombinant offspring will arry fwr harmful allls than ithr parnt.
This is <lnlytru if rats of harmful mutation ar rmarkaly high, with vry
individualarrying on avragmor than on nW harmfu| mutation. o tst
th mutation thory dirtly, sintists nd to stimat how many nw
harmful mutations ppar in h individual. This is a diffiult quntity to
msur'ut stimatsar n()Wavilabl fr a fw organisms. Hurans sm
572 es

to hav a high nough rat of harmful mutation to mak sx usful, ut th altrnativ forrs of a
rat sms to too low in Worms and flis. An important dtail is that s hangs unfold in volu
only hlps to inrase avrag fitnss if hzrrmful allls intrat synrgistially, mor thorolrghly or ar
that is, if thy bom poportinatly nror hamful as mor of thm u- dation of svlras
mulat in a gnom. A waknss of th mutation thory is that it dos not T r i n d i n s t t i s t i 1l
sm to aount for ologial pattrns in th distriution of asual linags rivd his dotoral d
as wll as th Rd Qun hypothsis ds. who was a studnt of S
\Jhna gnom rpliats, it tnds to dtriorat. utations our and work on th samplirrg<
thy tnd to harmful. Th nvironmnt also tnds to gt Wors from ya hav no ffts on th fit
to yar: your nemis gt ttr at attaking you, spially parasits and opd y Motoo Kimur
..Motoo Kimura'' nd ..
pthogns with short gnration tims that volv rapidly to adapt to your
gnotyp. Ths two prosss tnd to mak mintirining stati gnom biohmial and r-nol
oth diffiult (du to mutation) as wll as a ad ida (du tl parasits).S s p i s .O n o f t h r i t i t
sms to an adaptation to dal with on or anothr of ths prolms, or al distriution of alll
prhaps to dal with oth of thm at on. Thr has n a rnt trnd to- triutions xptd und'
ward omining th Rd Qun and mtrtational mdls and rognizing wns drivd ths
tat sx has n-rultipladvantags.vn Gorg Williams, whos qust for an aout th nattlr of mut
individual-lvl advantag of sx launhd th modrn study of its volu. to alld, not Only al
tion, hs ondd that its spis-lvladvantagsmight ruial aftr all pirial data' but also p
(Williams 1992|. Asxual spissm to b muh mor pron tl xtintion thortial advans wt
than sual spisH . n , m o s t s u v i v i l l gs p i sa r s x u a l , n d m a n y , i n - utral thory nd the
luding mammals, sm tO hv faturs that prvnt thm fom vr - allls among a st of n
oming asual. Although th prolm of sx has rntly bn rdud and gn flow.
from a mystry to a mr ontrovrsy' sintistsstill nd a good dal mor Thr of wns's otl
infrrnation to undrstnd xtly how sx hlps irrdividual oganisms to i n r p r i n g h g l r i
hav mor sussfuloffspring and how it prvnts populations from going alll vailal in a popl
trnt. pndd on th prisst
nti load lays bnath
B|LIoGRAPY th so-alld ldn's
Bll, G. 7982. b strpitf Ntur.Brkly:Univrsityof lifornia Prss. ost of so many unfit g
amilton,!. D. 2001.Nrrot,tlRldsof GeLnd.ol,2,h uolutioof Se, indd nutral. wns's
oxford: Oford UnivrsiryPrss. ntiststhought aout th
Judson,o.2.002. Dr. Tti'sSduito Il rti.Nw York: tropolitan for th fftiv popult
Books.
ar rally ontributing t
K l n d r a s h vA,. s . 2 0 0 1 .S n d U . T r e d si n G e t i s1 7 : 7 5 _ 7 7 '
as of onfusion, spt
Ridly, \'4, 1994. Th Rd Que: Sd tb uolutio of Hum Na,re. Nw
York: amillan. popultion siz, a quest
\Wismnn, A. 1889. sssuplttHerditd Kidrd BiologilProblems, his <lllgusdrivd n
Ofod:lrndonPss. and human disass,m
Williams, G. . 7992.Ntrl Sltil: Domis, Luls,d hIleges.oford: asd on th nonrando
Oxford UnivrsitvPrss. -B.B.N.
wns has a lgnda
strutor.H hs inspir
'Warrn Australia, prinlarily at
wns, J. (b. 1937) Univrsity of Pnnsylva
Although thy may sm sotri at first glan' many of th stirtistial as- tistil Soity of Austr
pts of ppulation gntis'suh as th prisdistrir'rtionof fitnssamong of th Royal Soity.

)n to mak s usful' ut th
\n important dtail is that sx
I al1lsintrat synrgistia1ly,
rarmful as mor of thm au-
Ltionthory is that it dos not
jistribution of asual linags
riorat.utations our and
o tndsto gt wors from yar
i you, spially parasits and
volv rapidly to adapt to your
maintaining a stati gnom
lad ida (du to parasits). Sx
anothrof ths prolms, or
rr has n rnt trnd to-
:ionlmodls and rognizing
;.!illiams,whos qust for an
;h modrn study of its volu-
rtags might ruial aftr ll
muhmor pron to xtintion
r i s s u l ' a n d m a n y ' i n -
at prvnt thm from vr -
s has rntly n rdud
:istsstill nd a good dal mor
hlpsindividual organisms to
r v n t sp o p u l a t i o n sf r o m g o i n g
:y:Univrsityof alifornia Prss.
,d.o|.2,Tbuolutionof Se.
retio.Nw York: tropolitan
tis17,.75-77 ,
,tolutioof m Na're. Nw
'idredBiologilProblms.
, Luels,d hllges.oford:
-B.B.N.
|an,many of th statistial as-
is distriution of fitnss among
ues 57
altrnativ forms of a gn' play vital rols in dtrmining how gnti
hangs unfold in volution. Fw sintists hav unovrd ths vital rols
mor thoroughly or arefully than \WarrnJ. wens, whos work is th foun-
dtion of svral aras of rsarhin volutionary population gntis.
Traind in statistial gntis at th Australian Ntional Univrsity, wns
rivdhis dotoral dgrtn 1964 undr th suprvisin of P. A. P. Moran,
who was a studnt of Sir Ronald Fishr. wns is known primarily for his
work on th sampling distriution of nutral allls, whih ar allls that
hav no ffts on th fitnss of thir arrirs. Th nutral alll thory dvl-
opd by otoo Kimura and Tomoko Ohta (se th alphatial ntris
..Motoo
Kimura'' and ..Tomoko Ohta'' in this volum) wddd th study of
iohmial and molular polymorphisms to pattrns of variation among
spis.On of th ritial tsts of this thory ompard th osrvd statisti-
ai distriution of alll numers and frqunis at a singl gn to th dis-
triutionsptd undr nutrality. In a srisof paprs in th arly \970s,
wns drivd ths ptd distriutions undr a varity of assumptions
aout th natur of mutations. Th wns Sampling Distriution, as it am
to alld, not only allowd th nutral thory to vluatd against m-
pirial dt, ut also providd th foundation on whih svral additional
thortial dvans wr uilt. Ths inludd various rfinmnts of th
nutral thory and th dvlopmnt of mthods for using th distriution of
alllsamong a st of nighoring populations to stimatrats of migration
and gn flow.
Thr of wns's othr ontriutions stand out. First, h showd how
intrprting th gnti load, th diffrn twn th fitnss of th st
alllavailal in a population and th avragfitnssof all othr allls,d-
pndd on th pris statistial distriution of fitnss. Th onpt of th g-
nti load lays nath svral ontrovrsis in volutionary gntis, from
th so-lld Haldan's dilmma (how an an volving population ar th
ost of so many unfit gnotyps)to th liklihood that molular variation is
inddnutral. wns's ritial study of th gntiload hangd th way si-
ntiststhought about thsissus.Sond,wns drivd pris prssions
for th fftiv population siz, whih is a masur of how many individuls
ar rally ontriuting to th nt gnration. This work larifid svral ar-
as of onfusion, spially surrounding th fft of atastrophi drass in
population siz, a qustion of intrestin onsrvation iology. Third, h and
his ollagus drivd nW mthods for dtting assoiations twn gns
and human disass,mthods that solv long-standing statistial prolms
asdon th nonandom sampls that mrg from mdial rords.
w n s h a s a I g n d y p u t a t i o n s s u p r l t u ra n d l a s s r o o m i n -
strutor. H has inspid gnrations of studnts throughout his arr in
Australia, primrily at LaTro Univrsity, and in th Unitd Stats at the
Univrsityof nnsylvania.H rivdth . J. Pitman dal from th Sta-
tistialSoity of Australia ln 1996 and' in 1999,ewas ltd as a fllow
of th Royal Soity.
574 obiolog

|LIoGRAPHY
wns,!. J. 2004. tbmtilPopultiGetis.2ndd. Nw York: Springr.
.V.
wns' J., and G. R. Grant. 200|. Sttistilthodsi Bioinformfis.Nw
York: Springr. -IT

obiology
With th advnt of spa ploration, two filds of rsarh-origin.of-lif
studis nd th sarh for tratrrstrial lif-wr mldd into a singl nw
disiplin. In 1959, Nol Piz_winning miroiologist Joshua Ldrrg
oind th trm eobiolog for this nw intrdisiplinary fild, offing th
rational that on ndd to know th onditions undr whih lif,s origin
had n possil on arth in ordr to know what kind of hmial ondi-
tions to look for on othr plants. Furthrm<lr' h argud that strilization
of sparaftwas ruial for thy landd on othr plants,lst ontamina-
tion of thos worlds y arth miros or vn organi moluls from arth
rndr it impossil forvr to know with rtainty what kind f nativ or- i1
ganis or organisms had n thr originally. At first lrgly an Amrian ;.tl i,
;i:lif&
sin austh National Aronautis and Spa Administration (NASA)
I',
litr'
l-
Was th singl largstsour of funding, work in thsaras had th potntial
:t;
.s0.ar,
,iS
to b politiized th old as did all th sin rlatd to spa -
ploration. The Sovit Union was highly srtiv,for xampl, aout what
strilization standards,if any' had n applid to its intrplantarymissions
targtdfor impat or soft landing on th moon' Vnus' and ars. Similarly,
th Unitd Stats was highly srtiv aout how its prtis in strilization
was rlatd to lassifidgrm warfar rsarh.Gradually, howvr, intrna.
tional onfrns sponsord y th ommitt for Spa Rsarh
(osPAR) dvlopdintrnational sparaftstrilizationstandards,vntu-
ally inorportd into te 1967 Outr Spa Trty. Intrnationl origin-of-
lif onfrnsand a wid rang of NASA-sponsord mtingson th oigin
of lif, organi moluls in spa,Pramrian palofossils'nd othr topis
also gradually onsolidatd an intrnationalommunity of sintistsworking
on pis f th puzzl as disparat as iohmistry,gohmistry,plantary
gology, and radio astronomy. In 1968 th first oiology journal gan
..ll}':Rti
puliation (now titld rigis of Life d uolutio of tbe Biospber).In ^RI' {.R
As l,l'il-\ '
1972 exotologists foundd thir first profssional soiety, th Intrnational N0 I

Soity for Study of th origins of Lif (ISsoL). only with th thnolgy o{

spa xploration, ths sintistsforsw, would it om possil to d- Bingsfrom outrs
vlop a tuly univrsal sinof iology-to undrstand (o tst) what uni- drawing, y !7illia
on Mars,'' osmop
vrsal laws istd for iohmistry and volutionary history.
at th tim whn th
From th bginning, oiology pursud aadmi laoratory rsarh as
on ars Wrth p
wll as th dvlopmnt of spifi instrumnts to fly <rnsparaft, suh as assumthat voluti
th two U.S. Viking Mars landrs'In7976 thsraft found no lar vidn th invitlouto
for lif at th \4artian sufa ut providd nough data to suggstthat if lif
did xist on rs, it would probaly only found dp nough low th
surfato shilddfrom solar ultrvilltrays. In addition, th devlopmnt
 obiolog 575

.ls.2nd d. Nw York: Springr.

thods i Biiformtis. Nw
-1.T.

:lds of rsarh-origin-of-lif
.wr mldd into a singl nw
roiologistJoshua Ldrrg
disiplinary fild' offring th
tions undr whih lif's origin
what kind of hmial ondi.
'r,h argud that strilization
1 o t h rp l n t s 'l s t o n t a m i n a -
n organimoluls from arth
. r a i n t yw h t k i n d o f n a t i v o r -
v . A t f i r s t l r g l y n A m i n
SpaAdministration (NASA)
in thsarashad th potntial
th sinrlatd to spa x-
tiv,for xampl, aout what
. . dr o i t s i n t r p l a n t a r ym i s s i o n s
on' Vnus' and Mars. Similarly,
row its xprtis in strilization
h ' G r a d u e l l .h o w v r .i n t r n a -
,mmitt for Spa Rsarh
stri1izationstandards, vntu-
Traty.Intrnational origin-of-
o n s o r dm t i n g so n t h o r i g i n
a n p l o f o s s i I sa.n d o t h r t o p i s
:ommunityof sintistsworking
:mistry,gohmistry' p1antary
first xoiology iournal gan ''|lli
uolutio of th Bilspber).|
AR|i.RAl}; '\l.tlRs lN n1Irll llY AR l lril,\.o R!is'il''- Us' ANI)
s ',il.\. :\S Nf} |l1. \Ytt.l' t} (}vRlil) wltl ']n^llRs ()R PL,R. l Is
N{) l'ss Rn.\s(}NI}l,F: 0 ql,ll(ts' tNs'\D () A llANI}, t,
;sionalsoity,th Intrnational (iR(}(1t' {'ll }i^('I.l:s ()R R.ln{)s('ls l,| oRANs,'

) L ) .o n l y w i t h t h t
would it om possil to d-
o undrstand(or tst) what uni-
lutionaryhistory.
aadmilaoratory rsarh as
:nts to fl on sparaft, suh as
Lsraft found no lar vidn
:noughdata to suggst that if lif
found dp nough low th
ays.In addition,th dvlopmnt
h n o l o g yo f
Beingsfrom outr spa hav long n th stuff of fantasy and nightmr.This
.!lls,..Th
drawing,y !7illiam R. Ligh, appardin H. G. Things That Live
on ars'', osmopolitgzie,Marh 1908, 35_343.It appardjust
..anals''
at th tim whn thrws muh popular disussionout whthrth
on rs Wrth produtsof ral intlligns.Not how oftn suh piturs
assumtht volutionis progrssivnd nonontingnt,with humanlikforms
th invitaloutomof th pross,anywhrand vrywher.
576 obiolog

of radio astronomy thniqus to sarh for signals from intllignt tratr. in th dirtion of iol.
rstrials (STI) was part of oiology fronr th outst, although ongrss faily quikly ut ..nith
ut off NASA funding for STI wok in 1992. B th 1980s and 1990s, nw stats(2000, 109).Il.s
astronomial thniqus wr dvlopd to sarh for plants of stars ynd arly in lif's history wl
th solar systm' with th first suh plant ing dttd in 1995. Laora- ating as oth an inforr
tory work has pusud numrous goals, inluding study of th organi om- lism.
pounds fond i metorits, turnd moon oks, and (via sparaft)
omts' and study of anint roks for fossilizd miros nd/or organi |BLIoGRAPHY
moluls that might ssoiatd with on-living organisms. This work Dik' S., and J. . Strik.2
has ld to pushing bak to 3 illion yars or mor th oldst known fossil mi- strobiolog.Nu, Br
ros from arth, narrowing th tim window fo th pross of hmial Fry, I. 2000. The mrg
uruit'l,,. Nw BruIl
volution to hav ourrd. Howvr, it has simultanously rva[dthat
Grhurm,L. 1987,Si
arth is ing bomardd y organi moluls formd in hug irmounts in Nw York: olurnr
intrstllar spa' so that th original prursors of lif on arth nd not all zn, R. 2005. Gesis:
hav bn synthsizd hr from purly inorgani starting matrials. oiol- Hnry Prss.
ogy rsarh has also spun off numrous othr important disovris: Lynn L d r gJ ., l 9 6 0 . h i
argulis's thory, now univrsally aptd, that ukyoti lls aros via 940.
.Wos's L v l o kI, . 1 9 7 9 . G i :
formation of srial symbioti olonis of pokaryoti lls; Carl dis-
Prss.
ovry of th Arhaa' a grop of atria as diffrnt from othr tria as Mllgu|is.L. |970.origin
thy ar from ukaryots; Jams Lovlok's Gaia hypothsis, whih laims Prss.
that all lif on arth ats as a oordinated homostati systm maintining i I l r 's . I 9 5 3 .A p r o d t r
onditions within th limits lif n tolrat; th thory that a hug stroid onditions.Sie|1
o p i n , A . l 9 3 8 . T h o r i g
impat at th nd of th rtaous priod was rsponsilfor th mss - 'W.s,
. 1987.Btrialt
tintion that inludd th dinosaurs; and th ..nular wintr'' hypothsis, Wolf,A. 2002. Grmsin s
pditing that a similar mass tintion ould follow a nular War of any imagination,1958-19
signifiant siz, and spurring on muh of global limat modling sin.
Th brodning of th rsarh agnda rsultd in th xoiology fild ing
rnamd astroiology in 1996.
Following th Soviet biohmist Alksandr Oparin, origin-of-lif rsearh
assumd in th 1950s that arth's primitiv atmosphr must hav n
hmially rduing' that is, it ontaind no fr oygn' arbon mostly s
mthan, and nitrogn as ammonia. Th 1953 Ury-Millr xprimnt
showd that Spontnous synthsis of amino aids and othr rlvant io-
moluls ould our rapidly undr suh onditions, vn assuming rzrndom
hmistry. By th lat 1970s, howvr, xoiology rsarhsuggstdarth's
arly atmosphr was mor likly to hav n hmially nutral (or pr-
haps vn slightly oidizing), undr whih onditions th synthsisof organi
moluls is drastially rdud. Altrnativ pathways of hmial volution
ar undr invstigation, inluding synthsis of organi polymrs using lays
as tmpltsand talysts;high-tmpratLlr, high-pssurhmistry suh as
might our at undrsa hydrothmal vnts, whr many rsarhrs now
think lif may have first originatd; and omining mtorit-drivd organ-
is to s what kinds of liflik protolls will produd. It is now widly
blivd that th hmistry tht ll frm inorgani prursors to tlr first
biopolymrs ours nonrandomly' mor rapidly, and spontanouslyundr
widr rans of onditions than was livdin th 1950s and 1960s' Biasd
 obiolog 577

]nalsfrom intllignt xtratr-
:h outst,althoug ongrss
By th 1980sand 1990s, nw
h for plantsof stars byond
ing dttdin |995. Laora-
t i n gs t u d yo f t h o r g a n i m -
t roks, and (via sparaft)
i z d m i r o s n d / o r o g a n i
-living organisms' This work
r th oldst known fossil mi-
w for th pross of hmial
simuitanouslyvald tht
s f o m di n h u g a m o n t s i n
,rsof lif on arth nd not all
ri startingmatrils.xobiol-
:r important disovris: Lynn
t h a t u k a y o t i l l s a o s v i a
aryoti lls; arl Wos's dis_
Iiffrntfom oth atri s
aia hypothsis,whih laims
l m o s t r i s y s t m r n i n a i n i n g
h thory that a hug astroid
rs rsponsilfor th mass x-
..nularwintr'' hypothsis,
d follow a nular war of any
l a l l i m a t m o d I i n g s i n
ld in th oiology fild ing
in th dirtion of iologially rlvant moluls' lif proaly originatd
fairly quikly ut ..nithy han nor dsign,''as sinhistorian Iris Fry
stts(2000, 109). Rsarh has also foused on th RNA world, period
arly in lif's history whn RNA istd for protins ut was apal of
ating as oth n information-arying rnolul and a atalyst of mtao-
lism.
BlBLIoGRAPY
Dik' S.' and J. . Strik.2004. The Liuig Uniurse:NASA d th Deuelopmentof
strobiolog.Nlv Brunswik'NJ: RutgrsUniversityPrss.
Fry, I. 2000. Th mrgof Life ln the rth: A Historil d Scientifi
ouruitll.Nw Brtrnswik,NJ: RutgrsUnivrsityPrss.
Grham'L. |987. Sitl,Philosophd llm Bhuiori tb Souit IJnion.
Nw York: olumbiUnivrsityPrss.
Hazn,R. 2005. Gsis:he SitificQust flr Lif's rigins. Nw York: Josph
nr Prss.
Ldrrg, J. 1960.oiology:Approahsto lif byondth arth.Si|32:
93400.
Lovlok,J. 1979. Gi: Ntu l-ooh t Lif o I,rtb.London: Oford University
Prss.
argulis' L. 1970. rigi of ukroti elis.Nw avn' T: Yal Univrsity
Prss.
illr,S. 19.5.A produtiorr of minoiridsurrdrpossiblprimitivarth
onditins.Sie||7 : 528*529.
oparin,A. 1938.Tlrc rigi of Lfe. Nw York: Mamilln.
.Wos,
. 19 87. Btrial volution.i cro bi ol lgi l Ruit't-t
s 5 1': 221'-271'.
\(olf,A. 2002. Grms in spa:Amrin lif sintists,spapoliy, and puli
imagination, 19 58-1964.lsis 93: 183-205. -/..s.
.

oparin, origin-of.lif rsarh

l atmosphr mllst hav n
fr oxygn, aron mostly as
l953 Ury-illrxprimnt
aids and othr lvant io-
ditions, vn assuming random
'logy rsarh suggstd arth's
:nhmially nrrtral (or pr-
d i t i o n st h s y n t h s i so f o r g , n i
,thwaysof hmial volution
f organi polyrnrs using lays
righ-prssur hemistry suh as
, whr many rsarhrs now
lining mtorit-drivdorgan-
[ produd. It is now widly
organi prursors to th first
lly, and spontanouslyundr a
n t h 1 9 5 0 s n d 1 9 6 0 s . B i a s d
 alld indpndnttl
for omparativ ioll
populations of th sal
F dsndant rlationsh
long-trm rats of grr
larg th populatilns
ollagus hv br:n
and som of ts I
iology.
In addition to ths
Josph(. t942)
Flsnstin, thi1t xplord th vol
Although all sintistsaspir to l-rilnghow thir ollgusthink alut an tion mong llls at
issu, only fw manag to do so. A slt fw hang thir ollagus' whih tightr :rnd lo
thinking on mo than on issu.Josph Flsnstinhas hangd th thinking tions) volv and pl
of volutionary iologists on at last thr ritial issus. prosss.Linkag a
A studnt of plpulation gntiist Rihard Lwontin, Flsnstin wrot his lution of sxual rpr
dissrtation on mthods for infrring phylognis' riving his dotoral d- in this volum) and i
..Th Pattrn and Pro
gr in 1968 from th Univrsity of hiago' Aft postdotoal work at d_
inurgh with Alan Rortson, h dvotd th :rrly yars of his ar to Hankinson in this vol
population gntis;his dissrtationrsarhdid rrot appar in sintifijour- |n 1'992Flsnsti
nals until 1'97. 'fter this rfratory priod' h rturnd to th study f n s .I n 1 9 9 3 h w a s
phylognti mthods and has mad sminal ontriutions to phylogntis Arrian Soity ot }
and population gntis'along th W^ypromoting a synthsisof th two. ti<lnalAdmy of S
Flsnstin pionrd a sttistial approalr to phylognti infrn (s
..Phylogntis'' in this volum). H rognizd that, BIBLIOGRAPH
th alphatial ntry
whn infrring a phylogny' not only would th lngths of th branhs Flsnstin, l. 2004.If
subjt to sampling rror, ut so would th atual shap of th ntir tr. S l a t k i n , . 1 9 9 5 . h S
Ntrlist145:ii-r
Sampling rror introdusunrtaintyaout th idntity of los rlativsand
rnt ommon anstors'whih in turn ratsunrtinty aout th ordr
of pparan of major faturs in groups lik birds or fish. Nitlrr tradi-
Fishr, Ronald A
tional mthods Iror th mrging ladisti mthods onfrontd this polm
satisfatorily. Flsnstin introdud th statistial thniqu of maimum Sir Ronald Aylmr I
liklihood to th study of phylognis. Th initial rlutan of volr-rtionary gl:rnd. Fishr and his
biologists to mra th thniqt1Was ovrom whn Flsnstinshowd of ight hildrn
that ladisti mthods would point to th wrong phylogny wlrnvr diffr- ]athmatial gifts a
nt branhs of an volutionary tr priend suffiintly diffrnt rats of lriology. Baus of
haratr volution. This rsult has om a ttook xampl, and mai- Indd' som of his n
mum liklihood arrd rlatd mthods hav om th standard approahs or papr' giving Fish
for undrstanding volutionary rl:ltionships. his had.
Biologists a now fllwing Flsnstin'slad on two oth topis that Fislrr matriLllat
mrgd from his synthsisof phylogntisand population gntis.In th H g r a d u a t 6; n | 9 1
|ate 1.97s,svl volutionary iologists pointd out that traditional om- ol for anothr yr t(
parisons of haratr ssoiations among spis ould misintrprtd sts wr road' inlt
baus suh omparisons did not rogniz th phylognti rltionships | 9 | 4 h e a m s t i
among th spising ompard. Flsnstindvlopda pwrful nrthod pany in London. on
for intgrtingphylogntiaffinitisinto ompirativanalyss.His mthod, aus of his potlr r

s78
 Fisher 579

alldindpndntontrasts,and its dsndantshav om standard tools

for omparativ iology. In th arly 1990s, Flsnstin pointd out that
po;lulationsof th sarl spiesar prt of a ompl pdigr of anstor_
dsndantrlationships and that, through this pdigr,on ould stimat
long-trmrats of gnti hng among populations and how gntially
larg th populations hav n. thods dvlopd y Flsnstinand his
ollagushav n applid to svral prolms in volutionary biology,
and som of ts mthods hv om important tools in onsrvation
iology.
In addition to ths aomplislrmnts,Flsnstinwrot a sris of paprs
that plord th volution of gntilinkag, whih is th statistialassoia.
thir ollagusthink aout an tion among allls t diffrnt gns. amind th irumstansundr
:t fw harrgthir ollagus' whih tightr and loosr linkag (strongr nd wkr statistil assoia-
nstinhas hangd th thinking tions)volv:rndplord ho''v los linkag afftsa varity of volutionary
tial issus. prosss.Linkg mong gnsis vital lmntin plantions for th vo-
Lwontin, Flsnstin wrot his lution of sual prodution (sth alphtial entry ..volution of sx''
nis'rivinghis dotoral d- in tlris volum) nd it plays a ruial rol in spiation (s th main ssy
..Tlr
Aftr postdotoral work at d- Pattrn and Pross f Spition'' by Margart B. tak and Shla J.
:h arly yars of his arr to Hirnkinson in tlris volum).
did not ppar in sintifi jour- |n 1992 F.lsrrstin
was ltdto th Amrin Aadmy of Arts and Si-
d' h rturnd to th study of ns.In 7993 he was th sond ripint of the Swall Wright Award of th
l o n t r i l r t r t i o nso p h l o g n t i s Amrian Soity of Naturalists, and tn 1999 h was ltdto th U.S. Na.
roting a synthsisof th two. tional Aadmy of Sins.
h to plrylognti infrn (s
BIBLIOGRAPHY
is volum).H rognizd that,
l th lngths of th ranhs b Flsnstin,
J.2004. lfrrigPhIlgies. A: SinaurAssoiats.
Sundrland,
Slatkin,M. 199.5.Th SwallWrightAward:JosphFlsnstin.
Ameri
aual shap of th ntir tr. -I.T'
Ntltrlist145:ii-v'
th idntitof los rlativs and
ats unrtaintyaot th ordr
lik irds or fish. Nithr tradi- Fishr,Ronald Aylmr (1890_962)
rthods onfrontd this problm
t i s t i a lt h n i q u o f m a x i m u m Sir Ronald Aylmr Fishr was orn Fruary 17, 1890, in London, n-
initial rlutan of volutionary gland. Fishr and his twin rothr, who did in irrfany, Wf th youngst
)rol1-l whn Flsnstin showd of ight hildrn orn tr Kati H:rth and Gorg Fishr. Showing his
rong phylogny whnvr diffr- mathmatialgiits rly, Fishr would hang rh f of oth statistisand
rd suffiintly diffrnt rats of iology. Baus of xtrm myopia, his arly trining Was unonvntional.
a txtook ampl, and maxi. Indd,som of his most influntial arly tutring was don without pnil
omth standard approahs or Papr, giving Fish a rmarkirbl aility to do ompl mthmatis in
hishad.
's lad on two othr topis that Fishr matriultd at Gorrvill and aius ollg, amridg, in 1909.
s and population gntis. In th H graduatdin |972 with high honors in mathmatis,and he would stay
lointd out that traditional om- on for anothr yar to study physis and statistialmhanis.Fishr's intr-
spisould b misintrprtd stsWr road, irrludingastronomy' mathematis,physis, and iology. In
z th plrylogntirlationships 7914 e am statistiianat th Mrntil and Gnral Invstmntom.
in dvllpda powrful mthod pany in London. ontrary to lris dsirs, Fishr was kpt out of \World !ar I
)mprativanalyss.His mthod' ausof his poor visiorr. Btr,n1915 and l919 h taught mathmatis
580 Fisber

and physis at a varity of puli shools. |n 1917 h mrrid Ruth iln ulturl, d edi
Gatton Guinnss, with whom h would parnt ight hildrn. On of Fishr,s agriultural rsar
si daughtrs,Joan Fishr Bo, authord th most wll.known iography of fild's onptual f
him in 1978. lightd mainly y l
At Rothamstd xprimntal Station' twn I9I9 and 1933, Fishr good ompany in -
mad substantial ontriutions to statistisand gntis.In statistis,h in- omplishmnts did
trodud th onpt of liklihood (Fishr 1921). Th liklihood of a param- Royal Soity in 19
tr is proportional to th proaility of the data and it givs a funtion that 1955.In 1943 Fish
usually has a singl maximum valu, that is, th maximum liklihood. And ridg Univrsity; i
th nxt yar Fishr introdud a nW onption of sttistis,th aim of lg' amridg; a
whih was th rdution of data and th prolmati of whih Was th sp- lizath II. Numl
ifiation of th kind of population from whih th data am, stimation, and Aftr a luminous
distriution. Soorr thraftr, Fishr pulishd SttistiI ethods for R- of 72 onJu| 29, \|
serh Worhers (1925), in whih h artiulated mthods for th dsign and Fishr has en ds
valuation of primnts. Th 1925 dition was th first of many. In gntis, volatil tmpr. Fis
Fishr's first papr (in 1922| adumratd a mathmatial synthsisof Drwin- to his viws, drov
ian natural sltion with th ntly rdisovrdlaws of ndlian hrdity. writings hav bn
That, and othr work (.g.,on th volution of dminan), would ulminat mor fftivly or
in The Getil Theor of Ntrl Sletil (1930), on of th prinipal tion and his gnius
txts' along with thos of J. B. S. Haldan (1892-1964) and Swall Wright
(1889_1988),omplting th roniliationof Darwinism and Mendlism and BIBLIOGRAP
stalishing th fild of thortial populatin gntis and its appliation to B o x ,J . F . 1 9 7 8 .R . .
ugnis.Rmarkably,Th Gnticl Theor of Nturl Seletk was ditatd Fishr,R. A. 1921.o
y Fishr to his wif, Ruth, during th vnings.Th book was rvisd and ris- frm small sar
1922,n t
sud in 1958. In 1929 Ftsher was ltd to th Royal Soity.
diburgh42:3
Fishr lft Rothamstd to tak th Galton hair of ugnis at Univrsity 1925.Sttist
ollg, London, in 1933. Karl Parson (1857_196) was rtiring from th 1 9 3 0[ 1 9 s 8
hair' whih was thn dividd into two, so that oth Fishr and Parson's York: Dovr Pu
son, gon, ould b appointd. Fishr ontrolld th gntisstion; Par- 1935.h D.
'
son ontrolld th statistisStion.(Fishr had turnd down Parson'soffr Fishr,R. A., and F.
dil Rsr
of hif statistiianat Galton Laoratoris for th position at Rothamstd in
791'9.).Whnh took th Galton profssorshipt Univrsity Collg' h was
thoroughly mird in ontrovrsy ovr th foundtions of statistis with Karl
Fishs
Parson and his followrs. Fishr's rvolutionary work in statistisam at
around th tim that Parson's own wok was showing waknsss; un- Th word fisbes ls a
frindly omptition cata|zed th ontrovrsy, and Parson would tak ill hold. Most popl 1
flings toward Fishr, who had his own ill flings toward Parson' to his fs is usd, but thil
grav. Th ontrovrsy in sttistiswith Parson would not th only on- ths too ar fishs
trovrsy in whih Fishr was involvd. Bginning t 1929, Fishr and Amri- moving y ithr (
an physiologial gntiist Swall \/rightwould ngag in an animus-filld and/or unpaird far
ontrovrsy ovr th gnti asis of volutionary hang that would last un- gn (and rlassnl
til Fishr's deat ln 7962. gills'' gos a long ''
In 1935 Fishr's The Desig of perimezls appard and, lik SttistiI onsidrd fishs(L
ethods for Resrh Workrs' would pandd and rissud many living typs of fish
tims. Ths two works, and Fishr's Sttistil bls for Biologil, gri- of known fossil for

n 1917 h marrid Ruth iln
nt ighthildrn. On of Fishr's
r most Wll.known iography of
twn 1979 and 1933, Fishr
and gntis.In statistis, h in-
) 2 l ) . T h l i k l i h t l o do f a p r m -
data and it givs a funtion that
;, th taximurn liklihod. And
Lptionof statistis,th aim of
llmtiof whi was th sp_ lizabth II. Numrous othr honors wr stowd upon him'
:h th dta am' stimatiln,and
hed Sttistil ethods for Re-
ated mthods for th dsigrr and
W a st h f i r s to f r a n y .I n g n t i s '
rathmatialsynthsis of Darwin-
vrdlaws of ndlian hrdity.
of dorrrinan).would ulminat
ion (19\, on of th prinipal
(1892-1964) ad Swall Wright
f Darrvinism and ndlism and
on gntisand its appliation to
l lf Nturl Sltion was ditatd
gs.Th ook ws rvisd and ris-
th Royal Soity.
n hai of trgnisat Univrsity
857-196) was rtiring from th
;o that oth Fishr and Parson's
trolld th gntis stion; Par-
had turnd down Parson's offr
for th position t Rotharnstd in
ship at Univrsity ollg' h was
oundations of statistis with Karl
ionary work in statistis am at
.k ws showing waknsss; un-
.rsy, and Parson would tak ill
ll flingstowrd Parsorl. to his
,rsonwuld not th only on-
inning in 1929,Fiser and Amri-
would rrgagin an animus_filld
.ionaryhngthat would last un-
ts appardand, |Ike Sttistil
b xpandd rrd rissud many
istil Tbles for Biologil, Agri-
Fishs ''81
ultrl, d dil Reserh (with Frank Yats, 1938) volutionizd
agriultural rsarh. Fishr's work in statistis was rvolutionary at th
fild's onptual foundations. orover, Fishr work in gntis, high-
lightd mainly Th Geticl Thor of Nturl Seltiott' would, with
good ompany in Haldan and Wright, rvolutioniz iology. Fishr's a-
omplishmntsdid not go unrognizd. ws awardd th mdal of th
Royal Soity in 1938, th Dwin mdal in \948, and th oply mdal in
1955. In 1943 Fishr was appointed Balfour Profssor of Gntis at am-
ridg Univrsity; in 7957 he m prsidnt of Gonvill nd Caius ol-
lg, Camridg; and in |952 h was ratd Knight Bahlor y Qun
Aftr luminous ar as statistiianand iologist, Fishr did at th g
of 72 on Ju| 29, \962, in Adlaid, Australia, whr h had rtird i \9 59 .
Fishr has n dsridas harmig and Warm among frinds, ut with a
volatil tmpr. Fishr's tmpr' ombind with an unwavring ommitmnt
to his viws, drov him to hatd ontrovrsis with othr sintists. Fishr's
writings hav n dsrid as diffiult; muh of wlrat h ontriutd was
mor fftivlyonvyd y othrs mor apal of simplifying his prsnta-
tion and his gnius.
lLIoGRAPY
-b
Bo,J. F. 1978. R, . Fishr: Lif of Sietist.Nw Yrk: \Wily.
Fishr,R. ^. |921. on th ,.prlblrlr''f a offiintof orrlrtin
ddud
fronr small smpl.rt<l113_32.
|922. on th dominanratio. Prledigs lf th Rol Soitof
d i n b u r g h4 2 : 3 2 | - 4 1 ,
-]'925.
Sttisticlt|lodsflr RserhW<lrkrs. L<lndon:olivr and Boyd.
1930 [1958l.h GeetilThorof Nturl Sltio.2ndd.Nw
York: DovrPubliarions.
1'935.Te Dsig rlf primertts.London: Olivr and Bod.
Fishr'R. A., and F. Yats.\938' Sttistilblesflr Biologil' griultrl,d
-R..s.
edil Reserh. London: Olivr and Boyd.
Fishs
Th word fishs is aout s slippry as th oganisms to whih it rfrs r to
hold. ost popl gnrallygt a rathr similar mntal imag whn th trm
fs is usd, ut this rrly inluds thinking of ls or vn sharks, although
thstoo r fishs.Dfining a fish as ..a wtr-dwlling' akond animal
moving by ithr (or oth) latral ody undulations or osilltions of paird
ad/orunpaird fan-lik ody appndagsalld fins and whih otains oxy-
gn (ilnd rlassnmrous dissolvd wasts)vi spializdstuturslld
gills,' gos a long Way toward inluding all organisms that r sintifially
onsiderdfishs(Hlfmrrt al. 79971'Polg t al. 2005). So divrs ar th
living typs of fishs,howvr, rht this dfinition is not lzrrut. Irrlusiorr
of known fossil forms furthr strths yond auray almlst any attmpt
'82 ishs

at a omprhnsivdfinition. Tw rasons Ount for th nar impossiil- nutral uoyany r

ity of larly dfining what sms at first glan to a straightforward noun. rsponss of th vr
First, a numr of distint volutionary iinags,alit vidntilly drivd Wtr' the sparat
from ommon anstor, surviv today having normously divrgd in al- ihthys hav oftn
most all rspts through th influn of naturl sltion. Sond, fishs adaptation. For xa
(th plural applis to mor thn on spis,th singular to individuals of a than-watr odis,
singl spisno mattr how many of thm) ar th most iodivrsof living provid uyany. (
akond animals (vrtrats).xtant (living) fishs numr no lss than pokting of th s
27,000 namd spis,omprising aout 50% of all xtant vrtrats;thy ns dmonstrat t]
ar adaptd to inhait th nrly 71" <lf th arth's surfa ovrd y variation.
ons and sas,as wll as th additional 1 to 2Y" of th plant's surfa o- Th xtant hon
upid y laks, rivrs, and othr fshwatr and stuarin nvironmnts. typial shark is a<
F'ishsar found vrywhrthr is surfawatr' and oftn in sutrranan gnom indiats th
watrs as wll; thy vary in morphologial' physiologial, and havioral vnd Ostihthy
haratristisfrom volutionary adaptations to th widly varial ondi- spis in xistn
tions of th arth'sWatrs. siz, ut thy ar gt
Four vry distintiv fish linags (lads),plus anothr usually, if rlu. ntimtr in adult l
tantly, onsidrd ..fishlik,''ar tant today. A dozn, prhaps onsidr- low thm to thriv i:
ably mor, qually distintiv lingsar known only from fossils, som of ially in frshwatr
whih tnd k at last 480 rnillin yars to ony fragmntsfrom th Or- frshwatr haitats
doviian, and possily vn from imprssions in roks frlm th arly am. whil 58% of livinE
rian, anothr 40 million yars arlir. Th living linags ar th fishlik xtnt) is a mor
lants (phalohordats);th nar vrtrat ut konlsshagfishs dd som frshwat
(yiniforms), with 50 or mor living spis;th jawlss and truly vrt- Th ihlids of Sout
brt lamprys (Ptromyzontiforms),with aout 40 living spis;th arti- ios, with litrally
lginous sharks, skts' and rays (hondrihthys)'with at last 880 living svral gologially
spis;and th ovrwhlming majority of xtant spis, th ony fishs siv parntal ar of
(ostihthys),with varying stimatsof th numr of living spissom. imprintd on thir
wh around 27,000. Th living hagfishsand lamprys shar a ommon thir parnts in ht
primitiv trait with many fssil linagsin tht thy lak jaws and paid fins. rapid gnti isolati
Thy fd y pinhing or grasping with hrny tth loatd around thir oral Th random patt
opning. Thy us unpaird rionlik fins in th midlins of thir l-lik is also aundantly <
sallss odis to ontrol movmnt. Th hondrihthys and Ostihthys th vast group knov
not only hav wll-dvlopdjaws, usually rimmd y hardr-than-onmin- atfishs, lantrn an
ralizd teth, ut also paird fins that prmit normous thr-dimnsional almost all oral r
ontrol of movmnt. Dtaild study of fossils of oth groups shows similar Tlosts first appa
trnds through tim in th adaptations of th two lads to th natural sl- uniqu uniting fat
tion dmands lf watr as a mdium in whih to mov' to fd, and to avoid port of a portion of
prdation. Fins om mor flxil and al to assumvarying anglsof at- lar adaptiv adv
tak rlativ to th flow lf dns visous watr ()vr th fishs' ldis.Jaws on linag has
and assoiatdstrutursom lolsndfrom th rst of th had sklton dom and han na
so that thy may protrud toward food itms, and th oral-ranhial avity
oms inrasingly flxil sO that it may pandd to rt sution
that drws Watr' and th food it ontains, into th mouth. As fins and hy- |BLloGRAP
drodynami ody shap povid for inrasdagility, havy trnal armor FishBas:A GloblIr
of sals, spins, and had ons is thinnd or lost altogthr'making nar .htm.

:ountfor th nar impossiil-
t straightforwadnoun.
s, alit vidntially drivd
g normously divrgd in al_
url sltion.Sond, fishes
to individuals f a
r singr.rlar
] th most iodivrs of living
g) fishsnumr no lss than
of al[ -tantvrtrts;thy
l arth's surfa ovrd by
2olotlf t pln.ssura o-
nd stuinnvironmnts.
ttr'and oftn in sutrrann
physiologial,and havioral
to th lvidly vari:r1ondi-
plus an<lthrusually, if rlu-
r. A dzn' prhatrlsonsider-
lwn lnl frorrrfossils' som of
l ony fragmntsfrom th or-
in roks from th arly am-
living lingsar th fishlik
rt ut akonlsshagfislrs
:s; th jawlss and truly vrt-
, l u t4 0 l i v i n g s p i s ;t h e t i -
h y s )w, ith at last 880 living
)xtant spis, th ony {isl.rs
numr of living speis som.
rrrdlmprs shr a ommon
t thy lk jaws and paird fins.
tthloatdaound thir oral
n th midlins of thi l_lik
rondrilrthysand Ostihtl-rys
l m d y h r d r . t h n - o nm
in-
Litnornrous thr-dimnsional
ls of oth groups slrows similar
two ladsto th natural sl-
tto mov, to fd, and to avoid
: to assumvrying angls of at-
rovr th fishs' odis. Jaws
m th st of th lrad sklton
s, and tlr lral-ranhialavity
xpndd to rat sution
nto th mouth. As fins and hy-
:d agilitv, havy trnal armor
o l < l st I t t l g t h rr,n k i n g ,n a
Fishs 58
nutral uoyany rlativly sir to ahiv. Although dmonstrating similar
sponssof th vrtrt ody to th natural sltion dmands of living in
Watr' th sparat volutionary lads of th Chondrihthys and ost-
ihthys hav oftn usd vry diffrnt mhanisms to ahiv a ommon
daptation' For xarpl' to ovrom th fft of gravtt on thir dnsr-
than-wtr odis, hondrihthys hav volvd lag' oil.rih livrs that
provid uoyany. Ostihthys hav modifid an axiliary spiratory out-
pokting of th sophagus into a gas-filld uoyany laddr. Suh diffr-
nsdmonstrat th random natur of adaptation asd on han gnom
variation.
Th xtant hondrihthys r flsh atrs and gnrally larg organisms. A
typil shark is aout 2 mtrs long. vidn from analysis of th shrk
gnom indiats that thy hv volvd at a muh slowr rat than hav ad.
vand Osteihthys. This may aount for th rlativly small numr of
spisin istn today. Living ostihthys ar highly varial in dit and
siz, ut thy ar gnrally sallr than sharks' with som ing lss than a
ntimtrin adlt maimum lngth. Thir ditary radth and small siz al-
low thm to thriv in many mor haitats than any othr lad of fishs, sp-
ially in fshwatr nvironmnts. Forty.on prnt of all fish spis liv in
frshwatrhaitats, whih ar highly dividd and isolatd on from anothr'
whil 58% of living spis our in th sa, whih (in spit of its normous
tnt) is mor ontinuous hitat than th piolrtinntl fshwtrs. In-
dd som frshwatr ony fishs show astoundingly high rats of voltion.
Th ihlids of South Amria, Afria, and Suthrn India ar spially sp-
ios, with litrally hundrds of spis having volvd in situ in ah of th
svralgologially young Afrian Rift Vally laks. Baus thy hav xtn-
siv parrrtalar of ggs and young' ihlid offspring an om haviorally
imprintd on thir Parnts nd show a prfrn for individals similar to
thir parnts in hoosing thir own mats (assotativ mating), rsulting in
rapid grrtiisolation and sltion lading to nw spis.
Th random pattrn of volution without dirtion or lar sltivvalus
is also aundantly dmonstratd y fishs. Almost ll ostihthys long to
th vast group known as Tlosts, inluding ls, hrrings, salmons, minnows,
atfishs,lntrn and othr dp-safishs,ods, anglrfishs,basss,phs,
almost ll oral rf fishs' and so on. Thr ar 25,000 or mor spis.
Tlostsfirst appad r.rt200 million yars ago in th mid-Tiassi. Thir
uniqu uniting fatur is struturally rthr mino shift in th intrnal sup-
port of a portion of th tail (audal) fin that dos nor appar to giv thm any
lar adaptiv advantag ov othr ontmporary fishs. Nvrthlss, this
on lin:rghas n th gnsisof most fishsliving today, illustrating th ran-
dom and han natur of volution.
B|LIoGRAPHY
FishBas:A Gloal In{ormationSystmon Fishs.http://www.fishas.org/hom
.htm.
584 Fith

Hlfman, G. s., B. B. Colltt,and D. . Fay. 1997.Tb Diursitof Fiss, th tr showd th

ldn, A: Blakwll Sin. tans. A ritial fa
Maisy, I. G. 1996.DisourigFossilFishes.Nw York: Hnry Holt and
f th fat that gr
ompany.
Pough'F. H., . M. Janis,and J. B. isr.205.Vrtbrte \/hil th algorithm
Lif.7t d.PartsI and
, 1-156.Uppr SaddlRivr,NJ: Prntiall. -r.B.. fadd from us, dist
proah for ronstru
.Waltr last squrs has rr
Fith, rv[.(b. 1929)
Margoliash mthod i
Philosophr Isaiah Brlin (1953) haratrizdintlltualndavor as ing thir papr, basd on
ondutd y ithr hdghogs or fos. Hdghogs spend thir livs fous- givn what w know
ing intntly on a singl ohrnt issu, whil foxs onsidr numrous' at on of th most i
tims vn unrltd' prolms. For th most part' this dihotomy holds up twn so-alld lrth
wll for sintifi inquiry, ut thr ar notal xptions who upst th usd thm to distingu
.Walter
rul. . Fith, who managd to mak lasting ontriutions to th spiation vnts (or
fild of molular volution, is on suh ption. In partiula, Fith's a- (paralogs).This disti
rr has n distinguishd y groundraking advans in mthodology, th studis ut also for tL
kind of ffort that rquirs th digging aility of hdghog, as wll as fun- 1990s, th ortholog-
damntal onptual dvans aross a rang of topis, ahivmnts that n- tool for invstigato
ssitat th mntal agility of a fo. qund gnoms.T|
Traind as a iohmist, Fith has spnt th ttr part of his arr at th Sam' or vry simila
Univrsity of !isonsin and latr at th Univrsity of California at lrvin. hav divrgd in fun
!7hen h gan his omparativ rsarh on protin squns' th notion of omparisons an os
molular volution was littl mor than a fantasy;h is as muh rsponsil onflation of ortholo1
as anyon for th mrgn and dvlopmnt of this nsnt rsarh pro- omparison approa
gram into a major, lgitimat, and distint disiplin at th nxus of volu- pross y disrimin
tionary and molular iology. Togthr with asatoshi Ni, anothr (2000) allow for mor
luminary of molular volution, Fith foundd th Soity for Molular Bi. voiutionay iolo
ology and volution and its journal, oleulr Bhlog d uolution, lutionists ar prsn
whih rmains th prmir journal in th fild and is on of th highst- past vnts asd sol
ranking journals in th volutionary iology fild. Th ontriutions that sqrrnsof tant
Fith has mad ar numrous' so a mr sampl of what Fith has givn to suh as th fossils uti
molular volution is prsntd y fusing on his work in thr distint ar- y historians' moiu
as:mthods' onpts' and applitions. of tools that provid
A rowning ahivmnt of molular volution is th us of phylognti adpt' howvr, ao
mthods to ronstrut th volutionary histois of gns, protins, and, y vnts. In an audai
tnsion' spis. Whn rlatd protin squnes gan to aumulat in to dfy this trnd y
th lat 1950s and rly 1960s, thr wr no mthods availal to ana|ze fort did not rprsn
thm from an volutionary stndpoint. Numrial taxonomy mthods had influnza virus, and 1
only just n formalizd into algorithms, ut ths approahs ould not pidmis. Suh a pr
radily applid to sequn data. \Working with manul argoliash' Fith flunza strains to us
publishd a sminal papr in 1967 tat rportd th first distan-asd on a phylognetiap
mthod for phylognti ronstrution of protin squns.Distans wr nomi squnsto t
omputd as th proportion of amino aid diffrns twn all pairs of s- fstd itslf with th
quns undr onsidration. Th Fith'Margoliash mthod amountd to a that rprsntdth
lvr implmntation of th last squars statistial prinipl, whry a natd pidmis fror
phylogny was ronstrutdsuh that th distans along th ranhes of squn'Fith and l

7, h Diursit-lfFishs.
York:HnryHolt and
,rtbrt
Life.7th d. Parts I and
11.
intlltualndavor as ing
;hogs spnd thir livs fous-
: foxs nstirnumrous' at
part, this dihotomy holds up
Llptions who upst th
l lasting ontriutions to th
ption. In partiular, Fith's a-
; advansin mthodology, th
of a hdghog, as wll as fun-
lf topis,ahivmntsthat n-
: ttr part of his arr at th
vrsity of California t lrvin.
lrotinsquns,th notion of
ltasy;h is as muh rsponsil
rt of this nasnt rsarh pro-
isiplin at th nxus of volu-
with Masatoshi Ni, anothr
d th Soity for Molular Bi-
'ulr Biolog d uoltion,
ld and is on of th highst-
y fild. Th ontriutions that
lpl of what Fith has givn to
o n h i s w o r k i n t h d i s t i n t a r -
ution is th us of phylognti
lris of gns'protins, and, y
uns bgan to aumulat in
Lomthods availal to ana\ze
nrialtaonomy mthods had
t thsapproahsould not b
uith manul argoliash, Fith
portd th first distan-asd
'otinsquns.Distans wr
ffrnstwnall pairs of s-
goliash mthod amountd to a
statistialprinipl, whrby a
distansalong th ranhs of
Fitb 585
th tr showd th last diffrn with th pairwis protin squn dis-
tans.A ritial fatur of th mthod was a wighting shm that allowd
for th fat that gratr distans wr likly to rlativly lss rliabl.
!hil th algorithmi implmntation of th Fith-Margoliash mthod has
-I.B., fadd from us, distan mthods ar urrntly th most widly usd ap-
proah for ronstrutingphylognis,and th gnral ritrion of wightd
last squars has rmaind vial. Th utility and rliaility of th Fith-
Margoliash mthod is undrsord y th eukaryoti phylogny rportd in
thir papr, asd on ytohrom squns'whih is rmarkaly aurat
givn what w know now.
on of th most important onpts in gn volution is th distintion -
twnso-lldorthologs and pralogs.Fith oind thstrms in 197 and
usdthm to distinguish twngnsthat shar a ommon anstor du to
spiationvnts (ortholgs) and gns that divrgd via gn dupliation
(paralogs).This distintion is rlvant not only for asi molular volution
studisut also for th undrstandingf protin funtion. Indd,y th lat
1990s,th ortholog-paralog dihotomy am an indispnsalonptual
tool for invstigators trying to annott th poduts of ompltly s-
qundgnoms.This is ausorthologs tnd to nod protins with th
sam' or vry similar, funtions, whil paralogs oftn nod protins that
hav divrgd in funtion. Information transfr asd only on naiv squn
omparisonsan osur thsfats and lad to misannotationsasd on th
onflation of orthologous and paralogous squns.Mor nuand squn
omparison approahs that mra Fith's artiulation of th volutionay
prossy disriminating twnortholgous and pralogous rlationships
(2000)allow for mor rlial and usful annotatlons.
volutionary iology is an pliitly historial sin,and molular vo.
lutionists ar prsntd with th uniqu hallng of infrring th natur of
past vntsasd solly on ontmporanousdata' namly gn and protin
squnsof tant orgnisms. Dspit th lak of ass to historial data,
suh as th fossils utilizd y palontologistsor th anint tts diphrd
y historians,molular volutionistshav sussfullydvlopd an arsnal
of tools that provid a window to th past. volutinary sin is far lss
adpt, howvr, aout making prditions onrning th firtrrr ours of
vnts.In an audaious sris of studis,Fith and his ollagusattmptd
to dfy this trnd y atually prditing futur volutionary vnts. Thir f-
fort did not rprsnta mr aadmi xris.Th ojt of study was th
influnzavirus, and th ida was to try and prdit th ours of ftur flu
pidmis.Suh a prdition would provid a asis for sltingamng in-
flllenza strins to us in th prodution of a vain. To do this' Fith rlid
on a phylogntiapproah omparing th rlationship among flu strain g-
nomi squnsto th dats when thy wr olltd. A lar trnd mni-
fstditslf with th influnza phylogny showing a sussfultrunk linag
tht rprsntd th volutilnary history of th influnza strains that domi-
natd pidmis from on yr to th nt. Fousing on individual sits in th
squn'Fith and his ollagus(Fith et a|. 19971Bush t aL. 1999) were
586 Ford

al to idntify spifi variabl positions in th influnz squn that 1.96, a post h hll
ould b usd to hlp prdit thos strains that will aus futur pidmis. of All Souls ollgt
This old departur from th luidation of past rvnts)th status quo for al- ivd its Darwin Nz
most ll of volutionary iology, to th prdition of futur volr.ttionrytra- Ford am to th
itoris stands s a tstmnt to forward thinking arrd prsin1that hs st in th Lpidpt
makd l/altrFith as a trulv xDtionalmolular volutiollisr' thr of a populatio:
Their prsonal s
BlLIoGRAPHY ality rords going
Blin,I. 1953.Th tlghogd tb o.Londn:Widnfldnd Niolson. th papr pulishd
'W. aility of th popult
Bush,R. .' . A. Bndr,K. Suarao,N. J. o, llr-rd . Fith' 1999.
Prditinghvolutionof hunrr-r influnza. Sin286: 792|-1925, ltion, whih allolr
Fith, W. \4. 197.Distinguishinghomologousfrom analgousprotins.Sstmti h maintaind his i
Z o o l o g 1 9 : 9 9 - 1 1 3 . on ths animals ir
1'971,a. Rate f hangof onomitantlyvariblodons.lourI of
stsllrfor th sr
olulr uoltiolt |: 84-96.
1971.Toward dfiningth oursof volrrtion:inimun-r:lrrng for a Ford's earlistw
spifitrtopology.Sstmti Zoolog 2:406416' ly of a gn that af.
2000. Homology:A prsonalviw on somof th prolms.Trdsi G- cheureui. Thy sh
tis16 227_21. gn' but also n t}
Fith,W. M., R. M. Buslr.. A. Bndr,and N. J. Co. |997.Long trnrtrndsin
growth. Although I
th volutionof () HA1 humanirrflunza typA. Prldigs o|.tb
Ntionl Adryof SieesL]S 94: 7772,-7718. gns' te Gmmr
Fith,W. \4., and . Margoliash. 1967.nstrutionof phylognti trs.Sie Ford is prhaps
\55 279-284. was th first to off
Fith, !7. ., and . N1rkwitz.1970.An improvdmthodfor dtern-rining odon pard for Julian Ht
variabilityin gtrnd its appli:rtion
to th rtof fiationof mut:rtinsin
r ' o l u t i o n .B i o h t i , . dG t i s4 : > 7 9 - 5 9 3 . -r.../. Ford's ologiI
rn toBthr in th
spis in suh pro1
rurrnt mutatlon
Ford, dmund Briso (1901-1988) whih on morph i
. B. Ford, usually alld Hny y lris olla.usnd frinds, is st known and polymorphis
as th hif proponnt of th rsarhprogram known as ologial gntis. nti polymorphisr
Intndd to throw light on th rol of gnti divrsity in th adaptation of among loi ontrit
..su
organisms to thir nvironmnts, eologial gntis gins with osrva. what h alld
tions of variation in ntural populations. Brding studisthn dtrminth phnotyps' Batsi
mannr of inhritan of th diffrnt varitions. Cmparativ analyss of ampls of suprgn
diffrnt haitats and orrlations of nvironmntal and gti varials Although not gi
lad to hypothssout th rol of variation in volutionary adaptation. Fi- R. A. Fishr in th ]
nally' primntal manipulation of nvironmntal varials rsults in th th Sarlt igr m
onfirmation or rjtion of th hypothss.Ford's rsarhphilosophy is st Ford was al to tr
out in lris sminal ook, ol-lgilGentis. yarly nsuss.Fi
Ford spnt his rrtiresintifi arr at Oford Univrsity' H matriu- quny wr too g1
latd as an undrgraduatin 192, reeivinga BA in zology i |924 and a ontntion at tht
BS in \927. tle ws appointd Dpartmntal Dmonstrator in Zoo|og of fftivpopula
(1927|, thn Univrsity Dmonstrator (1929) and Lturr at Univrsity ol. that th original an
lg (1933). I 199, h ar Radr in Gntis, th first xpliit p- Ford was also in
pointmnt in that sujtin oford. H stalishdth Gnti l-aoratoris populations. For
in 1951 and rivd an appointmnt as Profssor of ologial Gntis in th hindwings of ,V

in th influnza squn that
rt will aus futltr pidmis.
ast vnts' th status quo for al-
tion o{ futur volutionary tra-
rinking and prsin that has
rolulr volutinist.
on: Widnfldnd Niolson.
.W.
'x, and . Fith. 1999.
\. Sic286: 1921-7925.
.om anlogusprotins. Sstmti
v:rriblodons.Jourl of
lution:inimum hang for a
):406416.
.reds
m of th prolms. in Ge-
| . o . 1 9 9 7 . L o n g t r m t r n d si n
t,vpA. Prodigs lf th
7.-7718.
:tirrof phylognti tres. Sien
vd r-rrhod for dtrmining odon
rttlf filrion tlf rnut.rtionsin
-r.K.l.
)a8usand frinds, is st known
21In known as ologial gntis.
lti divrsity in th adaptation of
al gntisgins with osrva-
dingstudisthn dtrmin th
iations.omparativ analyss of
rnmntal and gnti varials
ln in volutionaryadaptation. Fi-
)nmntal varials rsults in th
. Ford's rsarhphilosophy is st
o f o r d U n i v r s i t 1. H m t r i u -
rg a BA in zoology i |924 and a
rntl Dmonstrator in Zoo|og
9) and Lturr at Univrsity ol-
in Gntis, th first pliit ap.
alishdth Gnti LaL.loratoris
rofssorof oloial Gntis in
Ford 587
196, a post h hld until his rtirrntin \969. }-ord was lrda Fllow
of All Souls ollg in 1958. H was a ilow of th Royl Soity nd r-
eivdits Darwin dal tn 1954.
Ford am to th study of gntis and volution through his liflong intr-
st in th Lpidoptra. Bfor attnding Oford, h gan a study with his fa-
thr of a population of lit (phdrs) urii, th Marsh Fitillary.
Thi prsonal osrvationsmrad th yrs from 1917 to 1935, and lo-
lity rords going k to 1881 mke this an unusually lngthy study. In
th papr pulislrdwith his fathr in 1930, Ford attriutdth unusual vari-
ability of th population during priods of aundan to th rlaxation of s-
ltion,whih allowd suoptimal phnotypsto surviv.Throughout his lif
h maintaind his irrtrstin moths and ttrflis,prrlishingtwo volums
on ths animals in oliins's Nw Naturalist sris. Ilts Bltttrfljes ws a
stsllrfor th sris.
Ford's arlistwork at Oford was a ollaorativ study with Julian Hu-
ly of a gen that afftd th rat of darkning of tlr y olor in Gmmrlls
chureui. Thy showd that th irt dpnds not only on th Mndlian
gn,[.lutlso on th siz of th y, th tmpratur'and th rat of odily
growth. Although not th first to ttiut volr'rtionaryimportan to rat
gns'th Gmmrus studisprovidd an important arly ampl.
Ford is prhaps st known fr his wok on gnti polymorphism. H
Was th first to offer a ritial dfinition of th phnomnon in a papr pr-
pad for Julian Huxl,s The Ntu S1'5ngmtis(1940), rfind as follows in
Ford,ologil Gtis (|964,84). ..Gntipolymorphisrrris th our-
rn togthr in th sam loality of two or mor disontinuous forms of a
spisin suh proportions that th rarst of thr annot maintaind y
rurrnt mtation.'' H distinguishd twn transint polymorphism, in
whih on morp is in th pross of rplaing anothr, and stal or al-
nd polymorphism, maintaind by stailizing sltion.H notd thL1tg-
nti polymorphism oftn irrvolvd larg phnotypi ffts, los linkag
among loi ontriuting to thos ffts, and strong sltion, lading to
..suprgns.''
what h alld H intrprtdth ABo lood grous' Rhsus
phnotyps, Batsian mimiry in uttrflis, and htrostyly in plants as x-
ampls of suprgns'a position orn out y susquntrsarh.
Although not givn to mathmatial nrodling, Ford ollortd with
R. A. Fishr tn th |947 study of olor polymorphismin Pi dlmiul,
th Sarlt Tigr moth. Sin htrozygots at this lous ar distinguishal,
Ford was al to trak gn frquny and popultion siz in a long srisof
yarly nsuss.Fishr,s analysis stalishd that th hangs in gn fr.
quny wr too gt to rought aout y gr.rti drift lon, a point of
ontntion at that tim. Although hallngd y Swall \right on th asis
of fftivpopulation sizs,susquentwork on this olony has stalishd
t h t t h o r i g i n a l a n a l y s i sW s o r t .
Ford was also intrstdin th fftsof polygns on variation in natural
populations' For many yars h studid variation in th pattns of spots on
th hindwings of ill ;jrti,th Madow Brown uttrfly. Tlr pattrns
.'88 Flrm and Funtilr

wr mark:rblyonstant ovr wid aras of urop, ut thy sho,dgrat thn th fr-rrrtionstht '
hzrngsovr slight distarrsin sor.rthwstngland arrd th Isls of Sillv' of ftlrm. Russll rfrr
Spot pattrns on th lrg islands tndd tl similar, whil tlros from T h p r t l - { u n i r lpno s i t i r
smallr islands diffrd rdially. Ford attriutd tl"ronstany lf pattrn r i o r o h | sD e r w i r l
ovr tim to stailizing sltion and th diffrn btwn larg .rnd small natural tlrologians.Aft
islnds to th fft of avrirgingovr divrs haitts orr th larg islands. implrtanof naturl st
In addition to his gnti wok' Ford mad orrtriutionsirr svral othr in th i830 dat t
filds. pulishd on th hmistry of uttrfly pigllrnts'on th arhaol- Hilair (olman 1964
ogy of southwst nglnd' and on th ontnts of hurhs in th Oford forms of nimal odis
distrit. h d t b s r v d ,t h i r
Prhaps Forl'sgrastirrrilton th str'rdyof vari:lti<lnin natrrr:llpopula_ tlrat animirls slrardbst
tions has <.lmaout tlrrough his intations 'uvithhis studr-rts and ol- D a r w i l r r p t l r r dO l l t l ) i
lagus.H influndth work of Philip M. Shppard,yil A. lark, A..}. on th siJ of |.untion
ain, H. B. D. Kttlwll, Laurn M. Cook, Linolrr P. Browr, John R. G. funtion-was th aus(
Turnr' Ronilld A. Fishr, and many othrs.Ford ws prhaps too inlind to
It is gnrall1,akn<lu
it only his frinds and ollgus,ut, s Brya l:rrk (199.5,l62) maks
two gat laws: Unitl.
lar in his Royal Socit Biogrpbicl moir, ..still ologil Gee-
of typ is rnant that
tis . . . is nssary far for srious wlrkrs in th fild, still it is th lrst
in orgar-riings of t
and last ompromising sttmntof th .sltionist'viw, and still it is a joy
thir haits of lif. ol
to rad.''
d s n t . h p * s
y th illustrious ul,
|BLIoGRAPHY
ltion.For ntural s
lark,B. . l995. dmundBrisoFord. Bilgrphil tlirsof Felkuslfth parts of ah ing t<
RrylSoitl,of l'odl41z147_168,
having irdgltdthm
rd,. R., . B. Ford, lrndK. G. ]vlWhirter.1964'volutionrystrrdis on
illiurtl:Th Islsof Silly,1958_59.Hrdit1'19:471_488. ing aidd in sotn .
F . i s h r , R . A . , a n d . B . F o r1d9. 4 7 . T s p a d oaf g r r i l n t u oln d i t i o n s i n a th dirt tion tlf th
olony of th mth Plidomiul L-.Hrdit 1: |4_174, suitd t th sv
Fod,. B. 194.5.Buttr|lis, London:llrs. onditilns tlf ist
1964 ||9751.clllgit..l caetis.4th1.Londn:haprnirnnd ll. inlrritan of forlll
-]..
1 8 - 5 92, 0 6 )

Latr-nintnth-ntu
Forrn nd Function (dward s. Russll)
doutd th powr rlf r
dward S' Rr"rssll's For d Ft.mtilis uniqr.rlr,alulrsollr in th iormal, o somtilllsL
history of r-rimlmorphology. It is a livIy xploratin of th most imporrant o d i so f a n i m | s( s
morphologil thlristsfom th nintnththrough th arly twntith n- Form tld Futirl t
turis. How ould a rport on long-dad mophologists livly? Th livli. tts and rpr:oduti<
llss oms frorr-rth fat thr Russll organizsth thoristsaround ntral tratiss.((il1.lis of Rr
dt thi1tprsistdthroughout th nilrtnthlltuy and prsistsvn to- rrintnth ntury lrlrr.
day. Th dat is in th titl. Form nd Futio ould just as wll hv nals y lirtr historians
n titld orm ursus Futio. To put th mattr simply, whih am first, t n t i O ne t i t s p u l i . r
form or funtion? Why do odis of anirrralshav th ftlrms thy d<l?Do th morph<lllgists nd sorn
forms ist bzrusthy allo''v funtions to svl?[f s, frrntior-rlisnr
is sond itlr-ry,ws tht
favord arrd form is a v.prldut of funtion. or ar organismi forms or d u r i n g t l l r l yt w n t i
strutursa produt lf autonomlus fom-grlrtingprlsss?If th lattr, (sth main ssay ..Th
 tlm and Futrtin .t89

Lrpr'ut thy showd grat thn tlr funtions that ody arts srv ar th y.produts' llot th auss
..pu morphology.''
of frr. Rrrssll rfrrd to th prl-f<rrmpsitin s
,l.rndnd tlr Isls of Silly.
, similar, whil thos from Th pro-funtion plsitiontook various fornrs during th nintnthntury.
d th onsteny of pattrn Prior to hals Darwin, funrilnalistsWrtlologists,most of whom wr
n twnlarg Lrnd small ntural tlroltrgians.Aftr Dilrwin' most furtionlistsWr livsin th
itatson th larg islands. importanof naturl sltionor Lamarkism. Th ontrast was illustratd
ontriutilnsin svral othr in th 1830 dat twn Gorgs uvir irnd tinn Goffroy Saint-
|y igmnts,on th arhol- H i l a i r ( o l m a n l 9 6 4 ; A p p l 1 9 8 7 ; R u s 2 0 0 3 ) . u v i r l r i m dt h a t t h
s f hurhs in th oxford forms of animal odis and body parts wr ditatd y th funtions tht
had to srvd, tlrir so.alld ondirions of istrr.Goffroy laimd
f variationin naturl popula- that animals shard astrt ldy plr-rs, and lr insistd on th unity of typ.
s with his studnts and ol- Darwin rportd on this dat in o th rigi of Spis,ut am down
p p a r d , r ' ' r i l A . l a k , A . J . on th sid oi furrtion.Naturl sltion-a pr()SStht shps form ttl suit
,inolrrP. Browr, Jt>hnR. G. funtiln-was th aus of volutionary hng.
d was prhapstoo irrlind to It is gn|l,vakn<lrvldgltlrat all organi Lringshr,n formd on
a r r l r k ( I 9 9 5 , 1 6 2 ) m a k s tw() grat laws: Urrity of Tp, and th onditions of istn.By unity
,,stt||cologil Ge-
ir, of typ is rnt that fundarnntlirgrmntin strutur,wlrih W s
th hld, still it is th last in lrgani ings of tlr stll lass, nd rvl.rihis quit indprrdntof
i o r l i s rv. i w , n d s t i l l i t i s j o thir haits of lif. On my thlry,trnity of typ is plaind y unity of
dsnt.Th prssilnof onditilrrslf istn'so oftn insistd on
th illtrstriousuvir, is fully rrrrady th prinipl of natural s-
ltion. For natural sltion ats y ith now adapting th varying
l Mtirs lf FIlo's of th pats of alr ing ro its lrgni and inorgr-rionditions of lif; or y
hving adptd tm durirrg long-pst pildsof tim: th adaptations
4. volutionarystudis on eing aidd in som ass y us and disus, ing slightly afftd y
Irtlit1,1'9 47 |-488.
th dirt ation lf tlr trtrlonditios o lif, nd ig irr all ss
g r l l i n n t t l l o t l t l i i t l n isl l
|it1,1: 11_174' sujtdto th svral larvs of gIOWth. n, in fat, th law of th
nditiolls of istn is th higlrr law; as it inluds, tlrrough rh
hrllan nd il.
-onr.1tln: inhritalr tlf fonlr 1ptatiolls,r:rt rf Urriry of Tvp. (Darwin
-1.til.
1859,206)

Ltr-nintllth-ntury vtllutionistsswung k twrd frnr again. Tlry

ll) doutd th powr of nturl sltitlrr.Insted, thy livd that ruls of
iqulyvalualrsour in tlr form1,or somtillls mryologial, onstrution Wr rsporrsilfor th
loraionof th most irnportarrt o d i so f n i r n a l s( s R u s 1 9 9 6 a n d R i h a 12s 0 0 8 ) .
lrough th arly twntith n- Form d Ftio ontains a numr of usful translations of original
lhologists livly? Th livli- tts and rglrodutions of illustratins fro-r hard-to-find rrrrphologial
s tlrthoistsaround a ntral trtiss.(opis tlf Russll's rdrawings of rnorpholt>gialplats froln th
h nturyand prsistsvn to- nintnthntury hav ftn n passd lff as produtions of th origi.
titl oa|d jr-lstas wll hav nals y latr lristorinsand biologists.)Ironiallv, th ook rivd littl at-
-rattrsimpl, whih am first, tntion at its pulition in 1916. |t gan tl takn mor sriously y
tavth forrnsthy do? Do th morphologists nd slmvtllutionary thorists around 797.The rason' a
r srvd?If so, funtiorralism is sond irtllrv, was that th study of mtrrphlog \,\,ntthfoug'h a dry spil
t. or ar orgenismiforms or during th arly twntith ntury. Th modn synthtithoy of volution
rtingprosss?lf th lattr, (sth mairl ssy ..Th Hist<lryof r,olutionarvThought'' y ihal Rus
590 Form and Funtion

in this volum), whih was inauguratd in th 190s and wll stalishdy

1950, saw littl importan in traditional morphologial studis.owevr, a Frish, Karl Rittr
growing minority of volutionay thorists' som of thm ritis of th syn- Kari Ritt von Frish r
theti thory' wr ronsiding th imprtan of morphology during th t h . ss i d y t h r g l
1970s. On of ths,Gorg V. Laudr, a nw assistantpofssorat th Uni- numr of notd ad
-l
vrsity of hiago in 981' larnd that th Univrsity of hiago Prss was h a v d i s r i n g u i s h d
rprinting historially important sintifi tts. He nminatd Form d who inflund him mt
Futio and wrot a usful introdution. Laudr disussdwhy morphology ology at th Univrsity
had so littl rlvan to th synthti thory of volution' ut h rportd t th Univrsity of
that morphologists wr on again ginning to onsidr th nintnth- t n o f a r f u l p i
ntury topis of strutural pttrns and prinipls of organismi dsign. So ful osrvr of living
Russll,s 1916 ook was rprintd ausof th rbirth of intrstin mor- d v i s i n gs i m p l p r
phology around 1980. Laudr and othr twntith-nturymorphlogists onditions.
drw insights from Russll's history that wr rlvant to ontmporary d- W h n F r i s h W sa y
ats in volutionary thory (Laudr 1996; Amundson and Laudr |994). nows to adpt thi t
Russll himslf favord funtionalism ovr pur morphology, ut his book i n t o o n f l i tw i t h t h
prsntdth pur morphologists in a sympathti light. Rognition of th m o l o g i s t .S t u d y i n gh o
virtus of his ook Was not limitd to morphologists' howvr. rnst Mayr' ludd that fish and irr
arhitt of th synthti thory of volution and a funtionalist to th or' othrwis olor-blind.
ommntd on Form d Futil: ,,Ter is proably no othr rnh of r a i s d n d k p t m n y t
iology for whih w hav as supr a history as fr morphology: Russll l o f h a n g i n g o l o r a
(1916) is unsurpassdto this dy, rmarka1fr its frsh analysis of th pri- tim. This suggstdto
m a r y s o u r s ' '( a y r 1 9 8 2 , 8 7 9 n | 5 ) . h prodd to tst z
|BLloGRAPHY Frish's findings, and tJ
a dad. arly in th
Amundson,R., nd G. V. Ludr.1994.Funtilnwithoutpurpos:Th ussof
o l o r s n s i n h o n y
ausalrol funtionin volutionaryiology.Biolog d Phibsoph 9:
t o m a k F i s hw o r ] d .
44469.
Appl' T. . |987. The uuir-GoffroDbt:FrhBiolog i th Ddes F r i s h ' sf i r s t s t u d i s
bforeDri. Nw York: Oxford UnivrsityPrss. shp.H nt studi
.w.1964'
Colmn' Gorgesuuir' Zoologist:A Stud i the Histlrlf uolution mnts h disovrd tl
Thor.amridg,A: HvrdUnivrsityPrss. thir fllow workrsi
Darwin, . 1859.n th rigi of Sples.London:John Murry.
1920 he pulishd his
Ludr'G.. 1996.Th argumntfrom dsign.ln M. R. Ros nd G. V. Laudr,
ds.,dpttil,55-91. San Digo'A: AadmiPrss. s.H undrstood th
Mayr, . 1982. The Groth lf Biohgil Thlught.Cmidg,A: arvard dans, a round dn
UnivrsityPrss. th hiv. Initially h su
Rihards,R. J. 2008. Th Trgi Ssof Lif: rst Hkld th Strgglour and th waggl dan
ulltiorThought.hiago:Univrsityof hiago Prss. 1944 tha h disovr
Rus, M. 1996. od to : Tb pt of Prlgrssi uolutiorBiolog.
of rh food sour fol
ambridg,MA: Harvrd UnivrsityPrss.
23.Drtui d Dsig:Dos uolutitlue Purpose?amridg, lowing spring h arriv
A: Harvard UnivrsitPrss. prformd y bs rtt
Russll,. S. 19l 6 [1982|.Form d Futio.London:John urray. Rprint, th hiv, srvs to oll
hiago: Univrsity of hiago Prss. -R.. food sour.ln his fuI
p i v p o I a r i z dI i g h
t h m o n l o n g f l i g h t sr o
r i s h s p n m u h o |
d dR i h r d H r w

930s nd wll stalishd y
rlogialstudis.owvr,
n of thm ritis of th syn-
of morphology during th
lssistantpofssorat th Uni-
ivrsityof hiago Prss was
ts. nominated Form d
r disussdwhy morphology
of volutin, ut h rportd
] to onsidr th nintnth-
iplsof organismi dsign. So
lrrirth of intrst in mor-
:ntith-nturyrophologists
rlvntto ontmporary d-
,mundsonand Laudr 1994).
r morphology, ut his ook
h t i l i g h t . R o g n i t i o n o f t h
rlogists,lrowvr. rnst Mayr,
'nd funtilnalist to tlr or,
proably no othr branh of
y as for morphology: Russll
for its frsh anlysis of th pri-
urpos:Tlr ussof
rithout
iologd Phillsoph9:
|renhBiritlgin th Ddes
Prss.
Studitth Histlr1'of uolutil
Prss.
rn:JohnNlurray.
M. R. Rosand G. V. Laudr,
lemiPrss.
r. ambridg' MA: rvard
t Hkld t Strugglelt,r
[ hiagoPrss.
rogrssill,uolutiorB iolog'
t,'L1Purpos?arbridg,
lndon:Jlhnurray. o.o.'1*.o.
rish 5L)1
Frish,Karl Rittr von (1886_1982)
Karl Rittr von Frish was lrn in Vinn to a fantily distingr.rishd
on his fa-
thr's sid y thr snrations of physiians and on his mothr's sid y a
numbr of notd aadmis. A lovr of ilnimals as a youth, h wnt on to
hav a distinguislrdarr lls an xprimntal physiologist. Th tahrs
who inflund hirn most wr his unl, Sigmund nr' profssor of physi-
ology at th Univrsity of Vinrra, and Ril-rardtwig, profssor of zoology
at th Univrsity of Munih. Both irnprssdupon hirn th itil impor-
tan of arful xprimntation as a ornplmnt to his talrrts as a thought.
ful osrvr of living animals. Frish rspondd y oming mastr at
dvising sirnpl xpirnntsto tst th bhavior of animals undr natural
onditions.
.Whn F.rishWas a young rsarhr, his arly work on th ility of min-
nows to adapt thi oloration to dark or light kgrounls rought him
into onflit with th laims ot arl von ss, a prominrrt Munih lphthal-
mologist. Studying how various animals rspond to light, Hss had on-
ludd that fish and invrtratsspond to th rightnssof olors ut ar
othrwisolor-lind. Fish, oth s naturalist nd s somon who hd
aisd arrd kpt many diffrnt spis f fish' knw that som fish ar apa-
l of hanging olor and that rtin spiesar spiallyolorful at mating
tim. This suggstdto him that fish hav a snsof olor, a hypothsis that
h prodd to tSt and onfirm primntally. Hss, howvr, rjtd
Frish's findings, and th dt that nsud twn thm lrstdmor than
a dad. arly in tlr dbat Frish prodd to addrss th qustion of
olor sns in horrybs.Tlrr-rsgarrir whol sris of rsrhSrhat Wr
to mak Frish world-famous.
Frish's first studis f honysxmind th insts' snsof olor and
shap.H nt studid th s' sns of smll. In th ours of his pri-
mnts h disovrd that sout s ar somhow al to ommuniilt to
thir fllow workrs in th hiv th xistn of dsiral food sour. In
192 e pulishd his first rports on wlrat h alld th ..languag''of th
s. undrstood th s,..languag''to inr,,olvwo distinivkirrds of
dans,a round dan and a wggl dan, prformd y s rtr'trningto
th hiv' Initially he supposd that th round dan signifid ntar sour
nd th waggl darr signifid pllln sour. It ws not until th fall of
|944 tat h disvrdthat th diffrnt dans r rlatd to th distan
of th fold sor from th hiv and not to th natur of th food. Th fol-
lowing spring h arivd at th rmarkal onlusir-r that t waggl d:rn,
prforrrrdy bes turning from food sours rnor than 100 mtrs from
th hiv, srvs to ommunit oth th distan and th dirtion f th
food sour. ln his furthr studis of honysh dmonstratd that thy
privpolarizilight and in dditilrrlrv an ..intrnallok.'tht nls
thm on long flights to adjust to th sun's hanging pla in th sky.
Frish spnt mulr of his arr t th Univrsity of unih, whr h su-
ddRihLrrdrtlvig in \925 as profssor of zoologv anl dirtor of th
592 Frish

Zoo|ogia| Institut.In th rly 1930s, with hlp from th Bvarin govrn-

mnt nd th Rokfllr Foundation, Frish mad th Zoo|ogta| Institut at
unih th most advand sintifi ntr of its kind in urop. H man-
agd to rtin his position throughout th Third Rih' though not without
som diffiultis frorn th Nazi authoritis. Atir Allid on-ringdstroyd
muh of th institut in 1944, h ontinud his studis in Austria, first at his
summ h<lmin Brunnwinkl and thn as profssor of zoology at th Univr-
sity of Graz (|946_|950). H rturnd to th Univrsity of unih in 1950,
whr h remaind until his rtirn]ntin 1958. Among his lny honors was
te 197 Nol rize for Physiology or diin' whih h shard with Kon-
Th Galipagos
radLorez and Niko Tinrgn.
Th Gal6pagos Ar
lBLloGRAPl]Y
70 mils long) loat
.!.,
Burkhardt,R. Jr. 1990. Frish,Karl Rittr von. In Ditior of Sintifi dor, to whih thy
Biogrph'supplmnt 2, 17: 312-2. ativ) and of r
Frish,K. von, 1967. tsiologistRemntbrs.ListhGomrih,trarrs.Oxford:
-R.\B. plat driftd strl
rgmonPrss.
hot and thr is lit
1532, thy wr ini
Amrian novlist
mad a sour allusi
ground" (1987,12
volutionists thy a
Th islands soon
gos ing Spanish f
strs that rowdi
tintion' thanks t
ruts providd
sporadially by sai
first in th svnt
and thn in th ig
arry Spanish nam
Th Gal6pagos A
arond th glob, <
was fasinatd y
birds that liv on th
oth flora and faun
and signifian of
What did strik Dal
ganisms on th Ga
fat that th island
Non of this sm
.!hy
origins. woui
why South Amri
islands?
is uriosity pi
tions of th island
hlp from th Bavarian govrn-
n a d t h Z o o l o g i a l I n s i t u t a t
f its kind in urop. H man-
rird Rih, though not withut
\ftr Allid oming dstroyd
ris studis in Austria, first at his
,fssorof zoology at th Univr.
:Univrsit1'of unih in 1950,
8. Among his many honors was
;in,whih h shard with Kon-
o,In Dictior of Sitifi
th Gomrih, trans. oford:
-R.vB.
Th Gal6pagos Afhiplago
Th Galipagos Arhiplago is a group of 14 small islands (th largst aout
70 mils long) loatd on th quator aout 500 mils off th oast of ua-
dor, to whih thy long. Thy ar volani in origin (som are still slightly
ativ) and of rnt gologial ag. Thy wr formd as th ontinntal
plat driftd astrly. Harsh, with indr and ash undrground, th limat is
.W,strnrs
hot and thr is littl or no frsh watr. First disovred y in
152, te wr initially alld t,etds, or th nhantd Isls. Th
Amrian novlist Hrman lvill, who visitd thm on a whaling trip,
mad a sour allusion to this nam, dsriing thm as ..villy nhantd
ground'' (1987, 129), adding: ..an and wolf alik disown thm'' (127). For
volutionists thy ar far from vil and thy ar rtinly nhantd.
Th islands soon took on th nam of thir most famous dnizns' Gal6pa-
gos ing Spanish for tortois. Not just any kind of tortois, ut giant mon-
strs that rowdd th isls until thir numrs wr rdud to nar
tintion' thanks to th killing and taking y marinrs who found that th
ruts providd llnt far for long sa voyags. Th islands wr visitd
sporadially y sailors, hifly uanrs, nd wr givn nglish nams,
first in th svntnthntury aftr prominnt prsonags (lik King Jams II)
and thn in th ightnth ntllry aftr worthis of that tim. Today, thy
arry Spanish nams givn y uador.
Th Gal6pagos Arhiplago was visitd in 1835 y HS BegIon its trip
around th glo, arrying harls Darwin as th ship's naturalist. Darwin
was fasinatd y th wildlif, spially th tortoiss and th many small
irds that liv on th islands. As was his ustom, h mad larg olltions of
both flora and fauna, although as yt h did not truly raliz th importan
and signifian of kping sparat th individuals from differnt islands.
!hat did strik Darwin at on was th sming rlationship twn th or-
ganisms on th Gal6pagos and th South Amrian mainland, and also th
fat that th island dwllrs gnrally had slight ut distintiv diffrns.
Non of this smdto mak sns from a rtionist viwpoint of organi
.Why
origins. would th organisms similar to th mainland organisms' and
why South Amria rather than Afria? And why th diffrns twn th
islands?
His uriosity piqud, Darwin mulld ovr th iogographial distriu-
tions of th island spimns, onntrating spially on svral spis of
59
594 The Gldpgos rhipelgo

mokingird that h knw am from diffrnt islnds and that smddif- David Lak, who throu
frnt physially. thought als<raout th tortoiss tht th arhiplago's Ga16pagosjust ftlr\x
govrnor had dsridas distintivto thir rsptivislands.As h trid to siv olltion of th h
put in ord th nrany small linhs tht h had lltd, h saw that thy Amrirr usunr oi N
too rvalddiffrnson diffrnt islands. ould thsall rnrvaritis Lk at first put .rll sig
or Wr thy somthing mor, in fat and in thory? War yars on his {inding
Whn Darwin turnd to ngland' h gav his spirnns to spialists to was no dout mulr infl
atalog. At this point Darwin flt oligd to slip ovr tO n volutionary po_ onlmist rnst a.vr. I,:
sition. Th irds partiularly wr undoutdly diffrnt spis, and this thal
prisly at th tir-l-r
ould plaind naturally only on tlr supposition tht original foundrs th synhsis Sstmli
hd om to th arhiplago and hangd and volvd as thy and thir su- D r i's F ihs (1947 )
ssorsmovd from islnd ro island. A ruial part of this thinking was tht rlught on y strrrggl
volutionav hang involvs division of populations and susqtrntspi- Lak's ook has now gi.
tion. Darwin thrfor always thought of volution's history as ing funda- linkd Dawin with th
mntally on of splitting and divrging. Its pattrn was oral-lik (his first nat fft of making
mtaphor) o a tr of lif (th mtaphor tht h mad f:rmousin O the ori- md Drwin an r,ol
gin of Spcis).Hr lr diffrd fro arlir volutionists wo thought of mokingirds (sSLrllt
volrrtion as ssntiallya mov up a singl lin, with variations along minor Aftr Lk,th Gli1
sid branhs. i n t h n o - D a r w i n i a ns y
Although th Gal6pagos took somwhat of a sondary rol as his think- e t n i l t U r o f t h ' l t
ing nraturd, Darwin always usd islarrd iogogaphy as a ky support of lorving British thinkinE
volutionism, frquntly irknowldging th importan of his visit to th fator. In Amria, in lir
Gal6pagos. Praly in part aus of this signifian,th grat Swiss- tur lf organisms' thr
Amrian ihthyologist Louis Agassiz, th most important post-orlgin sin- olog,vof th islnds
tist wlro nvr irptdany forrr-rof volutionism, visitd th Galipagos lat was plord and strtlr
in lif in 1B7Z' Peraps unsurprisingly, h flt that is spial rationist vi. (196I), who ma.la lon
sion of lif's history was vindiatd y th Gal6pagos and its inhaitants. xistnof altrnativ
Agssiz thought thr was insuffiinttirn to produ so varid display on Th British Wr unll
islnds gologially so young (sLarson 2001, spiallyhaptr 4). ow- until th 1970s' Th lrr
vr' no on ally followd in his path' Soon th Gal:ipagosArhiplago was t o g t h rw i t h e s t l s
stalishdas a priz as for volutio, although, ln lin with th gnral otnon of th lss
post-rigi disrditing of natural sltion, most ,ould hav agrd with 199]r:B. R. Grant and I
Agassiz that sonr othr mlranism was nddto prodr-rth aninlals and 2 0 0 7 ; W i n r 1 9 9 4 ) .V
plants as thy ar found today. ringd and watlrd anr
Suh ws th stat of affirs through th 1930s, wl,rnDarwinian sltion tions nd physial han
was mlded with Mndlian gntis and modrn volutionary thory. No- fts rought on b dro
Darwinism, or th synthti thory of volution, was orn. Howvr, thr Works onsantly' nl l
was no an imnrdiat rush to ring th Gal6pagos undr th nt of naturl s- nt. orovr' tllr s
ltion. Thr was a gnral fling, spially among Amrians' that small availailitis ar lr sign
differns twn groups r rarly adaptiv. Th population gntiist S- omptition twndi
wall Wright (19\, 792) gavethortialaking to this prjudiy arguing Although th Glipa1
that mny diffrns ar du to so-alld gnti drift, th randolrnssthat is arl1,samn whl took
.!7
rought y th vagaris of rding. Tlr diffrns twn Glipagos or- II who uilt nn airl
ganisms wr oftn put down to just suh a aus.No on dnid that sltion is now ralization y s
was important ovrall, ut not in th kinls of dtails rvald y th islands. pOrtnt natural phrron
'Ih
ky figr'rrin hanging tlris opinion WS an nglish shooltahr, forts ar ing md ttl

:nt islndsand that smd dif-
tortoissthat th arhiplago's
rsptivislands.As h trid to
had olltd,h saw that thy
] o u l d r h s a l l m r v i t i s
,hory?
his spir-rrns to spialists to
slip ov to an volutionary po-
dly diffrnt spis' and this
lposition that original founds
d volvdas rhy and thir su-
al prt of this thinking was that
u l a t i o n sl r n d s u s q u n rs p i a -
llution's history as ing funda-
pttrn was oral-lik (his first
h mad famous in o tbe ori-
:r volutilnistswho thought of
n , w i t h v r i a t i o n s l o n g m i n o r
lf a sondarrol as his think-
rgogphyas a ky support of
: impotan of his visit to th
is signifian, th grt Swiss-
ost impotantpost-orryi sin-
lnism,visitd th Gal6pagos lat
: l r t h t h i s s p i l r a t i o n i s tv i -
Gal5pagos and its inhabitants.
o produso varid a display on
) 0 1 ' s p i a l l y h a p t r 4 ) . H o w -
r th GalripagosArhiplago waS
though, in lin with th gnral
l' most would hav agrd with
ldd to produ th animals and
L930s,whn Darwinian sltion
odrn volutionary thory. No-
ltion' was orrr. Howvr' thr
>agosundr th rrt of ntural s-
lly among Amrians, that small
v. Tlr population gntiist s-
k i n gt o t h i s p r j u d i y a r g u i n g
ntidri{t,t randomnss that is
ffrnsrwnGaldpagos or-
tus.No on dnid that sltion
f dtilsvzrldy th islands.
L was an nglish shooltahr,
Tb Gldpgos Arhiplgo t9t
David Lk, who through th notrrgnrtrt rfJulian Huly lvnt to th
Gal5pagos jrrstfoWorld lWar II to stlld,Ytlr irds. H thn gan ir nras_
siv olltion of th irds-finhs partiularly-that was ownd y tlr
Amrian usutl of Natural History in Nw York. \riting up his findings,
Lak at first put ll signifint hng down to drift. Rflting during th
Wr yars on his findings, Lak swung ntirly to a sltivxplanation. H
Wasno dout muh irrflundby th Grman-orn, Amrian-rsiding t.
onomist rnst ar. Lak roomd with ayr and his family in Nw Yok,
prislyt th tirrltlrrrtlayr ws finishing his pro-sltionontriution to
th synthsis Sstttt"ttisd tle rigi of Speies (1942). Lk irrgui in
Dri's Fitlbs\1947) that all diffllslrtwnth spisar ad:rptiv,
rought on y struggl twn groups for sours.Although th titl of
Lak's ook has now givn th popular nn] to th finhs and has publily
linkd Darwin with th island inhaitants,it also had th somwhat unfortu.
nat fft of making popl think that it ws th finhs lusivly tht
mad Darwin an volutionist. At lst, if n()t mor important, Wr th
mokingirds (sSrrllowa 1982,).
Aftr Lak, th Glzipilgosirds Wr on oi tlr prim pis of vidn
in th no-Darwinirrsynthsis.Howvr, ontrovrsyorrtinudovr th -
at ntur of t sltivfor that lrad otlght on diffns.Lak, fol.
lowing British tir-rking,had mad omptition twn groups th k1'
fator.In Amri, in lin with gnalthinkig in that ountry aout th rr.
tur of organisrs,thr was mor nthusiasl for a position that mad tlr
ology f th islnds mor signifiant.This viw of sltion and its ffts
was plord and strongly dfndd y alifornia iologist Rort Bowman
(1967),whl mad long visit to th islnds in 19.52_195.H found that th
istnof ltrnirtivfold supplis Was th ky firtor in sltivhng.
Th British wr ltt-tit-nglrssd,
and tlrus mrtrs rrrrindat an impss
Llntiith \970s. Tlr htls:rn]-and-wif tm f Ptr and Rosmary Grant'
togthrrvith ir sussionof oworks ilnd studnts,gan what lras sirr
omon of th lassi long.trm studis of tlatural sltion(Grant l986,
1997;B. R. Grnt nd P. R. Grarrt1989;P. R' Grant and B. R. Grant 1995,
2007; !inr 7994)..sorkingon th islt D;lhn Major (s figur), th1'
ringd and wathd and rordd vry ird, noting th numr of flutua.
tions and physial lrangsfronr on yar tO th nxt' spiallynoting th f_
ftsrought on y drought and y rord rinfzrll.Thy found that seltion
works onstntly' irrrltlrat hngs an our rapidly fror on yar tO th
nxt. orovr) thr stllsto truth in th lz.rirthzrtfood supplis and
vailabilitisirr a signifiant fatol. in sltir,hang, ut not luding
omptitioll tlvndiffrnt groups.
Although tlr Glipagos has n attrd,vhuman oupany' from th
arly samn wlro took th tortoiss,through th Amrin fors in \orld
War II who uilt alr airas thr' to th prsnt uadorian fishrmn,thr
is now ralization y sintists and govrnmntl authoritis of what an im-
portant natural phnomnon th Gal6pagos Arhiplago rprsnts.Srious f_
forts ar ing md to prott and onsrv th islnds and thir inhitants.
596 h Glipglsr..hipelgl

lvill, . 1987.Th
Pros Pis,18.J
of Hrnluil
ds.vanston,II,:
S u l l o w a yF, . J . 1 9 8 2 .
th Histor of Bkl
Winr,J. 1994.ThF
York: Knopf.
Wright, S. 191.volu
1 9 2 . h r l
vlutiorl.Pro

Galton, Franis
Franis Galton was
to pursu a rr ir
study mathmatis'
Darwin and Galton
Th islt Daphn jor in th Glipagls Arhiplago,whr Ptr and win (1731-1802), a
Rsmry Grant did thir dirds-longstudy of Drwin's finlrs.Th would a ruil
volni origins oi th islt, hill with th top lown off anl ratr formd fat, it was Galton's
within, ar larly visibl.It has n possilto work out rlationshrps
th ida f improvin
twntlr dniznsof th islnds from th squntialordr in whih th
h oind th trm e
islands wr formd, y volrriativity' s th rth'spltsslippd aound
its surf. Sevrl ,Yarsaft
Galton organizd an
uropan ploratior
latituds,longituds
Now thy ar on of th most popular of dstinations for tourists. Darwin's ation of numriala
visit to th island, rathr than lvill's, has provn to hav th longr- him. H pulishdhi
lasting fft. in 1852 and was awa
BIBLIOGRAPHY aus of his pris,
ook aout his jlur
Bowman, R. 1961. orpholrlgiI Diffrtitio d dpttil i th Gldpgos
book for amatur an
Fihs. Brkly:Univrsitr,of lifor-riPrss'
Grant, B. R., and . R. Grant. 1989. uolutionr D,mics of Nturl Popul- triumph. It wnt thr
ti.l:,hLrg tus Fih of tb Glipgos, hiago: Univrsity of hiago Phonix Prss rissu
Prss. Galton Was an ativ
Grant, P. R. 1986. lklg d utllutilof Drtui's Fihs' Printn, NJ: yars. FIis ommnt
Pritrton Univrsity rss.
mattrs (.g.,latitud
199l. Ntural sltionnd Darwin's finhes.Sitifimri, Otor:
82-87.
intrstd in mtor
Grnt, P. R., nd B. R. Grnt' 1995. Prditing mirvlutionar rsponssto hartristiof a hi1
dirtional sltionon hitalvriation. u oluti on 49 : 241 -25 1. Th part of Galtor
27, Hl d Wh Spis ltipl: Th Rditil of Drtui's Fichs. aftr h had rad Dal
Printon' NJ: Prirrton Univrsity Prss. and ultivatl plants
Lak, D. |947. Dru,'itis Fihs.amridg: anrridg UrrivrsityPrss.
able to Galton that t
Larson, . J. 2001. uoltirl's Workshp: od d Sin ln t Glipgos
Islds. Nw York: Basi Books. to dmonstrt that .
ayr, . |942, Sstttttisd th rigi of Spis. Nw Ytlrk: olumr hritd. H attmpt
Univsitv Prss. Talnt and hat

iplago,whr Ptr and
f Darwin'sfinhs.Th
lown oif and a ratr frmd
o work out rltionships
quntilordr in whih th
afth'splatsslippd around
inationsfor tourists. Darwin's
ls provn t hav th longr.
d dpttil i th Gldpgt.>s
ss.
Dmis of Ntrl Popul-
s. hiagl:Univrsity of hiago
i'sFihs.Pinton,NJ: yars. His omrlnts at soiety mtings frquntly dalt with quntitativ
rcs.Sitifimeri, otor:
rvolutionaryrsponssto
,oltion49:241-251.
Rditlttlf Dui's Finhes. aftr h had rd Darwin's tbe rigi of S1leies.Baus donrsti animls
bridgUnir,rsityPrss.
d Sic tl th Gldpgos
;les.Nr,l,York: olumia
Glton 597
lvill,H. 1987.Th nntadas, or nhzrntlIsls.In PizzTlsd tbr
PrrlsePis'189-1860:h Nrth'strtrNbrrditiltt<! tb \Yritigs
of rmluill.Vl. 9. . Ha1'fod,A. A. adougall,ad G. T. Tansll,
ds.vanston,IL,:NorthwstrnUnivrsityPrss.
Sulloway,F. J. 1982.Darwin nd his finhs:Th vlutionof a lgnd.Jlurlof
the istrof Biolog 15: 1-53.
.Winr,
J. |994, Th tskof th Fib: Stlrlf urllutioi ut,le. Nw
York: Knopf.
in ndlianppLrlations.
Wright'S. 1931.volr-rtilrr Gnetis16:97-159.
|932. The rllsof nrlttationrirlirrg,
rossrding nd sltirr
tn
volutin.Prldings of th Sitltl ongrssof Gtis1: 56_66.
-.R.
Galton, Franis (1'822_1911)
Franis Galton Was born nar Birmingh:rm,ngland. H originlly intndd
to pursu a arr in mdiin, ut h swithd to amridg Univrsity to
study mathmatis' inflund y his Ousin harls D;rwin (1809_1882).
Darwin and Galton wr grandsons y differnt marriags of rasmus Dr-
win (1731_1802),a physiian' sintist,po' nd invntor. harls Darwin
would a ruial influn on Galton throughout muh f his arr. In
fat, it was Galton's rading of o the o rigi of Speis (1 859) that gav him
th ida of improving tlr human ra throllgh sltivrling,for whih
h oind th trm ltniL.s.
Sevral yas aftr grduating from amridg with a ordinary dgr,
Galton rganizd and finand an Afrian pdition that rsultd in th first
uropan ploration of northrn Namiia. H made arful masurmntsof
latituds, longituds' and altituds, rflting his liflong intrst in th appli-
ation of numrial and quantitativ mtho.lsto whatvr happnd to intrsr
hinr. pulishd his rsr'rltsin tlourttl <lith Rrryl Gogrphil Soiet
in 1852 and ws wrdd a gold mdl by tlr soityth saryr largly -
aus of his pris, quantitativ work. Tropil South A|ri (1853), Galton's
ook aout his journy,Was a suss,tTbe rt of Trul (1855)' a guid-
ook for amatur and profssional alik who vnturd into th ush, provd a
triumph. It wnt through many ditions, growing iggr ah tim. In 2001
Phoni Prss rissud th fifth dition, whilr had first n pulishd tn 1872.
Galton Was an tiv nrmr of th Royl Gographial Soity fo many
mattrs (.g.,latituds,oiling point thrmomtrradings).H also am
intrstd in mtorology and disovd th antiylon, wathr fatur
haratristiof a high-prssursystm.
Th part of Glton.s arr for whih h is probaly st rmmrd gan
and ultivatdplants wr th prduts of sltivrding,it stndrson-
al to Galton tlr;rtth human ra -riglrt improvd sinlil:rrly. Th ky ws
to dmonstrat that dsiral human tits suh as intlltual aility wr in-
..Hrditary
hritd. H attmptd to do just this in a two-part artil ntitld
alnt and Charatr,'' pulishd in a popular priodial alld ill's
598 Glto

gzine (1865)' and in th ook that followd, Hrditr Gicts(1869). In th Unitd Stats,p
In both th artil and th ook Galton trid to dmonstrat that wht h r- involuntary strilizati
.h
frrd t as ..tlntand haratr'' wr inhritd. ok trad pdigrs dfiint or
..fblmi
of judgs, statsmn'and othrs, Galton's asi assumption was that if h followd with intrsl
.$hn
pikd an minnt judg, for xampl, tht judg'slosstmal rlativs(.8., th Nazis am
fathr nd son) had a gratrhn f birrgdistinguishdthan thos furthr that rsultd in th str
.World .!Var
rmovd (.g.,grndfathr' grandson). !7omn wr not inludd, proaly II, ugni
.!hil
rfltingoth Vitorian prjudi and ml dominan in Vitorian soity. urop. ugni
Galton onludd tht his rsults suppotd his hypothsis.Although othrs rditd with implrtal
pointd out that nvironmnt (.g.,th fathr'sgood position might nsura printing, mtorolgy,
plum jo for his son) <luldalso ndrli th orlation' Galton attmptd
|LIoGRAPY
t o d i s m i s sr h i s o n t n t i o n .
Familiar with th ll urv y th tim h wrote Hreditr Geius, Ga1- Brooks, NL.24.tr
ton alr.rltd hypothtial normal distriution for th estimatd 15 million Glto. Nw Yok:
Bulmr' |vl.20o3.Frn
mais in th Unitd Kingdom aording to thir natural ailitis.H also was
J o h n sH o p k i n sU n r v
anious to ollt rea| data that h ould analyz statistially, partiularly Forrst,D.W. |974. Frt
with rgard to inhritan.Baus this was not asy to do with human - York:Tp[ingrPuh
ings, h didd to us swt pas as a modl bausthey wre asy to grow Galton,F. 1852.Rnt
and supposdly slf-frtilizing. In his primnts h found that sd diamtr Jourl of th Ro
was normally distriutd, ut th sd diamtr fonr rogny of lag.sdd | 8 5 3 .T r p i Sl
|855 . b rt o1
and small-sddplants driftd toward th man f th population as a whol.
outris.London:
H alld this ..rgrssionto th man,'' a statistial proprty that has n r- 1865.rdita
patdly dmonstratd sir-r(.g., in th as of diffrnt lirsss of mutual 1'8_327.
funds, suh as ons spializing in growth vrsus intrntionl stoks). Galton 1869. Herditr
also disovrd that whn h graphd th diamtrs of parntal sds vrsus l889. NturlIr
21.Th rt
progny sds,th points fll on a stright lin. had drawrr th first rgrs-
outris.London:
sion lin and from it h alulatd th first rgrssion offiint. Latr, Galton G i l l h a m 'N . W . 2 0 0 1 .
otaind th hunran data h so dsird whn h orgnizd an anthropomtri Birth of ugzlis.N
laborator at th 1884 Itrnationl Halth xhiition lrld irr South Knsing- K v l s ,D . 1 9 9 . 5l. t h ]
ton' London. Th laortory ropnd in th Sin Gallris of th South Prss.
Knsington Musum aftr th hiition losd. ollting quantitativ dat Kiihl, s. 1'994'Th Nzi
Paul, D. B. 1995.otr
from oth pants and lrildrn, he was now abl to drnonstratgrssin
H i g h l n d sN. J : H u n
to th man for masurabl human haratristis (.g., hight). Y/hn h Parson,K.1914-1930.
plottd forarm lngth against hight h disovrd anothr irnpotnt statis- C m r i d g : m r
tial onpt' orrlation (i..' tll mn hav long forarms)' lulatd R i l l v .P . R . l 9 9 l . S
th first orrlation offiint, myriads of whih hav n omputd sin. tll UitdSttcs.B.
Galton lso am intstdin fingrprints and thir lassifiation; his work
was ntral to th dvlopmnt of fingrprinting as a fornsi thniqu.
Galton olltd many of his most important firrdings in Nturl Iheri-
Gam thory
tnc (\889). H also aquird his first protg6es'inluding Krl Prson. A Th study of nimal
fin nrathmatiian,Parson Wnt far yond Galton in formulating statis- t w ni n t u i t i o n a r r d
tial thory. anwhil, Galton was promoting ugnis nd th ida was invstigtion. fforts t
gaining tration irr the prvailing Soial Drwinist nvironmnt at th nd of al tools alld gams
th nintnth ntury. Positiv ugnis nouragd th sltiv rprodu- A gam in this sns
tion of th supposdly fit, whil ngativ ugnis aimd to liminat th - ing whn th outom
prodution of thos dmd unfit. Unfortunatly, ngzrtivugrrisprvaild. oths. hus gam l

'd, Hrditr G eius (1869).
dmonstrattht what h r-
d.Th ook trad pdigrs
;i assumption was that if h
;'slosstmal rlativs (.B.,
stinguishdthan thos furthr
l Wrnot inludd' probably
r m i n a n i n V i t o r i a n s o i t y .
s hypothsis.Although othrs
lgood position might nsur a
o r r l r i o nG, a l t o n a t t m p t d
rote rditr Gis, Gal'-
n for th stimtd15 million
r natural ailitis. LI also was
alyz statistially, partiularly
lot asy to do with humn b-
)austhy wr asy to grow
rts h fund that sd diamtr
r frm progny of lag_sdd
Lnof tlrpopulation as a whol.
;tial proprty that has n r-
{ dif{rntlsss of mutal
us intrnationl stoks). Galton
mtrsof parntal sds vrsus
. H had drwn tlr fist rgrs_
ssionoffiint.Latr, Galton
l orgnizdn inthropomtri
hiitionlrld in South Knsing-
l Sin Gallris of th South
d. llting qllantitativ data
a l t o d m o n s r r r r g r s s i o n
ristis(.g.,hight). Whn h
|vrdanothr impotnt statis-
long forarrs).H alulatd
rih hv n omputd sin.
rnd thir lassifiation; his work
ng s a fornsithniqu.
'ant findings in Ntrl lheri-
g6s,inluding Karl Parson. A
d Galton in formulating sratis-
ting ugnisand th ida was
linist rrvironmntat th nd of
ourgd th sltiv rprodu-
nis aimd to liminat th r-
ly' ngativugnis prvaild.
Gme heor 599
In th Unitd Stats, passge of state ugni strilization laws rsulted in th
involuntarystrilization of thousands of individuals rgardd to mntally
..flmindd.''
defiintor ugni dvlopmnts in th Unitd Stats wr
followd with intrst in urop and lswhr' partiularly in Grmany.
!hn th Nazis m to powr thy pssd n involntry strilization law
that rsultd in th strilization of hundrds of thousands of individuals. Aftr
!orld !ar II' ugni strilization gradually ndd in th Unitd Stats and
urop.rWhilugnisis Galton's most wll_known lgy, h also must b
rditd with important ahivmnts in filds s divrs as statistis' fingr-
printing, mtorology' and xploration.
BlLIoGRAPHY
Brooks'I\^.2004.trmsures:The Drk Visiosnd Bright Idsof Fris
Glto. Nw York: BlomsburPulishing.
Bulmr,. 2003. Fris Glto: Pioerrfrditvd Biontr.Bltimr:
Johnsopkins UnivrsityPrss.
Forst,D.w.1974. FrltisGltl:Th Lif d'V/<lrhof Vit:torinGius.New
York:TplingrPulishingCo.
Glton,F. 1852. Rntpditioninto th intrior of South-IWstrn Afria.
Jornl of tb Rol Ceogr1lhilSliet22:140-76.
1853, rlpilSouthfri, London:John urray.
1'855.Thrt of rul:r, Shiftsnd otriusuilblin Wild
otris.London:John urray.
1865.rditrytalntnd hartr ' mill'sgzi|2: 157-166,
t8-27.
1869. HrditrGeius' Londn: amillan.
1889.Nturl lherit.London: amillan.
20|. b Art of Trul,or, Sbiftsd otritlsuilblei WiId
outris.London: Phoni Pss.
Gillham,N. !. 2001. Lif of Sir Fris Glto: Fr-lmfrin plortio to the
Birth of gzis.Nw York: oxford l-Inivrsityrss.
Kvls,D' 1995.In the Nme of ugics.mridg,A: Harvard Univrsity
Prss.
Knhl, s. 1994. Th Nzi otloz.Nw York: Oford UnivrsityPrss.
Paul,D. B. 199.5.otrolligHutn rdit, 186.\to th Prst.Atlnti
Highlalrds,NJ: HumanitisPrss.
Parson,K. 1914_1930,The Lif, Lttrsd Lb<lursof Fris Glto, vo|s.
amridg:amridgUnivrsityPrss.
Rilly'P. R. 1991.Th SurgiISolutio: istorof Iuo|utr Steriliztitl
i
the Uitd S/a/es.Baltimor: Johns opkins Univrsity Prss. -N.vG.
Gam thory
Th study of animal havior thrivs n paradoxs, affling inonsistnis
btwen intuition and vidn that ngg attntion nd stimulat furthr
invstigation. fforts to rsolv suh parados rly inrasingly on analyti-
al tools alld gams.
A gam in this snsis a mathmatial modl of stratgiintration, aris-
ing whn th outom of an individual's ations dpnds on ations takn y
othrs. Thus a gam has thr ky omponnts. Fist, thr ar at last two
Gme Thor

intrating individuals, alld playrs. In an volutionary gam' th st of play-

The centicl Th
rs is an otyp' tht is, a population of animals in a givn ologial nviron-
mnt (for ampl, a populatin of spidrs in a grassland haitat and a riparian Sir R. A. Fishr's Te
population of th sam spis would form two diffrnt otyps). Sond, l 9 3 0 , i s l da s t
ah playr has a st of fasil stratgis tht is onstraind y th inform- ltion and Mndlian
tion strutur of th intration (for ampl, animals an ondition thir - Fishr from Fishr's di
havior on whthr thy ar ownrs or intrudrs only if thy ar awar of suh p r o l m s o f g n t i sa r
rols). Third, th pattrn of intration must wll dfind and aompanid Th first svn hap
by a formula for how ah playr's rward from th intration dpnds on its D a r w i n ' s m h a n i s mt l
stratgy and on thos of th othr playrs. In volutionary gams, th rwards sidrd th first two l
ar masurd in trms of xptd futur rprodutiv suss. mntal thorm of natu
For a gam to usful, it must possil to idntify a stratgyor strat- ths two haptrs
gis from among thos fasil as th ..solution'' for a givn purpos. In an F i s h r ' sf i r s t h a p t
volutionary gam' this is th havior xpted to volv y natural sl- tion with, altrnriv
tion. If a havior is fixd in a ral population, thn it must at last tru strats that on th Mn
that vry fasil altrnativ havior would yild a low rward, othr- o f a p o p u | t i o n ' sv a i
wis th altrnativ havior would hav sprad into th population. Thus sistnt prolm for D;
th rlvant solution onpt-introdud by John aynard Smith (1982)- had ld iologiststo ab
is tht of an volutionarily stal stratgy (SS), a population stratgy that w i n . s a p t a n o f l
yilds a highr rwd than any fasil mlltant stratgy (s also Dawkins trolIingmuttionb
r976). l n d i n gr h o r y .B a u
Sintistsstriv to unravl a parado y stalishingonditions for an SS mutation rats' Fishr
to ist in a modl population with assumd omponnts, and y analyzing viv naturl sltion
its proprtis whn it dos eist. If th SS in th modl population fails to F i s h r ' ss o n d h a ;
math obsrvd havior in th ral population, thn on or mor of th as- natural sltion.Thr
sumptions ar modifid-aout otyp' information strutur and stratgy m a t h m a t i s . h f i r s
..tr4althusianparamt
st' of pattrn of intration and rward. Th SS is ralulatd as oftn as
nssary. In othr words, if a parado of animal havior ists, thn w of thir lif historis.l
hav wrongly gussd whih gam st modls how a ral population intr- Fishr partitions into
ats. T rsolv this parado' W must guss again-if nssary,rpatdly- masur of th rat of
until vntually w guss orrtly. natural sltion.For l
Gms ar valual austhy allow sintiststo plor th logi of a qual to th [additiv]
vrbal argumnt rigorously, ssuming iologially ralisti otyps' and to Fishr's ..fundamntal
dtrmin whn it is tru and whn it is fals. Gam thory oftn dmon- h i s n t u r a l s l i o nr h
strats what is diffiult to intuit. Fo xampl' gam thory has shown that l p r o o f t h t u m u l a
vitory y strongr animals ned not imply that strengthis ing assssdin naturl sltionon mr
ontsts' and has yildd a tst for suh n SS, namly, whthr nrgy r- B y n d l a r g .t h m i
srvs of losrs orriat positivly with ontst durations. For furthr dtails a s s ,s u h s d o m i n a
of this ampl and othrs, s strton-Gions and Adams (1998). ding thortial worl
pands his thorti
BIBLIOGRAPHY
U s so f g n t i v r i a
Dawkins, R. 1976. The SelfishGe. oford: Oford UnivrsityPrss. Th last fiv hpr
l4aynard Smith,J. 1982.uolutild th Tbor of Gms.amridg: h u m n p o p u l t i o n s 'p
amridgUnivrsityPrss.
n t r a l o s r v t i o ni s
strton-Gibbons, ., and . S. Adms.1998.Animal ontstsas voluttonary
gams. Americ Sietist 86: 334-341. -NI..-G. t i ss t u t u r st h i r
I n t h f i n a l h p r r .F i

volutionary gam' th st of play-
nals in a givn ologial nvirn-
a grasslandhaitat and a riparian
two diffrnt otyps). Sond,
Ltis onstraindy th informa.
, animalsan ondition thir -
Lrsnly if thy ar awa of suh
wll dfind and aompanid
.om th intrtion dpends on its
r volutionarygams' th rwards
'rodutivsuss.
ll to idrrtif)ra stratgy or strat_
rtion'' for a givn purpos. In an
ltdto volv y natural sl-
tion, thn it must at last e tru
uld yild a lowr rward, othr-
;pradinto th population. Thus
ryJohn Maynard Smith (19tt2)-
(SS),a population stratgy that
tutantstratgy(s also Dawkins
stlishingonditions for an SS
d omponnts'and by analyzing
i in th modl population fails to
rtion, thn on or mor of th s-
rformatin strutur and stratgy
]h SS is ralulatdas oftn as
I animal hvior xists, thn w
tdlshrv Ipopulation int-
; gain-if nssary'rpatdly-
sintistst plor th logi of a
ogially ralisti otyps' and to
fals. Gm thory oftn dnron-
rpl, gam thoy has shown that
. that strngthis ing ssssdin
r SS, nml, whthr nrgy r-
ltst durations.Fr furthr dtails
Gions and Adnrs (1998).
)xlrdUnivrsityPrss.
orof cms.mridg:
i. Anilrrrl
ont\ts
...
"""'Ifi]xZ-".
Th GntialThory of Natural Sltion 601
The Geeticl Thor of Nturl Selection (R. A. Fishr)
Sir R. A. Fishr's Tb Gentil Theor of Ntrl Selectio, pulishd in
190, is latdas th fist m;.orrorriiiationof Darwiniirn natural s-
ltion and Mndlian hrdity.Th 12-hptrmanusrtpt,ratd y Ms.
Fishr from Fishr's ditation, was th ulmination of Fishr's thught on th
problnrsof gntisand ntural sltiongun in th lat 1910s'
Th first svn haptrs of cetil Tbeor st out Fish's synthsis of
Darwin nrhanismof natrl slrionand ndlian gntis.Fishr on-
sidrdth first two hapts' on th natur of inhritan and th ..funda-
mntal thorm of natual sltion,'' th most important of th ook. [ndd,
thstwo haptrs aomplish th k pi of th roniliation.
Fishr's first haptr onsidrs impliations of a synthsis of natural sl-
tiorr with, altrnativly, leding nd ndlian inhritn. H dmn_
stratsthat on th ndlian thory' naturl sltionmay th min us
of a population's variaility. Th dmonstration importantly rsolvd a p-
sistntprolm for Darwin's thory of dsnt with rnodifiation, on that
had ld iologiststo andon ntural sltionas an volutionaryaus.Dar.
win aptnof blnding inhritnrquid him to imgin usson-
trolling mutation aus of normous mutation rats dmandd y th
lndingthory. Bausndlian hrdity did not dmand suh normous
mutation rats, Fishr was al to limint ths ontrolling auss and r-
viv natual sltionas an important volutionary us.
Fishr's sond hptr dvlops, mathematilly, his gntial thor of
natural sltion.Thr ky lmntsmay distilld from Fishr,s ..havy''
mathmatis.Th first is masur of avrag population fitnss: Fish's
..lthusian
paramtr,''th rprodutiv valu of all gnotypsat all stags
of thir lif historis. he sond is a masur f variation in fitnss,whih
Fishr partitions into gnti and nvironmntal omponnts. The third is
masurof th rat of inrasin fitnss,that is, th hang in fitnssdu to
natural sltion.or ishr, ..thrar of inrasof fitnssof any spisis
qual to th [additiv]gntivarian in fitnss'' (p. 35).This last lmntis
Fishr's..furrdamntalthorm o1 natural sltion,'and is th ntrpiof
his ntual sltionthory. Under this ruri, Fishr thn offrs a gomtl-
al proof that umulativ volution is primarily th rsult of low prssursof
natural sltionon mutations of small fft-
By and larg, th middl haptrs of Geticl The>rar plorations of
ass,suh s dorninan,sual sltin,nd rnimiry' tO suport th pr-
dingthortialwork' Nvrthlss,in th fourth and fifth haptrs' Fishr
pands his thortial disussion to mor grrral issus otlrning th
aussof gntivariation' irriudingrndom gnti drift.
Th last fiv haptrs of Getil Thor plor natural sltion in
human populations' partiularly soial sltiorrin lruman frtility. Fishr's
ntral osrvation is that th dvlopmnt of onomis in humn soi-
tis struturs th irth rat so that it is invrtd with rspt to soial lass.
In th final lraptr,Fishr offrs stratgisfor ountring this fft.Dspit
602 The Gnti Basis of vlutionaryhang

Fishr's spousal of this ugnis thsis in this part of th ook, h intnds th Having argud that
disussion to takn as an insparal tnsion of th prding part. Lwntin advans an
Tbe Getil Thor of Nturl Seletio is a point of dpartur in volu. tis is invitale. The
tionary thought, rsponsil in part for th origintion of thortial popula- rsult of a dpr,irr
tion gntis and what is ommonly alld th modrn synthti thory of and nutralist position
.!right's
volution. Getil Theor was followd y Swall and J. B. S. aility; thir fondnss
aldan's major works in 191 and 1,92,rsptivly. Fishr's viws on th pting r rjtingv
rol of natural sltionin volution ar widly aotdtodv. mrad hang and
ativly maintaind
|LIoGRAPHY
hld onvitions rgar
Fishr,R. A. 1930 [1958].h Geticlhorof NturlSltio.2nd d.Nw his ltrophorti dat
York: Dovr Publiations.Rlasdin a variorum dition by oford Univrsity inspird its transform
rssin 1999, editedy J. H. Bnntt.
Tbe Geti Bsis o1
Haldan,J. B. s. 192.The ussof uoltion.London:Longmans.
.Wright, duing volutionary g
s. 1931.volutionin ndlianpopulations.Gtis16:97-159.
-R..s. arful analysis of th .
apturs how gnti 1
Th Geneti Bsis of Euoltionr bge Lwontin, informatio
ody of data that wou
(Rihard Lwntin)
and rfind. In light of
In1974, Rihard Lwontin, a profssor at Hrvard Univrsity, pulishd an rlationship btwn t]
important assssmnt of th stat of volutionary gntis. Lwontin's 1974 tion, Lwontin argud
ook,The Geti Bsis of uolutior hge, onsidrd th trmndous plad with modls th:
hangs within evolutionary gntis wrought y th introdution of th- Indd, Tbe Gntic B
niqus from molular iology and th susqunt dvlopmnt of th nu- th ral prolms of
tral thory of molular volution. Lwontin's ook is notworthy for its standing th intrativ
analysis of th natur of volutionary gntis and th dynamis of sintifi
ontfovrsy and ontrovrsy rsolution. lLIoGRAPHY
Lwontin and his ollagu, J. L. Huy, hd touhd off a molular rev- Ditrih'. R. 1994.Th
lution in 1966 whn thy introdud th biohmial thniqu of l- lourl of th Histo
trophorsis as a mans of rsolving gnti diffrns at th molular lvl. ubby, J. L., and R. . l
htrozygosity in na
Lwontin livd that this nw ltrophorti analysis of gnti variaility
Drosophil psudoo
rsolvd th arlir disput twn th lassial and balan positions ovr
Kimura,. 1983.TeN
variability and sltion. Th high lvls of variaility dettd with l- ambridgUnivrs
trophorsis smd to support th alan position and th importan of Lwontin,R. . 1974.t
aland polymorphisms in volution. The lassil position's lif was that olumbia Univrsit
most mutations wr dltriousand most wr sltionpurifying. Thy strug- Lwontin'R. .' ndJ. L
htrozyglsityin n
gld to plain ltrophortivariaility until otoo Kimura and othr i-
htrozygosityin na
ologists gan to argu that most of th dttd polymorphism was in fat 542595-609.
nutral, nithr sltdfor nor against. In b Geeti Bsis of uoltior
hge, Lwontin mphasizd th ontinuity twn th arlir lassial-
alan ontrovrsy and th thn-raging nutralist-sltionist ontrovrsy.
Geetics nd the I
For him, th nutralist position was rally just th lassial position rought in
(ThodosiusDoz
lin with nwr moleular rsults. Whil this intrprtation of th ontrovrsy
maks a god as fo ontinuity in volutionary gntiS, it nglts th With grat harisma a
growing ontriution of biohmial rsarh to th dvlopmnt of molu- am a rallying poin
lar volution. gi of Spies (1'937\l

part of th ook' h intnds th
rsion of th prding part.
is a point of dpartur in volu-
.iginationof thortial popula-
:h modrn synthti thory of
.t/right,s
ly Swall and J. B. S'
sptivly.Fisher's viws on th
:ly aptdtoday.
f NturlSIctio.2ndd. Nw
' r u m d i t i o n y o f o r d U n i v r s i t y
London:Longmans.
n s .G t i s1 6 29 7 _ | , , - * . o . , .
hnge
lrvard Univrsity' publishd an
)nry gntis.Lwontin's 1974
ge, olsiderd th trmndous
;ht y th intrdution of th-
qunt dvlopmnt of th nu-
in's ook is notworthy for its
s and th dynamis of sintifi
ad touhd off a molular rvo-
iohmial thniqu of l.
Liffrnsat th molular lvl.
:ti analysis of gnti variaility
;sialnd lan positions ov
f variaility dttd with l-
position and th importan of
:lassialposition's lif was that
r sltionpurifying. Thy strug.
rtil Motoo Kimura and othr bi-
:tdpolymorphism was in fat
,he
Gnetic Bsis of uolutior
ity twn th arlir lassial-
utra1ist-sltionist ontrovrsy.
;t th lassial position rought in
intrprtationof th ontroversy
rtionary gntis' it nglts th
:h to th dvlopmnt of molu-
Gntisand th origin of Spis 60
Having argud that on ontrovrsy has n transformd into anothr'
Lwontin advans an plantion for why ontrovrsy in volutionary gn-
tis is invital. Th onntion and prsistnof ths ontrovrsis WaS a
rslt of a dpr, irrsolval idologial onflit. Advoats of th lassial
and nutralist positions hld a onsvativ attitud toward hang and vri-
aility; thir fondnss for staility ld thm to s sltion as a mans of a-
pting or rjting variations. Advots of th alan position' howvr,
mbradhang and variaility; thy favord th ida that variation ould
ativly maintaind in a population y alaning sltion.Ths dply
hld onvitions rgarding th valu of hang xplaind for Lwontin why
his ltrophorti data did not rsolv th lassial-balan ontrovrsy, but
inspird its transformation.
Th Geti Bsis of uolutionr hng should not b undrstood as r-
duing volutionary gntis to onfliting idologis. Lwontin prsnts a
arful analysis of th hllngs of produing a thory of gnti hang that
apturs how gnti prosss intrat and dvlop ovr tim. Aording to
Lwntin, information aout molular level hangs was finally providing a
ody of data that would allow thoris of volutionary gntis to tstd
and rfind. In light of nw molular data, Lwontin laimd that th ntir
rlationship twn thory and fat would hav to ronsidrd. In addi-
tion, Lwontin argud that thoris of individual gns would hav to r-
plad with modls that onsidrd gnoms and thir ompl intrations.
Indd' h Gtic Bsis of uolutionr hge nds with th harg that
th ral prolms of volutionary glltis an only addrssd y undr-
standing th intrativ ontt of allls.
ILIoGRAP|.]Y
Ditrih,. R. 1994. Th originsof th nutralthoryof molulrvolution.
Jourl of th l{istor of Biolog 27: 21_59.
Huby,J. L., and R. . Lwontin.1'966.A molularapproahto th studyof gni
htrozygosity in naturalppulationsI. Th numrof alllsat diffrntloi in
Drosophilpsudoobscr. Gtis54 577_594.
Kimua,. 1983. Tb Ntrl Thor of olulr uollltio. amridg:
amridgUnivrsityPrss.
Lwontin,R. . 1974. h Gti Bsisof uoltiorhge.Nw York:
ColumiaUnivrsityPrss.
Lwontin,R. ., and J. L. uy.1966. molularapproahto th studyof gni
htrozygosity in naturalpopulations.II. Amount of variationand dgrof
htrozygosityin naturalpopulationsof Drosophil psdoobsur.Gtics
54:595-609. -.R.D.
Genetis nd th rigi of Specis
(ThdosiusDozhansky)
.!ith
grat hrisma and a passion fr th subjt' Thodosius Dozhansky
am a rallying point for th volutionary synthsis. Getis d the ori-
gi of Speies (197) was his prinipal ontriution to that movmnt' and
604 Gntisand th Origin of Spis

it am an ssntial tt in volutionary iolgy. On on |evel, Getis H o w p o p u l a t i o n sb t

is a splndid summary of volutionary gntis in th 1930s' oth zoologi- th l930s. any disi
al and otanial. It rought togthr divrs information and ratd an mhanisms'' wr pr.
asy-to.undrstndmodl for voiution lading to nw spis.It also pr- hanial' physiologi
sntd an unxptd lvl of sophistiation and orrfidnto iologists i n t h a t d i s u s s i o n .G e
suspiious of volutionary studis. Lik no othr ontriution to the su- t a n o f g o g r a p h i a l
ject,,Getis gave a urst of nrgy to spiation studis as a resarh pro- twn two spis srv
gram. hapter in Getics is
i\4uh of Getis survyd knowldg aout th gnti strutur of pop- ssslos off th tap
ulations. Rent dvlopmnts in tehniqu had givn gntiistssnsitiv FoI|owing \0righ.l
tools for monitoring nW mutations, hangs in gn frquny, and hangs gn largly in trms
in hrmosom strutur. Dozhansky prsntd onsidrabl data on th th partiular alan
xtnt of this variation in oth laoratory and wild populations. H also d- adaptationist. vn wh
srid xprimentl systmsthat monitord hangsto th gntistutur h e t h o u g h t s l t i o nw :
of populations ovr tim and ovr hanging nvironmntal onditions. In tnsity. and it frqu
part, Dozhansky's survry lratdth hard-arndmpirial sid of pop- Dozhansky prsnt
ulation gntis.It also aimd to prsntth sujtas rigorous, with th vo. p s s u r sh a d nd i s
lutionary sintistsin full primntalontrol of thir data. spiation. Famously,
Dozhansky dfins volution in narrow trms: a hng in th frquny though it was trnal
of allls within a population. This is purposivly oprational. ceetis of- uls faild to rd so
frs many xampls of suh a hange.It also prsentsa dtailed disussionf ing a nw speis.
mhanisms driving thos hangs.This disussion,along with th latr dis- In th widr fram, J
ussion of isolation and divrgn,mak for th two thortial spins in volution within spi
Geetis. Dbzhansky argud natural sltion drivs adapttion and is a rprodutivly isolat
prinipal agnt in shifting th gnti strutur of populations. Sltion uss goris (.g.,nw gn
th raw matrial of gnom variation rsulting from random mutation as wll additional mhanism
as from th many othr gnti and hromosoml posssat Work within sss of mirovolutitl
organisms. Ky to Gntis's tm
Although potnt' sltionwas not Dozhansky's only volutionary mh- and mpiril studis
.sright's..shifting
anism. H was hvily inflund y Swall alan'' th- priod's pproah to
.Wright's
ory of population gntis and notion of adptiv landsaps (s q u s t i o n s n d u i l t a r
\iright 197, 192). !right's idas als mphasizd population siz and this with los tis to r
strutur' migration, mutation prssur,nd han. uh of a whol hap- I n p a r t . D o z h n s k yr
tr in Geetls is dvotd to dsriing Wright's approah. In fat, many vo- tal iologistsattaking
.Wright's
lutionary iologists larnd aout modls from rading Gentis. skptis from othr d
For Dobzhansky, shifting alan nd adaptiv landsaps Wer ssntial Sympathti ollagu
tools for xplaining th gntisof volution. tionary studis.Th pr
Spiation involvs mor than simply hang in alli fqunis. For i U n i v r s i r 'G t i
Dobzhansky, it rquird isolation and divrgn.This is th sond thort- o m n o u t | tf o r o
ial spin in Getis. Dozhansky amind oth prosssin dtail lading iology, that thory ur
to a dfinition of spisas rprodutivly isolatd and physiologially ina- 199).
pal of intrredingwith othr groups. on isolatd' divrgnrsultd Dozhansky pulish
from oth dirtional and random prosss.Dozhansky argud tht in th i n | 9 4 l n d a t h i r di n
ontinuum of lif, spisWr th only ..natural''units. Thir uniqu gnti w s i n t n d da s G
onfiguration, isolatd and following a puliar dstiny' Was ntral to that s i g n i f i n t h a n g st
thinking. s y s t m t i h a n g s l :
 Gntisand th origin of Spis 605

llogy. On n lvl, Getis How populations bam isoltd was a sujt of onsidral study in
s in th 1930s' oth zoologi- th 1930s.any disiplinsontriutd to that sr.rjt' and many ..isolating
: informatiorr and rtd an mhanisms''w proposd ('g., gographi, ologial, hvioral' m-
lg to nw spis.It also pr- hanial,physiologial). Anything tht prvntd gnti hang ountd
and orrfidnto iologists in that disussion.Getis survysprominnt ampls' strssingth impor-
rhr ontiution to th su- tan of gographi and physiologial mhnisms. Strility in hyrids -
'ion studisas a rsarh pro- twntwo spissvd as spial as study in this disussion. A whol
haptrjn Getics is dvotd to this topi, illr"rstratinghow isolting po-
t th gnti strutur of pop- ssslos off th t;rp of gn flow.
.!7right's
l d g i v n g n t i i s t ss n s i t i v Following shifting alan thry, Dozhansky plaind divr-
r gnfrquny'and hngs gnlargly in trms of sltion and drift, with irumstans influning
rtd onsidrabldata on th th partiulrrlan of fors. Impotantly, Dozhansky is not a kn.jrk
wild populations.H also d- adaptationist.vn whn it plyd a dtrminant ol in shaping organisms'
hangsto th gnti strutur h thought sltionwas many layrd.It frequntlyshiftd dirtion and irr-
nviroIlmntalonditions. In tnsity,and it frqrrntlyprssd populations in ontraditory dirtions.
-arndmpirial sid of pop- Dobzhansky prsntd xprirntal snarios in whih suh ontaditory
jtas rigrous,with th vo- rssLrrs hd n distinguishd.Dozlransky did not rqr.tirdivrgnf<lr
l of thir data. spiation.Famously, h namd Drosophil mird as a nW speis vn
.ms:a hang in th frquny though it Was xtrnally idntial to D. psedoobsur.Inthis as' individ-
ivly oprational. Getis of- uls faild to rd so th asnof frtil offspring was sr-rffiint for nm-
rsntsa dtailddisussion of ing a nw speis.
ssion,along with th latr dis- In th widr fram, Dozhansky largly rstritdhis fous tn Geetis to
r t h t w o t h o t i e sl p i n s i n volutionwirhin spisand to th form:rtion of spis,in his strit sns<lf
on drivs adaptlrtion and is a rpodutivlyisolatd omrunitis. To plain th oigin of highr t.
: of populations.Sltion uss goris (.g.,nw gnra' familis, and lasss),h saw no rson to invok
from ndom mutation as wll additional mlranisms!suggstingmrvolutiorrmrlv tndd th po-
mal prossst work within sssof mirovolutior.
Ky to Ceetics's impat was Dozhansky's <rnrnfor ojtivity,rigor'
nsky'snly volutionary mh- and mpirial studis. His ook ws old ssrtion <l[onfidtrin th
..shifting alan'' th- priod's approah to volutionary studis. Getis askd narrowly dfind
right's
n f adptiv landsaps (s qustionsand uilt arfully on thik vidnbas.Dobzhansky omind
rphasizdppr.rltionsiz and this with los tis to rnthmatilmdls nd rigorus primntal tsts.
:han.Muh of a whol hap- I n p a r t , D o z , h a n s k yw r o t a s a r p r s n t t i vof n w w v o f p r i m n -
''sapproah.In fat, many vo- tal iologists attaking volutiorrry topis. H also wrot with an y towad
nodls fronr rdirrg Getis' skptis from othr disiplins nious to avoid unsavory ompliations.
tiv IandsapsWre ssntial Sylpathtiollaguspointd to Getis as a sign of a nw r in volu.
tionary studis.Th produt of Dozhansky's 196 Jsup ltursat olum-
rng in lll frqunis. For bia Univrsit, Getis rvivd the olumia Biologial Sris,whih would
n.This is th sond thort- boman outlt for othr ky ooks in th making of modrn volutionary
th prosssin dtail lding iology' that thory usually rfrrd to as th volutionry synthsis (Cain
'latdand physiologially ina- 1993\.
isolatd,divrgnrsultd Dobzhansky pulishd a sond dition of Gtis d tb rigi of Spcis
Dozharrskyargud that in th in |94| and a third in 1951. Flis Geetis of the uolutior Procss (1970)
l . .u n i t s .T h i r u n i q u g n t i was ir-rtnddas Gtis's fouth dition' ah nw dition inludd sor
iar dstiny, was ntral to that signifiant hangs to Dobzhnsky's viws. In partiular, finding signifiant
systmatih:-rngs of hromosoml vrizrtionsin th wild, Dozhal-rskygrw
(t06 c offro Sit- ilir

to s a muh strongr rol for sltion in th volutionary pross. Using h i s r p a t dI i p s r v i

on of his powrful mtaphors' StphnJay Gould (1983), no grt lovr of or priphral naturalis
univrsal adaptationism,rfrrd to this as th ..hardning''of th synthsis, politial onsrvatism.
although othrs (for ampl, Rus 1999) s it as il natural dvloprnntof Buffon's hints onrn,
a sintifi thory in th fa of nw vidn. ivd vrtbratsas (
tims lmost invisil
|BLIoGRAPHY
ratd studis on th v
ain, J. A. 1993.ommn rllrns and ooprativ srllutions:
organizrtionl pos mammalian aud
ativityin volutionarystudis|96_\947./sls84: 1_25. Hgl and provd' in L
Dobzhansky,T. 197. Gtisd tb rigi tlf Spies.Nw York: rllumia
ogy stalishd y Ril
UnivrsityPrss.(Sonddition] 941, third dition1951.)
-. 1970' Ctislf th urllutilrPrtlss.Nw Yrk: ColumiirlJrrivrsit mhanil onstrint
Prss' for th diffnt..|
Gould, S. J. 1983.Th hardninglfth synthsis. In M. Grn,d.,Dimsiotlsof a n d v n l s s s .D i f f
Driism.amridg:amridgUnivrsityPrss. diffrnt animal form
Rus, . 1999. str of 1'stris: Is uolutio Soil ostrutil? 1 8 2 0 s a n d 1 8 3 0 sh t
amridg,A: HarvardUrrivrsity Press.
ditions mightmhan
Wright,s. 1931.volutionin ndlianpopultions.Gtis16:97_I59.
|932.Th rolsof mutation'inrding, rossding gan dispositions, mod
and sltintn
volution.Prodigsof th Sith Itrtiologressof Gtis|: thortial oldnssm:
56_366. -I.. all animl forms. His
vrtrats gav uvi
h a t d ,f a m o u s d i s p u t
Goffroy Saint-Hilair,tinne (1772-1844) s i n si n l 8 3 0 . A I r h
tinn Goffroy, known as Saint.Hilir, was orn in tamps on April 15' Wnt to Cuvir. many I
1772. Sent to Paris to study for holy lrdrs,h ntrd mdial studis in ordr anatomy.
to fulfill his sintifi amitions. Plad undr th patronag of th rystallog-
B|LIoGRAPHY
raphr Rn6-JustHaiiy, whom h savd from almost rtin dath in Sptm-
r 1'792, in Jun |79 Geoffroy Was pplintd profssor of zoology at th App|,Toy A. 1987.Tht
bforeDuiz. oftl
newly ratd us6um n:rtional d'histoir naturll, not without som latd
o r s i .P . | 9 8 8 .T h A g -
opposition. |n 1795 h invitd Gorgs uvir to shar his own aprtmnt in Brkly:Univrsityr
Paris; thy would writ fiv innovativ joint ssays.Unlik uvir, who stayd Goffroy Sint-Hilair,.
hom to ttr his arr' in 1798 Goffroy aptd th invit:rtion to join th arvis.
pdition to gypt organizd y Gnral Napll6on Bonapart. His work on L G u y d r . . 2 0 0 4 , i ,
th fauna of th Nil rgion ws notd in Fran, as Wr his privatl Ommu- Nturlist.hiago:l
niatd spulations on lif and ltriity, inrasingly suspiious in th on-
srvativlimat lading to th Napoloni mpir. Bk to aris in 1802,
Gillspi,JohnH.
shortly to lav again for Spin and Portugal, h was adly lagging hind
uvir. H was mad a mmr of th Institut only in 1807 nd was appointd John Gillspi has n
profssorat th nwly ratdsinfaulty in Pis in 1809. Th ulk of his attrns of molular
major work was pulishd aftr 181.5,nd during th 1820s Goffroy ld th sltion in varial n
sintifi opplsition to th onsrvativ govrnmnts in powr, of whih u- w o r k h a s i n l u d dt h
vir was a suppotrand t tims a rprsntativ. During th 180s lris prsis- d s r i n t u r I s l t
tnt ttaks on his dasdformr frind, his mystial Ovrtons'and his v r i o u s p a t t n so m o |
philosophial prtnsionsontriutdto his isoltionwithin th Prisian sin- ditions of thos' and o
tifi ommunity, though his rptation was onsidral thrlughruturop. A studnt of Knil
Goffroy's Philoso1lhielto|11iqu (t818), his ulogis f Lmrk, and l970 fom th Univrs
 illespie 607

'lrvolutionar,vpross. Using his rpatdlip srvi to transformist thoris mad him hro of radiirl
o u l d ( 1 9 8 ) ,n o g r e t l v r o i or priphal naturalists who ausd Cr'rvirf prostituting his sin to
r ..hardnit-lg'' of th synthsis' politial onsrvatism. Dply inflund y IJauy's rystallography and y
: it as nturl dvlornnt of Buffon's hints onrning th unity of animal typs, Goffroy initially on-
ivdvrtrats:ls ing omposd of th sam st of organs and prts, at
tims lmost invisil in som fmilis, fully dvlopd in thrs. His l-
ratd studis on th vstigil prsnin rptils of all th parts that om_
ivsolutiotls: Organizrtionl pos mammalian auditory organs liitd th admiration of Goth and
ls 84:1-2-5. Hgl and provd, in his ys' th thory of analogus, or, in th trminol-
Spcis.Nw York: llr-r.ri ogy stalishedy Rihard own, homology. livd that strutural or
: l i r i o 1n9 5 1 . )
mhanial Onstr.lints during dvlopmnt miglrt hv b rsponsii
. . \ w Y l , k :( o l t r l n h i . l I l l i r ' s i r y
for th diffrnt ..alaning'' of various sts of parts in diffrnt farrrilis
In . (irn, ed.,Dimsios of and vn lasss.Diffrnt ratios of prssion of an organ or parts indud
ity Prss. diffrnt animl forms, thus produing diffrnt funti<lns. During th
l Soil ostrtio|1? 1820s rrnd 1830s h toyd with tlre ida that hzrngingnvironmntal on-
ditions might miranially afft th mryo, thus prodrring modilid or-
'>ns.Gtis | 6: 97_1 59.
gan dispositions' modifid funtions, and modifid animirls. His growing
:rossrdingand sltion rn
nl ogrss of Gtis 1z thortialoldnss md Goffroy xtnd his thory of th unity of type to
-J.. all animal forms. His laim that zrrthropodswr uilt on th Sam plan as
vrtratsgav uvir th opportr-rnityto ngg his f<rrn-rr frind in a
'2-1844) hatd,famous disput on th uniry of typ that startd t th Aadmi ds
sinsin 1830. Although historians hv rpatdly ssrtdthat vitory
s orn in tamps n April 15, Wnt to uvir, many in urop sidd with Goffroy and with philosophial
rrtd mdi[studisin lrdr antOmy.
th patronagof th r1,strrllog. BIBLIOGRAPHY
almost rtain dath in Sptm.
Appl, Toy A. 1987. h uuir-GeoffroDbte:reh Biolog in the Dds
rtd profssr of zollogy at th
bflrDri. Oford: oxford UnivrsityPrss.
urll,not withtrr"rt som ltd osi'. 1'988.Th g of Lmrk:uolutiorhoriesi Fr,1790-180.
r o s h e r h i s t , r v t t p r t m n ri l l Brkly:Univsityof lifornirss.
sas.Unlik ltvir,who st;ryd GoffroySaint-Hilai,. 1818.Pbilosophietlmiq.Paris:lvlquignon-
]ptdth invitation to join th Marvis.
pol6onBonapart.His work on L Guydr,H.2004. ti GoffrlSit.ilir,1772-1844: Visilr
Nturlist.higtl:Univrsityof hiagoPrss. -P..
.t .ls Wrhis privtlyomnltl-
:rasinglsusPii()usin tlr rn-
Flmpir.Bak ttl Paris in 1802, Gillspi,John H. (. L944|
l' h was adiv lagging hind
orrlyin 1807 an]ws appointd John Gillspi has n th most prominnt hi.rmpionof th argumnt that
i r r a r i si n 1 8 0 9 . h u l k o f h i s pattrnsof molulr volution r plaind bttr y th ation of ntural
rringth l820s Goffroy ld th sltilnin vrial nvionmntsthan y tlr so-alld nutrl thory. His
llmnts in polvr, of whih u- work has inludd th dvloprntof omplx mathmatial modls that
i v .D u r i n gt h I 8 0 sh i s p r s i s . dsri natural sltion undr varying onditions and th omparison of
his mystial ()vrtons'nd his vrious pattfns of moleulr vriation within and among spisto th pr-
l l t i t t rl ' i t h r h P a r i s i l r l rs i n - ditilnsof thos, arrd otlrr, modls.
l l s i d r h lth r r l u g ' h o uru r t l p . A studnt of Knihi Kjima, Gillspi rivd his dotoral dgr in
\970 from th Univrsity of Tas at Austin. approahd th qustion of
), his ullgisof Lamark, and
608 Gillspie

molular variation y onsidring rlativly simpl modls of how ntural

Goth, Johann.W
sltion might oprate whn som forms of a protin wr favord under
some nvironmntal onditions and othrs under diffrnt onditions, with Johann !olfgang von (
onditions hanging from on gnration to th nt. In this arly work' orn in 1749 in Frankfi
Gillspi gan xploring th onditions undr whih varying nvironmnts p i r . H o m p l t dD i .
.Wrthr)
ratd balaning seltion, form of sltion in whih fitnss diffrns io 1774. F
among allls that ontrolld protin variation ar anld out ovr th struggl with sulariza
long trm y th flutuations in nvironmntal onditions. At first glan, asts and roadns th
this anllation of fitnssdiffrnswould sm to dvolv into th nutral Goth's death in.Wim
thory, whih postulats that th majority of diffrns among protin Goth's rputation .
variants produ no diffrns or only slight diffrns in th fitnss of in- prominn as a natur
dividuals that arry thm. Although in som ass modls of varial sl- th produt of vrs' a
tion prdit th sam pattrns as do nutral modls, many of th long-trm l. Goth's approah
pattrns of gn sustitution ar diffrnt. In partiular, modls of varial the skelton and th
sltion mak diffrnt prditions aout th xistn of a molular as a whol' dspit th
lok, whih is th apparntly rgular pattn with whih on form of pro- knowldg of th parts
tin rplas anothr within a groupr suh as mmmals or irds, ov long th whol. In 1784, G
priods. Gillspi's work showd that whthr a lok mrgs dpnds formulatd a vrtrll
upon th pris distriution of fitnss diffrns and th timsal ovr skull ould drivd
whih on masurs sustitutions. Ths rsults stand in stark ontrast to Goth, ould rog
many aspts of widly aptd nutral modls, alling thos modls into urrd.
qustion. Goth's approah to
Bsids offring th most omprhnsiv altrnativ to th nutral thory, xpns of us form
Gillspi is rsponsil for a numr of othr insightful ontritions. a ull has bn givn l
has dvlopd statistial approahs to tsting hypothss aout volution- opd th horns that h,
ary rats for oth molular data and data on quantitativ haratristis, th tlologial approa
suh as ody siz or morphologial traits. Gillspi's plortions of fitnss izd th adaptationist's
disounting-how an individual might ompromis prforman in on on- ial trait in trms of sl
dition with th nd to prfom in many, variabl onditions-ld to a num. aout loating ral
r of advns y othrs in th ara of lif history volution in a varial manul Kant, who fam
nvironmnt. His studis of th volution of mutation and migrtion rats in gani natur who would
variabl nvironmntshngd iologists'viws of how volvalthos rats to tlologial languag
might b. tory lmnts ar to
Th highly sophistiatd mathmatis that Gillspi mployd has r- vr, distans himslf 1
stritd his radrship among gnral volutionary iologists, many of natur of an organism rs
whom may know him mor for his dlightful introdutory ttook on pop- nvrthlss planator
ulation gntis.owvr, his work on protin voltion is th major altr- human prin.
ntiv to nutral thory and, as suh, oupis a prominnt position in th Th kind of thing Go.
disiplin. th organization f plal
Th modifiation thr
BIBLIOGRAPHY
as diffrnt plants. H s
Gillspi,J.H. |991. T ussof olularuollttio.Nw York: Oford ing various plants as th
UnivrsityPrss. a paradigm of suh a pl
24.PopultilGtis: oisGuid' 2nd d. Baltimor:Johns tions of th laf: ..All tnl
Hopkins UnivrsityPss. -J.T'
th lot of thm suggs
lady, onvy to you in .

y simpl modls of how natural
lf protin wr favord undr
undr diffrnt onditions, with
to th nt. In this arly work,
d whih varying nvironmnts
tion in whih fitnss diffrns
rtion r anld out ovr th
:ntal onditins. At first glan,
smto dvolv into th nutral
y o f d i f f r n sa m o n 8 p r o t i n
ht diffrnsin th fitnss of in-
assmodls of varil sl-
l modls,many of th long-trm
[n partiular' modls of varial
t th xistn of a molular
:n with whih on form of a pro-
as martnalsor lirds, ovr long
Lthra lk mrgs dpnds
:frnsand th timsal ovr
sults stzrndin stark ontrast to
rodls'alling thos modls into
altrnativto th nutral thory,
'hr insightful lntriutions. H
:inghypothssaout volution-
l o n q a n t i t a t i v h a r a t r i s t i s ,
illspi'sxplorations of fitnss
promis prforman in on on-
rrialonditions-ld to a num-
history volution in a varial
I m u t t i o na n d m i g a t i o n r t s i n
iws of lrow volval thos rats
ihat Gillspi mployd has r-
zolutionary iologists' many of
rl introdutoyttook on pop-
ltin volution is th major altr-
'pisa prminnt position in th
|uolutiln,Nw Yok: oxford
)uide.2nd d. Bltimor: Jhns
-1.T.
Goth 609
.Wolfgang
Goth,Johann von (1749-|832)
Johann r/olfgangvorr Goth, wll known for his litrerrymastrpis' was
orn in 1749 in Fr;rnkfurt am ain, during th tim of th Holy Romn m-
'V/rtber
pir. H ompltd Die Leid ds juge (Th Sorrows of Yor.rng
Wrthr)in t774. Fust I, Goth's luminous, rih treatmnt of uop's
strugglwith sulaization,was first pulishd in 1808. Fust Il, whih r-
sts and roadns th pisods of Fust 1, was ompltd a eaf for
Goth'sdth in Winrar in 1832.
Goth'srputatiOn as a pot should not' howvr, lips his insight and
prominnas naturl sintist.Indd, Goth livd that sin was
th prodr'rtof vrs, and tht art nd th study of natur wr inspara-
l. Goth's approah to nrorphology is a fin as in point. For Goth'
th sklton and th parts of th organi body wr to lookd upon
as a whol, dspit th antomist's prolivity for disstion. Physioiogii
knowldgof th parts is dpndnt upon th (artist's)vision of th form of
th whol. In1784, Goth Wrot an ssay on th intrmaillary on and
formuiatda vrtral thory of th skull' maintaining that th ons of th
skull ould drivd from th vrtra. Th initial form, ording to
Goth, or"rld rognizd dspit rh onsidral hang tht had o.
urrd.
Goth's approah to morphology dos not mphasiZ usl filis at te
pnsof us formlis. Tlr produtiv pproah is not to maintain tht
a ull has n givn horns to butt, ut to ask how h might hav dvl-
opdth horns that h, in fat, so uss.Goth's rmarks, and his rjtion of
th tlologialpproah, to som xtnt ar prursory to thos who riti-
izd th adaptationist's program-t usinssof undrstanding vry iolog-
ial trait in trms of sltivprssurs.Nvrthlss, Goth is mor snguin
abotlt loting ral planatory lmnts in organi natur than, say, Im-
mnulKant, who famously dnid that thr would vr a Nwton of or-
gani natur who would plin vn a singl lad of grass without an appal
to tlolgiallanguag.Kant is, prhaps,orrt' if the appropriat plana-
tory lmnts ar to undrstood in trms of us |fiiens. Goth, how-
vr' distans himslf from this kind of viw y maintaining that whil th
naturof n organisrnis not xplaind ntirly in mhanial trms, th ar
nvrthlssplanatory forms that an apprhndd on th asis of
human xprin.
Th kind of thing Goth has in mind an illustrated y his thinking on
th organization of plants. Th laf, aording to Goth, is a asi organ.
Th modifiation throf issusin th various parts of a singl plant, as wll
as diffrnt plnts. sms to hav thought that by systmatiallyosrv-
ing various plants as th di1fringinstntiationsof a dvlopmntalpinipl,
a paradigm of suh a prinipl' on ould undrstand th possil prmuta-
tiorrsof th laf: ..All thir shapsa alik, yt non th sanras th nt and
th lot of thm suggstsa srt law, a sard puzz|e. ould l only, swt
lady, nvy to you in a word th happy solution.'' Th pot's ouption is
610 Goldshmidt

frquntly thought to prss onisly in vrs that whih is diffiult tl ar- Baus of his wor
tiulat. So, too, th iologist in Goth smsto sk to xprss th elusiv pointd as on of th f
forms that undrwrit th priens of natur. fr Biology in Brlin in
frns twn gog
BIBLIoGRAPY
ras only rpsntd
Kant' I. 2000. ritiqu of th Poturof ldgmt.P. Guyr,d. amridg: lutionary transformati
ambridgUnivrsityPrss. mutations. His idas .
Rihards,R. J. 2002. h Rlmtiocptilof Lif. higo:Univrsityof
phnoopis and hom
hiago Prss.
Smon,D., nd . Zajon,ds.1998.Goth'sWof Sic:A hrlmolog phnotypi hangs.c
lf Ntur. Alany: StatUnivrsityof Nw York Prss. usd to dsrithsr
Stur,D.2002.In dfnsof xprin: Gth'snaturalinvstigations
and phors in th history o
sintifiultur.In [,. Sharp' ed.,mbridgompnion to Goth, 160-168. Mayr's Sstmtis
amridg: amridg Univrsity Prss. -1.K. part in ordr to rfut
ntly in th ontxt
tyi hangs ar oft
Goldshmidt,Rihard Bndit(1878_195s)
ntworks of gns, a r
Rihard Goldshmidt was on of th most ontrovrsial gntiistsof th shmidt's arlir idas
twntith ntury. Known as a skilld primntalistand a rativthorist' found rfug ftr his r
his idas about th natur of th gn and th rol of systmimutations (or
|BLloGRAPHY
larg rrrangemntsof hromosoms) in volution hav n onsidrd
hrtial, whil at th sam tim thy hlpd to rystalliz th so-alld modrn Ditrih , . R . 2 0 0 3 .R i
synthsis' whos partiipants wr' at th vry last, unitd in thir opposi- Nture Ruit,us Gt
G o l d s h m i d tR, . [ 1 9 4 0
tion to Goldshmidt. or rntly' StphnJay Gould (1982, in his intro-
S. J. Gould. Nw Hl
dution to th rprint of Goldshmtdt's Tb teril Bsis lf uolutio) and 1'958.Tbeorti
othrs (.g.,Ditrih 2003) hav rvaluatd Goldshmidt and argud that 1'96'1d u
his idas dsrv at last as muh onsidration as th prvailing dogma of Mayr, . 1942.Sstm
adaptationism. Univrsityrss.
Rihard Goldshmidt Was orn in 1878 in Frankfurt am Main into a pro-
minnt Grman Jwish family and grw up in a wll-to.do miliu. H gan
his studis of mdiin in Hidlrg' attnding lturs of oth Otto
Gould, StphnJa
Biitshli and Carl Ggnaur. owvr, his ral intrstwas zoology, and in Stphn Jay Gould w
1898 h movd t unih t study with Rihard Hrtwig. His first aadmi ultural stting that n
appointmnt Was as Hrtwig's assistant in Munih. As part of his tahing ionolasm, and his lI
dutis h organizd th laoratory in omparativ anatomy' whih in part Columia Univrsity h
xplains his vast knowldg of animal morphology' anatomy, nd ell iol- Norman D. Nwll. F
ogy. In his own rsarh h fousd on prolms of gntisand ytology, ollaorator, Nils ll
spially on th prolm of s dtrmination. Glldshmidt workd with on whih h would sr
th Gypsy mot Lmtri, whose gographial variants, whn matd to In his arly work, (
ah othr, produd an assortmntof so-alld intrmdiatintrss.To growth of individuals r
ount for ths intrmdiatforms, Goldshmidt prposd a dynami and Using quantitativ m
quantitativ modl of gntisthat was asd on th undrlying physiologi- portion of individual <
al ation of gne produts, hen th nam physiologial gntis.Th asi ats and timing f tht
ida of physiologial gntis Was that diffrnt variants of gn produts powrful tool for ana
should hav diffrnt physiologial proprtis' Suh as ration tims, whih xtint organisms.Mo
an aount for th diffrnt osrvalphnltvos. provid a potntial ml

that whih is diffiult to ar.
l s kt o x p r s st h l u s i v
Juyr' d. amrilg:
/e. hiago: Univrsity of
of Si: Phltltolog
Prss'
'tural invstigatilnsr-rd
,)mpdia|1 to G lth,',\:f.
-1es8)
rtrovrsial gntiists of th
rtlisa t n d r t i v r h o r i s t ,
ll of systmimuttions (or
ution hv bn onsidrd
ystallizth so-lldmodrn
l a s t ,u n i t di n t h i o p p o s i -
r y G o u l d ( 1 9 8 2 .i n h i s i n t r o -
:ril Bsis of uoltttion) and
}ldshmidtand argud that
1 as th prvailing dogma of
ankfurt am Main into zrpro-
wll-to.d miliu. H gan
ding lttrrs of lroth otto
I intrstwas zoology' and in
d Hrtwig. His first aadmi
nih. As part of his tahing
'tiv anatomy, whih in part
logy, anatomy' and ll iol-
ms of gntis and tolg,
rr. Goldshmidt workd with
l variants, whn nratd to
d intrmdiatintrss.To
nidt proposd a dynami and
n th undrlying physiologi-
gsiologialgntis.Th asi
nt variants of gn poduts
suh as ration tims, whih
:yps.
Gould 611
Baus of his work in gntis and physiology, Goldshmidt was ap-
pointd as on of th founding dirtors of th nw Kaiser \Wilhlm Institute
for Biology in Brlin io \91' Soon his intrstslso inludd th gnti dif'
trnstwn gographi as of Lmtri. Ths diffrns twn
ras only rPrstd mirovolutionary vllts. To aount fo lrgr vo-
lutionary transformations, Goldshmidt postulated th istn of maro-
mutations. His ids on this sujt wr also motivatd y his study of
phnoopisand homoti mutations, oth prsntinglarg oordinatd
phnotypi hangs. Goldshmidt's notion of ..hopful monstrs''' whih h
usd to dsri ths rrovl phnotyps' is on of th most unfortunat mta-
phors in th histoy of biology and was soon ridiuld y his ritis. rnst
1.r's Sstemtics d the rigi of Speis (|942) was writtn in major
prt in ordr to rfut Goldshmidt. Howvr, as w hav land mor r.
ntly in th ontt of volutionary dvlopmntal iology, larg phno-
rypi hangs ar oftn th rsult of volutionry hangs in th rgulatory
ntworks of gns, onpt that is, at last in spirit, vry los to Gold-
shmidt'srlir idas. Gldshmidt did in 1958 in Brkly, wh h had
found rfug aftr his migration from Nazi Grmany.
B|LIoGRAPHY
Ditrih,M. R. 200]. RihardGlldshmid: monstrsand othr..hrsis.''
Hofr"rl
Ntur RuisGtltis4: 68_74.
Glldshmidt, R. [1940]1982.The terilBsisof uoltio.Introdutio y
S. J. Gould. Nw Hvn,T: Yal UnivrsityPrss.
1958. ThortilGtis.Sttl:Univrsiyof Washingtonrss.
.Washington
196.I d ttt of th Iuor Trir.Seatt|:
Univrsityof rss.
ayr, . 1942.Sstmtisd th origin of Spis.Nw York: olumia
Univrsity Prss. -.D.L.
Gould, Stphnla $94|_2002)
StphnJay Gould was orn nd raisd in Nw York ity, in a fanrily and
ultural stting that nurturd his intrst itr natural history' his instiriv
ionolasm,and his lft-laningpolitis. H attnddAntioh o1lg,nd at
olumia Univrsity h arnd his PhD undr th dirtion of palontologist
Norman D. Nwll. H also am assoiatdwith his liflong find and
ollaorator' Nils ldrdg. Harvard alld, and Gould joind th fault
on whih h would srv throughout his arr.
In his arly work, Gould fousd on angs in shap that ou in th
growth of individu:rlsnd in th volution of animal spisand highr taxa.
Using quantitativ mthods f allomtry, showd how hangs in pro-
portion of individual haratrs' in rlation to ody siz, rflt shifts in th
ratsand timing of thir dvlopmnt (sGould |966). This provd to a
powrful tool fr analysis of adaptiv funtion nd pattrns of volution in
tint organisms.Morovr, ths shifts in timing' known as htrohrony,
provid a potntial mhanism for rapid volutionry hang, triggrd y
612 Gould

modst hangs in rgulatory gns. Io tog d Phlog fl977\,

Gould assssdth history of th study of htrohrony and showd how it
was rltd to an mging fild that would sotln om known as voiu-
tionary dvlopmntaliology, now niknamd vo-dvo.
Studis of triloits and land snails providd ldrdg and Gould with vi-
dn showing that spis prsist for long priods of tim with littl if any di-
rtional hang. owvr, volution an our rapidly in loally isolatd
populations, giving ris t nw spisthat may sm from th fossil relrd
to hav appard instantanously.ldrdg and Gould (1972) advoatd a
modl of ..puntuatdquilirium'' as an ltrnativto grdualism, that was
impliit in most onvntionl ounts of volution y natural sltion (s
also Gould and ldrdge 1977,199I. This hypothsispromptd a livly on-
trovrsy that was nvr fully rsolvd;oth pattrnsof hang an shown
to our. any palrrtologiststndd to favor puntutd quilirium,
whras most gntiistshld to gradualism, oth groups swayd y th n-
tur of thir data and th timsalsovr whih it is osrvd.(Stinsand
Ayala 1981 provids gntiists,rsp()nsto plrntuatdquilirium. A his-
torial ovrviw of th ontrovrsyan found in Spkoski nd Rus 200s.)
Gould alwys mphasizd th rol of historial irumstanin volution.
In ..h Spandrlsof San Marl and th PnglossinPardigm:A ritiqu of
th Adaptationist Programm'' (1979), oauthord with RihrrdLwntin,
Gould hallngd th notion that vry asptof frm and havior of living
organisms an plaind dirtly in trms of ntural sltion. Naturl
arrirs or onstraints of anstral dsign and pathways of dvlopmnt'
haratrs with no immdit adaptiv funtion that mrg aS y-produts
of sltion for othr faturs,and alvall han must takn into a-
ount. (Slzr 799 analzs this papr frlm a litrary prsptiv.)Rlat-
dly, Gould was a liflong nthusiastfor th idasof th Sottishmorphologist
D'Ary N7ntworthThompson, who always argud that rulr animal form A partiularhro of St
.Wntworth
Thomsor
ould xplaind as th rsult of th laws of physis and hmistry without
d Form, shtlwingho
rfrnto natural sltion(sGould \971\'
illyfish(B) h:rvth s
Gould was a vigorous opponnt of th onpt f volutionary ..prrgrss.'' paraffir.r(A), wnt fr t
To him, it implid a prdtrminddirtion that has no sis in prinipl l nssarilyvnth pr
on th vidnof lif's vllutionaryhistory. hang is unprdital.any ..ordrftlr f
hristnd
fators influn th fts of individual spis, ld to divrsifiation of 1 8 5 ) .T h r i s s o m n
major groups of organisms' or us mss tintions tl-rathav priodially th laim is tht phn
lard th stag, making way for th volution of nw asts Of haratrs adaptivnor nonadp
ar adaptivut tht s
from a fw survivors. Gould (198s) is arong th svral paprs that takl
progrss. Full Hous: h Spred of lle from Pltl tl Dri (7996)
is full-lngth disussiorrdirtd at th gnral radr as muh as t th pro-
fssional volutiontst. volutionary hangl
Building on th Onpt of spis sltion laoratd y StvrrStanly rt ffts of naturl
(1975,1979)' Gould argud that pattrnsof volution in lads (groupsof r- pos an pandd v
latd spisthat shar a ommon anstor)aris from a pross of sltion ritria opratst su
diffrnt in kind from that whih urs in volutilnat th lvl of popula. tion is a fous of urr
tions and spition. If this marovoluti<lninvolvs mor than th sum of tion opratsamong (
 Could 6l.l

ttog|1 ild Phlog fl977),

rtrt>hrlty and shlwd how it
j soon tlmkn<rwnas volu-
md vo-dvo.
ld ldrdgand Gor"rldwith vi-
riodsof tim with littl if arry di-
our rpidly in loally isoltd
may sm from th fossil rord
: and G<luld(1972) adr,oted
trnativto gradulism, thar was
volution y naturl sltin (s
hypothsispromptd a livly on-
pattnslf hrrgan shown
l favtrr puntuatd quilirium,
r, oth !]roupsswyd y th n-
hih it is tlsrvd.(Stirrsnd
o puntuatdquilirium. A his-
lund in Spkoski arrd Rus 2008.)
:oriaIirumstl1nir volr"ltion.
lglossinParadiglrl:A ritiqu of
'uthordwith Rihard l-wontin,
:t of ftlrtrand vior of living
ms of nturl sltion. Nirtul
and pathways of dvlopmnt,
tion thirt mrg as y-produts
r l l h n l l l u s t h k n i n t < -l -
lm litrry prsptiv.)Rlat- AB
idasof th Sottislrmorphol<lgist
A partiulirrhro of StphnJav Gould ws th SottishmorphlogistD'ry
s argud that muh irnimal fom
WntwortlrTlrorn;lsor-r.Illr-rsrti<ls
lik tlrs,from hompsln oiz Grlth
of physisand hmistry withlut
d Fori, showing how (apparntlypurly s funtion of physialfors)
7r). jllyfish(B) hv th samshapas fallirrgdrops of ink in wat,or oil tn
nptof volutionarv ..progrss.'' paraffin(A), wnt far t onvinGould that naturl sltionis not th sol or
n that l-rtrs
nl ilsis in prirripl or nssirrilyvn th prim r.rslflrirrvolutilt-r.
Suh phrromnalrirvn
y. hng is unprdital.any hristnd..ordflr fr'' y th thortialiologistStuartKauffman (|995,
pis,lad to dir.rsifiatit>nof 185).Thr is slmmiguityin Thompson and in latr writrs as to whtr
tintiIrstt h;rvpriodially thlim is that phtlmnlik rh shapof th jlIvfishr nithrvy
daptivntlr non:rl:rptiv(arrdhnsltilnis trtltinvolv1)
or whthrthy
lution of nW asts of harzrtrs
areadaptivut tht sltinWasnot nddtl flrmthm. SGould 1971.
)ng th svralpaprs tht takl
,
fro Pltl to D,lr',i (1996)
lral 1as mtrh s at th pro-
volutionaryhang within irh spis,it is thry dlupldfrom th di-
ion la<latd y Stvn Stirnly rtfftsof naturl sltiot-t,
ating on ppulations.This ld Gould to pro-
irr lirds (grorips of r_
| vlltrrior-r pos n pandd voltrtionrythorv, in whih sltion sd on distint
) arisfrom a pross of sltion ritriaOpratsat sussivlvls in th taonomi hirarhy. Spissl-
. volutitlnat th lvl of popula- tion is a fous of urrnt rsrhand primntltsts.Th ida that sl-
n invtllvsmor thlln th sun of tion oprtsm()nglds has ),tto translt.linto tstaIhypothss.
614 cold

Baus h hallngdprvailing viws, som rightly saw Gould as a riti Spkoski, D., and . I
of neo-Darwinian orthodoxy. Nonthlss, h was an ardnt admirr of Univrsity of hi
harls D:rrwin, singhis own work as an pansion of Darwin's ntrpris Stanly, s. . 1975. A
Ntil Adem,
and no l-nnsa rfutation of it. Gould's arlir work is most assilto
|979. rl
thos sking arr intodution to lris idas;his finl mgnum opus,T Stru- Stbins, G. L.' and F'
tr oi uolutior Thor1' Q002), is a rilr ut unwildy tom, mor rad- Since 21'3:967_
r I yq u a r i dt h a n r d i n i t s n t i r t y .
In addition to his original rsearh, Gould pulishd numrous ssays' most
notly in th magazine Nturl Histor. Ths wr olltd in sris of Grant, B. Ros
st-sllingooks, starting with uer sice Dr,in (1977a). Othr works ad- and Grant, Pt
drssd msllrsof human brirr siz and intllign(Gould 1981)' th izarr
Roslary and Ptr
tint irnimals of Camian sas (Gould 1'989|,and th rlation rwn si-
win's finhs in th (
n and ligion (Gould 1999|. H tstifid as an xprt witnss i th Arkan-
..rtion formd on of th
sas ourt as whr sin'' was shown to a rligious dotrin.
dsriing th fdin
Gould nvr hsitatdto join an intlltual fight. As a result, Gould amth
populations; doum(
prmir puli fa of volution in th Unitd Stats during th last qurtr of
trns of mt hoi
th rwntith ntrrry.
strutillg th phylog
th signal as study
BIBLIOGRAPHY Th Grants riv
.'J7. with honors from I
Allmn' D., P. . Klly, nd R. . Rlss,ds.208' StephJ Gluld:Rfl-
tilsltt His Vie of l-if. Nrv York: Oxford UnivrsityPrss. ridg. Thy mt an
ldrdg,N., nd S. J. Gould. 1972.Puntuatdquiliri:r: to phylti
An altrnativ lumia. P.R.G' tool
gradulisrrr. In T. J. . Shopf,d.'odIsi Plobillrlgy, 82-115. olumia; B'R.G. t
Srll-alliso: Frmirn'ooprnd o.
dotoral yar at Yal(
Gould, S. J' 1966.Allomtryand sizin ontognyirndphylogn'Bkllogil
Ruitl,sof the mbridgPhiloslphilSoiet41'z587-640. Univrsity of Mihig
1971. D'Ary Thompson and th sinof forrn.Nelu Litrr istor 2: positions at thsun
229-258. island id populati
1977,.l.uer si Druitt.Nw York: Nlrton. gos and morphlog
1977.togd Phllg1,. amridg'A: arvard Univsitrss.
diffrnt islands.B.R
1981.Tbistsr of n. Nw York: Nrrrton.
1988.on rplingth idaof progrsswitlr an oprationalnotionof among individuals i
dirtionality. In . H. Nitki, ed.,uolutilr Progress,319-338' hiago: aly pprs on sl
Univrsityof higoPrss. patri spiation. Th
1989. WderfulLif: h BurgssShld th Ntur of Historl' thm to th Galipag
Nw York: Norton. hallngsof logisti
|996, FilI Hous: Th Spredlf llfrtlm PItoto Drtui.
Th Grants hav r
Nw York: r.rgn.
|999. Rtlcksof gs:Scind Religio i tb Fullessof Lif. luding the Darwin,
Nw Yok:Ballanti. and tlr Darwin-.W
2002' Th Strturelf uoltiorThor.amridg,MA: Harvard Grants wr ltd
UnivrsityI)rss. thy wr oth lt
Gould, S. J., nd N. ldrdg.1977, Puntuatd quiliria:Th trpond rrrodof
2007, P.R.G. in 200
voltionronsidrd. Pllbilllg
3: 115_151.
1993.Pr-rntuatd quilibrir"rmotnsof ag.Ntur 66 Z2.3-Z27. S i n s( B . R . G . i n 2
Gould, S. J., and R. . L,lrrti.7979. h spandls of San ro and th
BILIoGRAPl
Panglossin pradigm: itiqtrof th adaptationist programme,Proedigs
of th Rl Soitof Llndrl'SrisB 205:581-598. Grant,B. R., and P. R.
Slzr,.J.,d. 1993. Udrstdig SitProse.alison:Univrsiryof wisonsin Popultion:h L
Prss. higoPrss.

som rightly saw Gould as a riti
ss, h was an ardnt admirr of
r pansion of Darwin's ntrpris
s arlirwork is most assilto
his final magnum opls, Tb Struc-
ih ut unwildy tom, mor rad-
d pulishdnumrous ssays' most
Ths wr olltd in a sris of
Dri (\977a|. Othr works ad-
rtllign(Gould 1981), th bizarr
p$9),nd th rlation twn si-
l as an xprt witnss in th Arkan-
ts shown to a rligious dotrin.
fight.As a rsult, Gould am th
dStatsduring th last quartr of
ds.2008. StphJ could: R|l-
.ord Univrsity Prss.
ld quiliria: An altrnativ to phylti
ls i PIobiolog, 82-II5.
ny and phylogny. Biologil
ioiet 41: 587_640.
r of form. Netu Literr Histor 2:
Norton.
ridg,A: Harvard Univrsity Prss.
1lrk:Norton.
;s with an oprational notion of
utior Progrss, 3 1 9_33 8. hiago:
le d the Ntr of Histor.
ll[rom Plto ttl Drin.
gio i t Fullnss of Lif.
hor. amridg, A: rvard
:d quilibri:Th tmpo and mod of
l5-151.
i age.Ntur 366: 223_227.
ndrlsof San Mar and th
dptationistprogramm. Prodings
205:581-598.
.Wisonsin
-lse.Madison: Univrsiry of
Grt nd Grt 615
Spkoski, D.' and . Rus, ds. 2008. Th Pleobiollgil Ruolutio. tago
Univrsity of Chiago Prss.
Stanly,S. . 1975. A thory of volutin aov th spis |v|.Prodigs of th
Ntiol Acdm of Scis 72: 646-65.
1979. rouolutio, Pttr d Pross. San Frniso: W. . Frman.
Stbbins,G. L.' and F. J. Ayala. 1981. Is nw volutionary synthsisnssary?
Sic2I3: 967_971'. -R.D...
Grant,B. Rosmary(b. t936)'
and Grant, PtrR. (b. 196)
Rsmary and Ptr Grant ar rnownd for thir long-trm studis of Dar-
win's finhs in th Gal6pagos Islands. Thy and thir ollaorators hav pr-
formd on of th most omprhnsiv studis of any group of spis,
dsriingth fding ology, rding iology, and numrial dynamis of
populations;doumnting natural sltionon morphologial traits and pat-
trns of mat hoi; disovring hyridization twn spis; and ron-
strutingth phylognti rlationships among thm. Thir work stands as
th signal as study of volutionary iology.
Th Grants rivd undrgraduat dgrs in 7960, Rosmary (B.R.G.)
with honors from dinurgh and Pt (P.R.G.) with honors from am-
bridg. Thy mt and marrid whil ttnding th Univrsity of British o-
lumbia. P.R.G. took his dotorat jn 1964 from th Univrsity of British
olumia; B.R.G. took hrs in 1985 from Uppsala Univrsity. Aftr a post-
dotoral yar at Yal, P.R.G. hld faulty positions at MGill Univrsity, th
Univrsity of tr4ihigan' and Printon Univrsity whil B.R.G. hld rsarh
positions t ths univsitis.P.R.G.'s arly work fousd on th ology of
island ird populations, inluding pattrns of distriution aross arhipla-
gos and morphlogial variation mong spisand among populations on
diffrnt islands. B.\G.'s arly work on Darwin's finhs xamind variation
amng individuals in bill morphology and song haratristis'lading to hr
arly paprs on sltion on ill morphology and th possiilitis for sym-
patri spiation. Th Grants' ommitmnt to studying Darwin's finhs took
thm to th Gal6pagos for six months of vry yar (sin 197), dspit th
hallngsof logistis' limitd dit, and th spartan lifstyl of living in tnts.
Th Grants hav rivd host of honors, individually and togthr, in-
luding th Darwin dal of th Royal Soity of London, th Balzan Priz'
and th Darwin-!alla Award of th Linnan Soity of London. Both
Grants wr ltd to th Amrian Aadmy of Arts and Sins in 1997
thy wr oth ltdas Fllows of th Royal Soity of London (B.R.G. in
2007, P.R.G. in 2003) and forign mmrs of th U.S. National Aadmy of
S i n s( B . R . G . i n 2 0 0 8 , . R . G . i n 2 0 0 7 \ .
|LIoGRAPY
Grant,B. R., and P. R. Grant. \989. uoltiorDmisof Ntrl
Popultio:Th Lrge ttlsFinh of the Gldpgos.hiago:Univrsityof
hiagoPrss.
616 Crnt

Grant, P. R. 1986. olog d uolutkl lf Drtui's Fis. Prtnton, NJ: Tbulr Vie of the P
Printon Univrsity Prss.
praisd Darwin's on t|
Grant' P. R., and B. R. Grnt. 27. I]o d Wh Spies ultipl: Tb Rdition
-].. thir dinurgh days t
of Drtui's Fihs, Printon, NJ: Printon Univrsity Prss.
trs srouting from sp
two mn diffrd. A s
Grant, Robrt dmond (793_1874)
and fw mournd his p
Robrt dmond Grant' an dinurgh-orn spong xprt, ir-rtrodudom-
|BLloGRAPHY
parativ anatomy to Britain in th 1820s and 180s. H inrporatd th
studis of Jan.BaptistLamak and tinn Goffroy Saint-Hilair' apt- Dsmond,A. 1984a.Ro
ing th ..transformism,''or volution, aompanying thir works. Grant also transmutationist. /O
indutd th young harls Darwin into Lamark's viws. Darwin, ord y 1984.Rort.
l,turson ..palo
mdiin at dinurgh Univrsity ln 1825-1827, studid Noth Sa invrt-
95_413.
rats with Grant, who had om an MD in 1814. As an old man writing |989' Tha Pliti
ls Autobiogrph, Darwin ralld Grant's admiration for Lamark. Rdil Lodl.ht
Grant hampiond Hnri d Blinvill's ontinuous hin of lif (rathr S l o e n , . R . l 9 8 5 .D r w i
than Gorgs uvier's disrt embrnchemets) and Goffroy's lif that th transformism. In D. I
N J : P r i n U
nrliv
sam organs orrd throughut th hain-hn Grant's laim in 1825 to
hav found th homology of th vrtrat's panras in mollusks. Studying
spongs in 1825-1828, Grant oind th group's nam, Porifra, and h
mootd a simpl, dfnslss,frshwatr form as th ..parnt'' of th armord Gray, Asa (1810-1
marin spis. Darwin xamind th larva of th llal ryozoan Flstr un- Asa Gray was on of tI
dr Grant. Grant livd ths lrva wr rlatd to th ,.ova'' of alga and ia. H ame on of
pointd ak to the spontanouslygnratd..monads'' lying at th juntion mnts drivd from th
of th plant and animal kingdoms. At dinurgh Darwin was alrady roah- of volution. A dply
ing issus surrounding th birth of individuals and spis' vn if h rjtd logial impliations of l
Grant's monadism upon bginninghis own vlutionary spulations. ural sltionould
As th first profssor of omparativ anatomy at London Univrsity in y a wis and nvol
1827, Grant alrtd th mtropolitan lite to th Lamarkian thrat. H Gray was orn in Sr
joind th Gologial Soity ounil in 1832, just as harls Lyll was pu- i n m d i i n n d p r a t i
lishing his antievolutionry volum Priniples of Geolog. Grant's fossil l- tahing so that h w(
tures promotd th ..dirtgnration'' of sussivspis,on giving irth Fishr Profssor of Nat
to anothr. And his studntsw amind on th onditins rspnsilfor stood that h would o
..originating
and ffaing th tmporary organi film on our plant'' (Zoo|- onsult with othr bot
ogy amination Paprs, 1857_58, p. 6, in ..Grant on Zoologial Sujts,'' Amrian plants. H w
ollg olltion DG 76, Univrsity ollg London). ( 1 8 3 8 _ 1 8 4 3 )a n d v
Grant took th Fullrian hair of physillogy at th Royal Institution in ot of th Northert
187, ut that markd his apog.H had lost his ounil sat at th Zoo1og- January 30, 1888.
ial Soity in 1835, whn th zoo's aristoratipatrons gav his onsrvativ Gray,s work on th
rival, Rihard own, pfrntial tratmnt. At th Gologial Soity, Own Amrian plants mad
ountrd th fossil impliations of Grant's transmutationism. Grant lost r- voatd y Louis Agas
sours and his finans ttrd.Th 1840s found him living in a slum, his liation of th ri
shay swallow-tail oat as unfashionl irs his lturs. A rising youllg lttr to Gray, datd S
Thomas Hnry Huly in 1852 mrly laughd t Gfant's ..ntriity.'' Darwin and A. R..sa
At th nd, Grant nvisgdplantary ooling as th motor to powr th vo. 264-267|. Gray sus
lution of warm-loodd mammals and irds from dinosaurs and otrosaurs.In taks from Agassiz at t

,'ri'sFihs. Printon' NJ:
Vh Spisltltipl: h Rdition
-J.T.
ln UnivrsityPrss.
4)
spong prt' introdud om.
nd 180s. H inorporatd th
r Goffroy Saint-Hilair' apt-
npanyingthir woks. Grant also
mark'sviws. Darwin, ord y
| 8 2 7 , s t u d i dN o r h S a i n v r r -
) in l8l4. As an old man witing
; dmirationfor Lamark.
ontinuous hain of lif (rathr
zts|and Goffry's lif that th
-hn Grant's lim in 1825 to
's panrasin mollusks. Studying
group's nam, Porifra' and h
m as th
..parnt''of th armord
l of th loal ryozoar' Flustr un-
..ova'' of alg and
rlatdto th
. . m t l n a d s .I.y i n ga t t h j u n t i o n
d mnts drivd from th gographi distriution of plants to dfnd th thory
urgh Darwin was alrady roah-
als and spis,vn if h rjtd
volutionaryspulations.
n t o m y r L o n d o n U n i v r s i y i n
to th Lamarkian thrat.
32, just as harls Lyll was pub-
ls of Geology. Grant's fossil l-
ussivspis'on giving irth
l on th onditions rsponsil for
:ganifilm on our plant'' (Zoo|-
..Grant on Zoologial Sujts,''
lgLondon).
o l o g y t t h R o y l I n s t i t u t i o ni n
ost his ounil sat at th Zoolog-
:ati patrns gav his onsrvativ
. At th Gologial Soity, Own
; transmutationism.Grant lost r-
)s found him living in a slum, his
. rising young
a s h i s | t u r sA
..ntriity.''
;hd at Grant's
ling as th motor to powf th vo-
l from dinosaurs and ptrosaurs. In
Gr 617
Tblr Vieu., lf the Primr Diuisios of tb iml Kigdom (1861) h
praisd Darwin's the origin of Speis whil rminding his old pupil of
thir dinurgh days togthr. But th bulr Vi,s ountlss volutionary
trssprouting from spontanously gnratdmonads showd how dply th
fwo mn diffrd. A sminal figur in th radial 1830s, Grant outlivd his ag,
and fw mournd his passingin |874'
lBLIoGRAPHY
Dsmond,A. 1984a.Rlrt. Grant:Th soialprdimnt of a pr-Darwinian
transmutationist.lournl of th Histor of Biolog \7: \89_223'
1984.Rort. Grnt'sltrviwson organidvlopmnt: Th Swiny
Lturson ..palaozlogy,''1853_1857.rhiusof Nturlistor 11
95-4\.
1989. Th Politis of uolutio:orpholog,ediie' d Rform i
Rdil Lodo, hiago:Univrsityof hiago Prss.
Sloan,P. R. 1985.Darwin,sinvrtbrat rogram'1826-1.86: Pronditionsfor
trnsfomism.In D. Khn, d.,Th Drii Hritg,71-120,Prinrl,
NT: Printon Univrsitv Prss. -4.D.
Gray,Asa (1810-1888)
Asa Gray Was on of th lding otanists of mid-nintnth-ntury Amr-
i . H m o n o f h r l s D a r w i n ' s m o s r a t i v s u p p o r r s u
. sing gu-
of volution. A dply rligious man, h also wrot xtnsivly on th tho.
logial impliations of Darwinism' attmpting to show how volution y nat-
ural sltionould ronild with th lif that th world was dsignd
y a wis and bnvolntGod.
Gray was orn in Sauquoit, Nw York, on Nvmr 18, 1810. H traind
in mdiin and pratid for a fw yars, ut aandond this for part-tim
tahing so that h would hav mor tim for otany. H was appointd
Fishr Profssor of Natural History t Harvard in 1842, whr it was undr-
stood that h would onntratlusivly on otany. H visitd urop to
onslt with othr otnists, ut his own work fousd mainly on North
Amrian plants. workd oo a Flor lf Nortb Ameri with John Torry
(1838_1843) and vntually produd a highly sussful ul of the
Botl of tbe Northr Uitd Stts (7848). H rtird tn 187 and died on
J a n u a r y3 0 , 1 8 8 8 .
Gray's wlrk on th lassifiation and gographi distriution of North
Amrian plants mad him suspiious of th ..rationist''position bing ad-
voatd by Louis Agssiz. harls Darwin onfidd in Gray for th pu-
liation of o tbe rigin of Speis in 1859, and an astrat of Darwin's
lttr to Gry' datd Sptemr5, 1857, formd part of th joint papr y
Darwin and A. R. rJallain 1858 (rprintd in Darwin and !7al1a1958,
264_267). Gray susquntlydfndd th thory of volution against at-
taks from Agassiz at th Amrian Aadmy of Arts nd Sins.
618 Group Sltio

Svral of his pprs, supporting volutionisrnOl1th sis of otanial g- othr volutioni.ry fo

ography, wr olltd in his Drii (I876). Gray saw how th disprsal quotd aov. Flrtunat
of plants from an riginl loation' oupld with adaptation lf popultions ontriut mor to th t
to th loal nvironmnts fund in thir nw homs, ould aount for th in th sond part of D
obsrvd distriutiorrof spis.Th similarity twnth plants of astrn group'' an volv v t
North Amria and Jpn ould plaind y noting that in alir go- among groups in a lar
logial tims, thsplants lrd n widsprad aross Asi and North Amr- among individals rvitlr
ia ut had n lrivrrsollthward during th i tgs,r,l,ithonly th two I t i s i m p o s s i l t o d .
isolatd fragmnts of th arlir population now rmaining. dsriing its trrrtrln
As a staunh Prsytrian,Gray was anious to show that Drwin's thor1' twntith ntu}.did n
did not dstroy th Agumnt from Dsign. In arsrislf artils (printdin tions straightforwardly
Drii) h gn by arguing that ny pross y whih spis uld gns to osystms.T
b adaptd to thir nvironmnt Was ompatil with th lif that th and was widly ritiiz
volutionary pross had n stalishdy a wis and nvolntrator. his ook dpttio r
Latr on' howvr, h ndd that th wastfulnssand rulty of natural lassi. lWilliams affirm
sltiongnratddiffiultisfor this approah, nd h suggstdtht vria- i I f r a m w r k , . r g r
tion is ld along nfiiallins y th rato. Drrvin ojtdto this viw vlv only by a r()
bausit lft Gay supporting th plsitionof thistivolr'rtiorrism,
in whih mpiriallaim tht
th dition of dvlorrrntis dtrmindv surlrnatualrathr thn ttl- pard to lvitlrin-gror-
ral mans. multilvl sltionth
am th onsnsusfor
lLIoGRAPHY.
tionary thoris f s
Darwin, C. R.' nd A. R. !7alla.19.'8.uolutiob NturISlctio, altruism, gm tory'
ambridg:amridgUnivrsityPrss.
altrnativs to group s(
Dupr,A. H. 1959. s Gr. amridg,A: HarvrdUnivrsityPrss.
Gray, A. 1'848,utll of th Bot of th Northr UrtitdS/a/es.Bostn: Th 1960s onsllsu
J. Munro. sltion is thortil
1876. Driitt:sssd RuitusPrtiigttl Driism.New ramtr valus to prv
York: Applton. vining mpirial r.il
) vlls.Nw York:
Torry,J., and A. Gray. l88.l84. Flor lf N'lrthlrit'',t.
.Wily -P'I .B. s l t n t i r . dso r l t l i l
nd Lltn-l.
argumnts g;rnto b .
diffiult to rrguill fav
thortial modls of g
Group sltion .s7illiams
ssors. hims
Charls Darwin idntifid a fundamntl prolm with soial lif and its po- traits suh as sx rtio
tntial solution in th following famous passag from The Dest of Nss 1991). All r,olu
(Darwin 1871, |66):..It must not forgottn that although a high standard istn of multil gl
of morality givs but :l slight or no advantagto ah individual mn nd his idntifid, oopriltiv
hildrn ovr th mn of th sam tri, yt tht rr inrasin th numr within th grtlups irrr
of wll-ndowd mn nd advanmntin th standarlof rrroralitywill r_ nvisiond y multilr'
tainly giv an immns zrdvzrrrtg to on tri vr anothr.'' A mjor vnt in v
This problm and its potntial solutiorr ar th asis of a thortil ovry that t singl
framwork in volutionary iology alld multilvl sltirn(Sorand \Wil- gulis 1970; Nlir,vrlrdl
son 1998). Baus ntural sltion is asd on letiv fitnss,a trait that only y small mrrttio
nfitsth group as a whol dos not hng in frqunywithin th group. spissymiti s
A trait that nfits ()thrsat th pns of th slf iltually drassin fr- l v l o r g e n i s m si n r h i
quny within th group and will ultimatly om xtint in th asnof alld, hav lurlrt

)nismon th asis of otanial g-
l876). Gra saw how th disprsal
d with adaptation of populations
w homs,ould aount for th
Lritytwnth plants of astrn
nd y noting that in arlir go-
:adarossAsia and North Amr-
g th i ags, with only th two
r nw rmaining.
ious to show that Darwin's thoy
In a srisof rtils (rprintd in
y prossy whih spis ould
npatil with th lif tht th
y a wis nd nvolntCrator.
.astfulnss and rulty of natural
oah, and h suggstdthat varia-
tto.Darwin ojtdto this viw
of thistivolutionism, in whih
y suprnaturalrathr thn natu-
tio b Nturl Sltil,l.
\: Hrvard UnivsityPrss.
tthr Uitd S,a,es. Bosron:
; Prtiig to Driism. New
. North Amri. 2 o|s. Nw York:
-P.J.B.
,rolmwith soial lif and its po-
assagfrom Th Dscnt of
ttn that although a high stndard
tgto ah individual man and his
yt that an inrasin th numr
th standardof morlity will r-
:iovr anothr.''
amth asis of a thortial ovry that th singl organisms of tody wr th groups of past ags (Mar-
:rultilvlsltion (Sor and \WiI-
sd on rlativfitnss' a trait that
ngin frquny within th group.
lf th slf atually drassin fr-
v omtint in th asn of alld,hav ourrd ptdlyand possily inlud th origin of lif itslf as
Group Sltion 619
othr volutionary fos. This dsristh first prt of Darwin's passag
quotdaov. Fortunatly, groups of individuals who nfit ah othr will
ontriutmor to th total gn pool than more slfishgroups' as dsrid
..for th good of th
in th seond part of Darwin's passag.Traits that ar
group'' an volv y naturl sltion, ut only y a pross of sltion
among groups in a largr population and oftn in opposition to sltion
among individuls within singl groups.
It is impossil to dsri th ()npt of group sltion without also
dsriingits turulnt history. any iologists during th first half of th
twntithntury did not shar Darwin's insight; thy assumd that adapta-
tions straightforwardly volv ell lvls of th iologial hirarhy, fom
gnsto osystms.This position am known s navgroup sltion
and was widly ritiizd in th 1960s, spially y Gorg . Williams in
his ook dpttio d Nturl Seltil(I966), whih am a modrn
lssi.Williams affirmd th importan of multilvl sltions a thort-
ial framwork, agring with Drwin that group.lvl adaptations an
volvonly y a pross of twn-groupsltion.Williams thn mad th
mpiriallaim that twn-groupsltionis almost invarialy wak om-
pard to within-group sltion. It was this mpiril laim that turnd
..th thory of individual sltion,'' whih b-
multilvl sltion thory intr
am th onsnsusfor th rst of the twntith ntury. Susquntvolu-
tionary thoris of soial havio, suh as kin sltion, riproal
altruism, gam thory' nd slfish gn thory, wr dvlopd pliitly as
altrnativsto grOup sltion.
Th 1960s onsnsuswas asd upon thr argumnts:(1) twn-group
sltionis thortiallyimplausil, rquiring a dliat ombination of pa-
ramtrvalus to prvail against within-group sltion;(2) thr is no on-
vining mpirial vidnfor group sltion;and (3) th thoris proposd
as altrnativs do not invok multilvl sltion in thir own right. All thr
argumntsganto qustiond,vn as arly as th 1970s, although it was
diffiult to argu in favor of a topi that for many had om taoo. Modrn
thortial modls of group sltion ar fr mor plausil than thir prd-
ssors.Williams himslf aknowldgdth importan of group sltionfor
traits suh s sx ratio and disas viruln (.!illiams 1'992: Wll'Liams and
Nss 1991). All volutionarythorisof soial havior must assumth -
istnlf multipl groups to iologially ralisti.!hn ths groups ar
idntifid, olprativ nd altruisti traits ar sltivly disadvantagous
within th groups and rquir twn-grlupsltion to volv, as always
nvisiondy multilvl sltionthory (Sorand.Wilson 1998).
A major vnt in volutionary thory ourd in th 1970s with th dis-
gulis 1970; aynard Smith and Szathmary 1995)' volution prods not
only y small mutational hang,ut also y singl.spisgroups and multi.
spissymioti assoiations oming so intgratd that thy om highr-
lvl organisms in thir own right. Ths major transitilns of lif, as thy ar
620 Grrlt,tpSeletiolt

groups of ooprativ molular rations. Dspit multilvl sltion th- Rihson,P. J.' and R. B
ory's turulnt past' it is th aptd framwork for studying major transi- rsformed um
tions. Th alan btwnlvels of sltionis not stti ut an itslf volv. Sor,., and D. S. Wilso
UselfishBhuior.
A mjor transition ours whn sltion within groups is supprssd,n- .!illiams,
G. . 1966.Ad|
ling etwn-group sltion to om th primirry volutionay for. urrt uolutior
Tlr paradigm of rnajor transitions did not appar until th 1970s and was 1992.Nturl Sl
not gnralizd until th 1990s, ut i.rlradyit sms likly to plain th vo- UnivrsityPrss.
lution of soial inst olonis (.Wilsorrand Hlldoblr 2005) and humans as Williams,G. C., and R.
th first ultrasoial prirnt spis (Boehm 1999). Th traits assoiatd with QurtrlRuie'o
Wilson, . o., rrdB. Hol
human morality Ppar dsignd to suppress sltion within groups' similar Proedigsof th N.
to gnti mhanisms suh as hro-losomsand th ruls of miosis, nabling
sltion among grollps to om th primary volutionary for. ultural
prosssan aus groups to om vry diffrnt in thir phnotypi prop-
rtis,vn whn thv ar gntillysirnilar.Multilvl sltionhirs om
th aptd framwork for th study of human ultual volution, in xatly
th sam way as th study of major transitions (Rihsn and Boyd 2004).
Th rvival of grop sltion might sem traordinary until w rmm-
r th:rtWilliams (|966) aptdmr.rltilvl as a thortial fram-
slti<lrr
work, riting group sltion on th sis lf an mpirial lim that it is
lwas wak oIrp:rrdto within-group sltion.Whn Williams hangd
his mind aout th thortial and mpiial vidn' h himslf rvrtd
ak to multilvl sltion. For fml.iasd s ratios (\Williams 7992'
49), he statd, ..I think it dsiral . . . to raliz th:rt sltion in fmal-
isd ndlin popul:rtionsfavos nrirls,nd that it is only th sltion
among suh groups tht an favor th fmal ias''' For th volution of vir-
..Th
uln in disas ol.]nisms(Williams and Nss 199|, 8), h statd,
volutionary outom will dpnd rn rlativ strngths of within-host and
etwn-hostomprition in pathogn volution.''
Unforrunatly, th fild as a whol hzrsnt yt ahivd a nW onsnsus.
Books nd artils Writtn from multilvl prsptiv appr alongsid
othr ooks and artils that ontinu to trat group sltion as ir rjtd
onpt. Ttooks hav en partiularly slow to rflt th many dvlop-
mnts that hav takn pla sin th 1960s. Th trr-rlnt histry of this im-
ortant sujt is rviwd y Borrllo (2005) in an rtil appropriatly
titld ..Th Ris. FalI. and Rsutionlf Grourr Sltion.''

BIBLIOGRAPHY
Bohm, '. \999' Hirrh i tb Forest:glitriismd th uolutio of
ttl tt ltrui sm. arrrridg,MA : Hzrrr,i-r
rd Univsit,vPrss.
"t
Brllo,. .200.5.Th ris,fall' nd rsurrtiln of goupsltiorr.
deuor
29:4347.
Darwin, . 1871. Th l)sctof n, tdSltilli Rltio,o Sex. Nw York:
D. Appltonar-rd .
arguIis,L. \970. rigi of ukrotil/s.Nw Havn,T: Yal Univrsity
Prss.
MaynirrdSmith,.|.,nd . Szthmarv.l995. Th jorrsitiosrlf Lil?. Nw
l,tlrk: !7. . Frman.
 Group Sltio 62

)ns. Dspit multilvl sltion th- Rihrson,P. J.' and R. Boyd. 24.Not b Gsll:olt, Itur
rmwork for studying maior transi- TrsformdHum uolutio.Chiago:Univrsityof hiago Prss.
.!ilson.
Sobr,., and D. S. 1998. I.JtoOthrs:Th uolutio d Pshologof
rtion is not stati ut an itslf volv.
UslfishBhuior.amridg,A: Harvard UnivrsityPrss.
on within groups is supprssd, n- Williams,G. . 1966.dpttiod Ntrl Seletio: ritique of Som
n th primary volutionary for. rrtuolutiorThought.Printon,NJ: PrintonUnivrsityPrss.
J not appar until th 1970s and was 1992.Nturl Sltil:Domis, Leuelsnd hllgs.oxford: Oxford
ady it smslikly to xplain th vo- UnivrsityPrss.
\fillims. G. .. and R. N4.Nss. 1991,.Thdwn f Darwinian mdiin'
lnd Holldolr 2005) and humans as
'm 1999\. Th traits assoiatd with QurterlRuituof Biolog66:1_22.
Wilson,. o., nd B. lldolr.2005.usoiality:Origin and onsquns.
'prsssltionwithin groups' similar Prodigsof the Ntiol dem of Sics\2: 167_1371.
lmsand th ruls of miosis, naling -D.S.\{1
primary volutiony for. ultural
ry diffrnin thir phnotypi prop-
nilar. Multilvl sltion has om
f human ultural volution, in xatly
sitions(Rihrsonand Boyd 2004).
smtraordinary until w rmm-
ilvlsltionas a thortial fram-
asis of an mpirial laim that it is
tp sltion. \Whn \/illiams hangd
rpirial vidn, h himslf rvrtd
,l-iasdsx ratios (.Williams 1992,
. to raliz that sltion in fmal-
nals,and that it is only th sltion
[malias.''For th volution of vir.
..Th
and Nss 1991' 8), h statd,
rlativ strngths of within-host and
volution.''
las not yt ahivd a nW onsnsus.
rltilvl prsptivappar alongsid
to trt group sltion as a rjtd
arly slow to rflt th many dvlop-
960s.Th turulnt history of this im-
lo (2005) in an artil appropriatly
r of Group Sltion.''

d th uolutio of
Zglitriism
arvardUnivrsityPrss.
of groupsltion.
iurrtion deuor

d Seltioi Rltioto S.Nw York:

]ells.Nw Hvn,T: Yal Univrsity

5, Th jor Trsitioso/ Lile. Nw

 Brlin. iillr's suii
Jna, who amhis
Siily, h fll in with .
Flrmann Allmrs (1
giftd paintr (s fig
srh for th lif .
( 1 8 3 5 - 1 8 6 4 ) ,h i s f i r s t
arr. With a small tl
his rsarh on th lit
of a pinhad that sr
Hakl,rnst (184_|9t9)
ing his hailittion z
rnst Hinrih Philipp August Hakl was harls Darwin's formost ham. rih Bronn's (1800-1
.lforld .!Var
pion at th turn of th twntith ntury. Mor popl prior to I onvrt. is rsarh
larnd of volutionary thory through his voluminous puliations than nt two-volum foli
through any othr sour. F{ts Ntiirlih Scho1lfgsgesbicht(Natural his- own illustrations. Th
tory of ration' 1868) wnt thrugh 12 inrasinglyaugmntdGrman di- volums y way of inl
tions (1868-1920) and ws translatd into th major uropan languags. rh ddd fiof
rik Nordnskild, in th first dds of th twntith ntury' iudgd it ..th traordinarius profsso
hif sour of th world's knowldg of Darwinisr" (\96,515). Th rum. akl's rilliant l
ling dtritus of this synthti work an still found sattrd alng th tn months suddnl
shlvsof most usd-ook stors.Di Wltr|ithsel(Th world puzzls, 1899), ovry h wrot his
whih plad volutionary idas in a roadr philosophial and soial on- rd. Rathr, h wou
tt, sold ovr 40,000 opis in th first yar of its puliation and wll ovr Darwinian promis o
15 tims that numbr during th nt quartr ntury-and this only in th onvition in onsid
Grman ditions. (By ontrast, during th thr dads twn 1859 and tion of Darwinian id
1890, Darwin's the rigi of Speiessold only som 39,000 opis in six l{ts Gnerell or1lh
nglish ditions.) B |972, Die Weltrtithsel had n trnslatd' aording to isms, 1866) laid dowr
Hakl's own mtiulous taulations' intr 24 languags, inluding Armn- for th rst of his r
ian, hins, Hrw, Sanskrit, and spranto. Th young Mohandas Gandhi Darwin but mor ar(
had rqustd pmission to rndr it int Gujarati; h livd it th sintifi ook Fiir Dri (18(
antidot to th dadly rligious wars plaguing Indi. Hakl ahivd many sition that th mryo
othr popular sllsss'and h produd mor than 2 |argethnialmono- phologial stags as t
graphs on various aspts of systmati iology and volutionary history. His Gnrell orpbollgi
studis of radiolarians, mdusa' spongs' and siphonophors rmain stn- with nologisms that 1
dard refrns today. Ths works not only informd th puli' ut thy ology' and a host of
drw to Hakl's small univrsity in Jna th largst sher of urop's grat dud tr diagrams t
biologists of th nt gnrtion, among whom wr Rihrd Hrtwig morphologial and tr
(1850_197) and his rothr Osar Hrtwig (7849_1922)' Anton Dohrn by advaning a Goth
(1840_1909)' Hrmnn Fol (1845_1892), duard Strasurgr(1844_1912), for th naturalist: Go,
!. o. Kvalvsky (1842_1883), Nikolai Mikluho-ac|a fl846-1888)' xprssions of th sam
Arnold Lang (1855_1'914),Rihard Smon (18.59_1918)'lfilhlm Roux ly (1825_1895)init
(185-1924)' and Hans Drish (|867_7947). Hakl's influn tndd orphologie' shorn o
far into suding gnrations of iologists' many of whom ralld rading In ordr to sk a w
his popular works as young studnts. rd a sris of popul
Hakl rivd his mdial dgr fom !ijrzurg in 1858, aftr whih phology. Th srisw
h plannd to do his hailitation with Johanns iillr (1801_1858)at Sch6pfugsgeschibt
622

]harls Darwin's formost ham.
[o popl prior to l/orld !7r I
s voluminous puliations than
h ijpfugsgesh i hte (N atural his-
:asinglyaugmntdGrman di-
th maior uropan languags.
l twntithntury' judgd it ..th
w i n i s m ' (' 1 9 6 . . 5 1 5 )T. h r u m .
:ill found sattrd along th
iithsel(T world puzzls, 1899),
[r philosophial and soil on-
r of its pLrliationand wll ovr
tr ntury-arrd this only in th
hrddstwn 1859 and
l d o n I ys m 9 . 0 0 0 o p i s i n s i x
had ntransltd,aording to
24 languags,inluding Armn-
to. Th yor'lngN1ohandasGandhi
ujarati;lr livd it th sintifi
ng Indi.akl ahivd many
or than 20 larg thnial mono.
ogv and volutionarylristoy. His
a n d s i p h o n o p l t o r s m a i n s t a n .
'ly informd th puli' ut thy
h largstshar of urop's grat
; rvhom wr Rihard Hrtwig
ig (1849_1922), oton Dohrn
ldr'rardStrsurgr (1 844-79 12|,
iklulro-Malay (1 846_1 888),
l n ( 1 8 5 9 _ 1 9 1 8 ) ,$ 7 i l h l m R o u
l1). Hakl's influn etndd
' n]anyof whorrrralld rading
t \Wiirzurgin 1858, aftr whih
l h n n sM t i l l r ( 1 8 0 1 _ 1 8 5 8 ) a t
Hkl 62
Brlin' Miillr's sLtiidld hil to turn to rl Ggnaur (7826-190) at
Jna,who mehis advisr. During his rsarhwork in southrn Italy and
Siily,h fll in with a group of Gman artists' among whom was th pot
Hrmann Allmrs (182I_1902), who am a liflong frind. Hakl, a
giftd painr (s figu on pag 624), rhought of giving up iologial r-
sarh for th lif of a Bohmian; only his trothal to Anna Sth
(1835-1B64), his first ousin, kpt him for'rsdon stalishinga profssional
arr..Witha small trat y Millr as his inspiration, Hakl onntratd
his rsarhon th littl-known groLrpof rdiolaria, ratursout th siz
of a pinhad that srtxoskltonsof unusual gortris.\7hil omplt-
ing his hilittiorr k in Blin, lr rad Darwin's rigin in Gorg Hin-
rih Bronn's (1800_1862) Grman translation and immdiatly am a
onvrt. His rshfinally yildd, in 1862, Die Rdiolrie-a magnifi-
nt two-volum folio having traordinarily autiful plats asd on his
own illustrations.Th ook won th admiration of Darwin, who rivdth
volums y way lf introdution to this nw disipl. Hakl's rsarh hd
th addd nfitof allowing him to marry Anna and to gin his lif as -
traordinariusprofssor in th mdiai shol at Jna.
Hakl's brillint ginning turnd drk in 1864 whn lris wif of igh-
tn months suddnly did. I suffrd i1 nrvous ollaps, and during his
rovyh wrote his p:rrntstht h ould no longr apt thir rligious
rd.Rathr, h would put his faith in somthing mor rlial,namly, th
Darwinian promis of progrssiv trensformation. H thn dvlopd that
onvition in onsidraldtail in a larg two.volum, thrtial applia.
tion of Darwinian idas to all ars of iology, inluding humn volution.
His cerll orphologi dr rgism (General morphology of organ-
isms, 1866) laid down th fundamntl onptions that h would ultivt
for th rst of his arr' H r-nadntral an ida that h found intimated in
Darwin ut mor rfirlly workd out ,v Fritz ijllr (7827_1897) in his
ook Fr Dri (1864), namly, th prinipl of rapitulation-th propo-
sition that tlr mbryo of a givn spis would pass through th sam ror-
pholgial stags as th phylum had in its volutionary dsnt. Hakl's
Geerll orphologie formulatd svral nW prsptivs, outfitting thm
with nologisms that gav his tratisa formidal ast: phylum' ontogny'
ology,and a ost of othr trms that had a shortr lif span' H als intro-
dud tr diagrams to illustrat th dsnt of spis and to suggst thir
morphologial and tmporal distanfrom on anothr. Th book onludd
y advaning a Gothan monism s th appropriat mtaphysil position
for th naturalist: God irnd ntur' mind and ody '*,rto rgadd as
prssionsof th sam undrlying Urstif . Darwin and Thomas Hnry Hux.
ly (1825_1895) initiallv sought to hav n aridgd vsion of t|le Gerlle
orphologie, shorn of its polmial ars, trnslatdinto nglish.
In ordr to sk ir widr audinfor his thortiltratis'Hakl dliv-
rd sris of popular lturs in 1868 summarizing his Darwinian nror-
phlogy. Th sris was pulishd th sam yar ndr th tjt\eNtrlih
Sh\pfugsgshiht,nd it ahivd immdit notority. In an initial
624 Hechl

rviw, Ludwig Rtiti

kl witl.r faud
rapitulatiOn-or t}
rprsntdvry your
ially idntial. Rtitir
as y using th san
(1870),Hakl r-rs
arly stag n1sid i
tl arrs,v()unn
and th hi.rrgrffra
Dspit th ontro
ful impt on its r.
rprsntd nin sp
vlopmnt' with th
.1
t roots io t|l Urt

t h h i g h s tl v l , a r
Jws, and Aras. Al
mon to nintnth-
an attitud whih or
gud, following his f
grads of humzrn m.
and that tlr urop
ingly ompl mind-
of .
In 1867, aftr vis
akl travld to
prformd th kinds
that would grn f:
latr. H also gan'
would yild a thr
spongs' |872). |n l
maintaind Was n
mpirial proof tht
Hakl also rgtrd
fl4#': organism, having tlr
mtzoans in gastru
rnst Hakl was a rillint tistand always filld his llkswith his owl
thory.
drawings.Tis on, rvalingl,v laldth ..Apothosisof vlutilna11,
Thought'' (from th supplmntto Haekl's\Ydrbildr,190.l)'is an Baus of his var
intrstingrfltionon his owtr privat lif-h was griving th dath f ir ivd th ommiss
young wInan' tO Whom h ws r"rotmarrid ut with rvhom h had just hd a lasssof orgrrisms
Passionat(spiritualut also phvsial)ffir-as wll .rsth angingtims. h omposd sr'
Sigmund rud'sThree sssrl tlle Thor lf Sulitl,(gratlyinflundy siphonophors, and
akl'sthinkirrgn rpitul;rtin) ws pulishdin th smy this othr author in th
pitur appad.
th high rgard of tl
That rgrd was als
rivd during his l:

L
lld his ooks with his owrr
rthosisof volutiona11-
mdrbildr' 1905), is an
ws griving th dath of a
with whlm h had just had
l wll as th hnging tims.
Sulit (grtly inflund y siphonophors, and radiolaria-with mor pags produd than y any
hd in th sm yr this othr author in th sris of bllgr rports. Th ommission indiatd
Hkel 625
rviw, Ludwig Rittimyr (1825_1895), an mryologist at Basl, hargd
Hakl with fraud. H osrvd that in illustrting th prinipl of
rapitulation-or th iognti lw' as it am known-akel had
rprsntdvry young mrvos of a dog, hikn' and turtl as morpholog-
ially idntial. Riitimyr maintaind' howvr, tht Hakl had mad th
asy using th sam woodut thr tims. In th nt dition of th ook
(1870),Hakl usd only on illustration of a vrtratmryo at a vry
arly stag and said it might as wll b th mryo of a dog, hikn, or tur-
tl aus you annot tll th diffrn. Th damag, howvr, was don,
and th harg of fraud would haunt Hakl for th rst of his days.
Dspit th ontrovrs, |trtiirlihe Shopfugsgeshicbte mad a powf-
{ul impat on its rdrs, spially on th topi of human volution. H
rprsntdnin spis of humn ings along a tr of volutionary d-
vlopmnt,with th Papuans and ottntots at th lowst rnhs, losr
to roots i te Urmsh, or ap-man' and with th auasian branh at
th highst lvl, arrying at th top rahs th ditrranans' Grmans,
Jws, and Aras. Although Hakel shard many of th raial viws om-
mon to nintnth-ntuyuropans, h was diddly not anti-Smiti'
an attitud whih on of his disfftd studnts hld against him. H ar-
gud' fllowing his frind, th linguist August Shlihr (1821_1868), that
grads of human mntal aility prssd grads of languag omplxity
and that th uropan and Smiti languages hlpd rat a orrspond-
ingly ompl mind-a gnral thsis that Darwin adoptd in The Desnt
of .
| 1867, aftr visiting Darwin and othr British sintists in ngland,
akl travld to th nary Islnds with thr rsarh assoiats.H
prformd th kinds of primnts on dvloping siphonophor mryos
that would garnr fam for Wilhlm Rou and Hans Drish som 20 yars
latr. H also gan work on a systmati analysis of alrous spongs that
would yild a thr-voium study, Die lks,ijmm (Th alarous
spongs' 1872). In this work, Hakl attmptd to provid what Bronn
maintaind was nssaryto show th viaility of Darwin's thory, namly,
mpirial proof that spisdsntWas mor than a thortial possiility.
Hakl also argud, mploying th ignti law, that in anint tims an
organism, having th strutur of a primitiv spong (and th form takn y
mtazoansin gastrulation),plid th anint sas. This am his gastraa
thory.
Bausof his various invstigationsof marin invrtrats,Hakl r-
ivd th ommission in th lat ]870s to dsri systmatially svral
lasssof organisms drdgd up y HS hllegr. ovr a 10-yar priod,
h omposd svral larg volums on mdusa, alarous spongs'
th high rgard of th sintifi ommunity fr his work in marin iology.
That rgadwas lso prssdy th many honorary dgrsand wards h
r i v dd u r i n .h i s I r t i m .
626 Ifckel

Rsntmnt y th biologially and rligiusly ortodox ontinud to build l868.Dl Na/r

against Hakl throughout th 1870s, and it has not aatd in to this day. In | 8 7 2 .D i l k s
t899, Di \Yeltr
I874, the Swiss mryologist \Wilhlnr His (1831_1904) ptrlished Use
H i s , W . | 8 - 4 . U l t s r i i
Krperform ud ds phsiologiscbe Problem ihrr ntsthng (ur odily ntsthug.Lipzig:
form and th physiologial prolm of its origin), whih rpatdth arlir Kruss,. 1984.rstI
hargs of fraud against Hakl and institutd nw ons. Among othr laims, Miillr, F. 1,864.iir Dr
is assrtd that Hakl had rprsntd th humn mryo with zrn ag- N o r d n s k i o l d, . I l 9 2 0 _
gratdl long tail- ontrovrsy tlrirt am knwn as te Sch'zfrg' Pulishing.
Pnnisi,. 1997. k
|n 1877, Rudolf Vihow (7821_1902) rjtd l-risontim studnt's fforts
Rihards,R. J. 2005.Tht
to hav volutionary thory taught in th low shools in Grmny. Virhow Hakl'sr,olution
hargd that volutionary thought attd soialists and ommunists' a laim Fesf uolt
tht uly thought quit surrilous aus of its inflammatory hilratr in 28.h rg'
Bismrk's Grmany-although, in fat, many Marxists (.g.,August Bl, uolutior hllug
R o u ,W . l l t 8 8 'B i t r i g
1840_1913) did find Darwinism ongnial. At th tun of th nrury, rli-
pthll,'gisllAtl'
gious opponnts of Hakl,s Weltriithsl and of his nwly stalishd 113-153,246-29
Monist Lagu rnwd th laims of falshood. Ths many hargs had
thir foundation in kl,s aknowldgd slip in 1868, ut thraftrthy
gaind for mostly from intlltul ralitrannd rligious dogmarism.
Haldan,J.B. s. (
Mor rntly,Danil Gasman (197 |) and StphnJay Gould (1977) argud
that Hakl's iology supportd Nazi raism, although thy onvnintlyig- John Burdon Sandrs
nord akl's philo.Semitism, an attitud quit unusual for th priod. N o v m 5 ' l 8 9 2 ' i n
ihal Rihardson rxamind Hakl's illustrations of emryos, and h t h p h y s i i nJ o h n S
too suggstdHakl's malfasan.Rihardsn omprd akl's illus- d a n ( l a r r i t h i s n
trations with photographs of mryos, and asily showd th dvirtions'How- t o a t s J. . B ' S . H a l d r
vr, Hakl had adaptd illustrations from thn-ontmporarysours.H th modrn synthtitl
showd that whn you lind up dpitions ndrdy xprts' th similarity n d S w l lW r i g h t ( l l
of volutionarily rlatd typs at irrlir stags of mryognsis am t i o h m i s t r y .I n e d
manifst-a phnomrron aknwldgd y tody's mryologists. ration- Wr1tr.
ists and Intllignt Dsign thorists hav itd th oldr Gmn litratur H a l d a n W s d u a
and Rihardson's photogrphs to indit not only Hakl ut all of volu- lg, oxford, in ngla
tionary thory. m t i s s h o l a r s h i pr
honors ii-rmathmati
lLIoGRAPY sis and phil<lsoplry. H
Drwin' . l860. Ubr die ntstchugder rtn im hir-nd Pflz-Raib i n s i r r d v l o p d
durh trlihZbtug odr,rhltugdr urt,lllhommnete Rssnim a n d s i s t r .l n l 9 0 l , w l
Kmpf m's Ds.H. Bronn, trllns.Stuttgart:Slrwizrrt,slr tur whr th iolog
Vrlagshndlung. rdisovrd prinipl
Di Gregorio,.2005. From Hre to trit:rst kl d SitificFith.
ing guina pigs. aldz
Gottingn:Vandnhokand Ruprlrt.
Drislr,. 1891. ntwiklungsmhnislr Studin.Zitshriftfilr h d i t yi n 1 9 0 8 . B y l
issnshftlic Zoologi 53: 160_184. th inheritan of tw
Gasman,D. 197|, Th Sietifirigis of Ntiol Soilism'Nw York: Sin W a r I s a m r n ro |
H i s o yP u b l i t i o n s .
lourI of Gtics.
Gould, S. J. 1977.t<lg nd Phlog'amridg,A: HarvardUnivrsity
l n l 9 l 9 , H l d e n l
Prss.
kl,, |862. Die Rttdiolri (Rhi<lp<ld
Rdiri):ie |l4togrphi.
2 vo|s. l i l i g n t I yo n l i I r k a g
Brlin:GorgRimr. ridg Univrsit, wh
1866. Grllorphologidcr rgism'2 vols.Brlin:Gorg Rimr. m ,w i t h F i s h r.

rsly orthodo ontinud to uild
: has not aatdin to this day. In
(1831_1904)pulishd Usr
n ihrer tsthug (our odily
.igin),whih rpatdth arlir
d nw ons.Among othr laims,
l human mryo with an xag-
m known as te Schtuzfrg.
:tdhis ontim studnt's fforts
.rshools in Grmany. Virhow
,ialistsand ommunists' a laim
o f i t s i n f l a m m a t o r y h a r a t ri n
ny arists (.g.,August Bl,
At th turn of th ntury, rli-
and of his nwly stalishd
hood. Ths many hargs had
slip in 1868, ut thrftrthy
r r n n d l i g i o u sd o g m a t i s m .
tphnJay Gould (1977) argud
r, althoughthy onvnintly ig-
quit unusual for th priod.
illustrations of mryos, and h
rdson ompard Hakl's illus-
sily showd th dviations. How-
thn-ontmporary sours. L{
ndrdy prts,th similarity
ags of mryognsis bam
today's mryologists. ratin-
td th oldr Grman litratur
: only Hakl ut all of volu-
im Thir- ud Pflnz-Rih
lr uru,llkommtrtRrlss im
;art: Shwizrart'sh
.nst ckl d Sietifi Fith.
lien, Zitsbrift fiir
l Soilism. Nw York: Sin
n b i d g
, A: Harvrd Univrsity
diri):i olgrpbie. 2 vols.
smn'2 vols. Brlin: Gorg Rimr.
Hlde 627
1868.Di NtrlihShlpfgsgeshicbt. Brlin:Gorg Rimr.
1'872.Di Klkschiimme.3 vols. Brlin:Gorg Rimr.
Bonn: mil Strauss.
1899' Di Wltrijthsl.
.W.
His, 1874.lJsrKorprflrmd ds phskiogischProblm ihrer
tsthug. Lipzig:Vog1.
Krauss,. 1984,rst Hkl.Lizig:unr.
Mtillr,F. 1864.Fr Dri.Lipzig:nglmann.
Nordnskiold,. |1920-7924|1936.Th Histor of Bbbg' Nw York: Tudor
Pu1ishing.
Pnnisi,, 1997.Hkl,smryos:Fraud rdisord. Sci277:1'435.
Rihards,R. J. 2005.Th asthtiand morhologialfoundationsof rnst
Hkl'svolutionary roit.In . Kmprinkand P. Dassn,eds.,The
Fsof uolutklni urtp'1860-1914.Amstrdam:Ptrs.
208.Th Trgi Sesof Lif: rst Hkl d th Strugglour
uoltiorThought.hiago:Univrsityof hiago Prss.
Rou,!7. 1888.Bitrigzur ntwiklungsmhanik dsmryo,rhiu fr
pthologishtlmiund Phsilltlgiud fr kliishdii 94:
113*153,246-291. -R./.R.
Haldan,J. B. s. (L892_|964)
John Burdon Sandrson Haldan, known familiarly as J. B. S.' was orn on
Novmr 5, 1.892,in dinurgh, Sotland, to Louisa Kathln Haldan and
th physiian John Sott aldan. J. B. S.,s youngr sistr was Naomi Hal-
dan (latr ithison). Th aldan family is dsnddfrom Sottish ais-
torats.J. B. s. Haldan is known prinipally aS on of th thr arhittsof
th modrn synthti thory of volutin, along with R. A. Fishr (1890_1,962)
and Swall Wright (1889_1988). Haldan also mad original ontributions
to iohmistry.In addition, h was a popul sin,politial, and fition
Wrltr.
Haldan ws duatd at th Dragon Shool, ton Collg, and Nw ol-
lg'oxford, in ngland. ntrdNw ollg in \911' y wy of a math-
matis sholrship Won at ton. A yar latr, Haldan won first-lass
honors in mathmatis and susquntly,in 1.914,first-lasshonors in las-
sis and philosophy. ldan did not possssa sindgr'yt his intrst
in sin dvlopd at n arly ag, mainly du to th influn of his fathr
and sistr.In 1901, whn aldan Was ight, h and his fathr attndda l-
tur whr th iologist A. D. Darishir (1879_1915) disussdth nwly
disovrdpinipls of ndlism. Along with his sistr,who was rd-
ing guina pigs, Haldan did xprimnts looking for pattrns of Mndlian
hrdityin 1908. B 1912, Haldan livdh had vidnof linkag (i..,
th inhritan of two allls togthr),and in 1915, for ntring !orld
lVar I as a mmr of th Sottish Blak \/ath,his viw was publishd in th
Jourl of Gtics.
In 7919, Haldn m a fllow of Nw ollg, Oford. H workd
diligntly on linkag, pulishing svral paprs. In t922 h movd to am.
idg Univrsity, whr h would stay until 192. t amridg, Hldan
.s7right,
am, with Fishr and a ky figur in th synthsis of Darwinian
628 lde

natural sltion with th prinipls of ndlian hrdity.Indd,Haldan al Institut. In his tr

would play no smll rol in uslrring in thortial population gntis.In Haldan argud that
1924,Ha|dan pulishd th first of a sries of 10 mathmatial paprs - of a sustitution load
ploring th reoniliation of Darwinism with Mndlism; th last in th sris in nvironmnt) oul(
was pulished in 194, just tw<l yars aftr publiation of his ook Te Haldan's laim was
ses of uolutio (\92|. Haldan was aw21
ses ws th apston txt not onl of Haldan's serisof paprs on vo- 1958, and in 1961 r,
lution; it was th apston tt of th origins of thortial population gn- my of Sins.Aft
tis. Haldan's prolmati was alignd with his ontmporaris;h wantd Bhuanswar, orissa
to undrstand th natur and signifian of natural sltirnin ndlian did not dampn Halt
..I know th
populations. Indd' th ook is known today prinrarily for its itial om- h wrot,
prison of Fishr's, !right's,and Hldn's own mathmatial plorations And it an hurt on 1
of this issu. But th ook is muh mor than tht. Haldan aomplishsa spot of laughtr, I an
synthsisnot mrly of Darwinism and ndlian hrdity, ut also, undr do our it i To hlp tl
th ruri of population gntis, of hromosomal mhanis, ytology, and dsrid as largr th
biohmistry. wit, and his politial
During his tnur at amridg, Haldan's work was by no mans r. som.
stritd to volutiotl' |n 1'923, ldane pulishd his famous Dedls, or
BlLIoGRAPH
Sine nd th Fture, a short popular ook on sin in its soial ontxt.
Th ook is said to hav inflund aldan's frind Aldous[{,u|e,s Brue lark, R. !. 1968./B5
ann.
N World (192). Haldan furthr ontriutd Haldan's prinipl, th
H a l d a n J' . B . S . 1 9 2
ida that shr siz oftn dfins what odily quipmnt animals rquir, in Trnh,nd Tru
hls |927 ssay ..On Bing th Right Siz''' in th olltion Possibl Worlds l924.A mth
d other sss' |n 1929, Haldan's spulationson th origins of lif wr rstiosof tb
prsint,oiniding with thos of Alksandr Oparin (1894_1980).In 1930, |927. Possibl
aldan's zmes laid down th foundations of nzym kintis. Dspit 1929.Th orig
193.ztlt
his frnti profssional pa, Haldan wd harlott Frankn Burghs in
|932. h t
1924. This asd som ontrovrsv aus hrlo was marrid at th 1934.A mtlr
tim (to Jak Burghs) arrd had to divor for marrying Haldan. j. B. s. 19 472429.
latr divord harlott and marrid iologist Hln Spurwa i |945. 1937.Th ff
Haldan lft amridg for Univrsity ollg, London, and am 1957.Th os
Haldan,J. B. s.' A. D.
hair of Biomtry in 1937. Also in 197,Ha|dan pulishd his pap, ..Th
Jourl of Gcti
ffts of Variation on Fitnss,'' arguing that th dlin of th avrag fitnss
of a population dr.rto mutation of th optiml alll was proportional to th
mutation rat but not to th indud drsin fitss.Haldan's onlusion
..prinipl Hamilton, W.D,
would latr alld th of th mutation load.'' At this tim, Hal-
dan bam a mmr of th ommunist Party and, in fat, wrot a numr William Donald Ha:
of politial paprs on N4arism for The DiI Worker. |n 1950, Haldan th twntith ntur
rk with th prty du largly t<lLysnkoism and to th rims of Sovit Darwin, partly a
ladr Josph Stalin. This was a priod irr th Sovit Union during whih si- Partly aus oth
n was politlizd and many dfndrsof Darwin Wr snt to th gulags. Prhaps th lagst
Haldan was i1warddth Darwin dl of th Royal Soity in 1953, ats that nfit oth
and h was wardd an honorary LLD from th Univrsity of Paris in 1956, ory suggststhat mu
Haldan lft Camridge in 1957 and migratd to India and ultimatly - wip out altruism,
am an Indian itizn. Th mov Ws a protst of British poliy during th whil th slfishindi
Suz risis. In India, Haldan Was a rsarhprofssor at th Indian Statisti- natur. Afrian kin1

lian hrdity.Indd, Haldan
ortialpopulation gntis. In
of 10 mathnratialpaprs -
Mndlism; th last in th sris
r puliation of his ook e
aldn'ssrisof paprs on vo_
o f t h o r t i apI o p u l t i o n g n -
his ontmporaris;h wntd
natural sltionin N{ndlian
ry pimartly fo its ritial om-
own mathmatialxplorations
n that. Haldan aomplishs a
dlian hrdity,ut also, undr
sonrlmhanis,ytology, and
's work was by no mans r-
lishd his famous Dedls, or
: on sinin its soial ontxt.
's
frind Aldous tlu|e,s Bru
ibutd Haldan's prinipl, th
y quipmnt animals rquir, in
a th olltion Possibl Worlds
ationson th origins of lif wr
r Oparin (1894-1980). In 1930,
ns of nzym kintis. Dspit
l harlott Frankn Burghs in
l harlott was marrid at th
formarring Haldan. J. B. s.
st ln Spurwy in 1945.
ol1g,London, and am
,.Th
ldan publishdhis papr,
th dlinof th avragfitnss
LalalllWas proportional to th
: in fitnss.Haldan's onlusion
utation load.'' At this tim, Hal-
Ltyand, in fat' wrot a numr
ill Workr. In 1950, aldan
lism and to th rims of Sovit
: Sovit Union during whih si-
)arwin Wrsnt to th gulags.
l of th Royal Soity in 1953,
th Univrsity of Paris in 1956.
rtd to India nd ultimatly -
)tstof British poliy during th
t DofSsort th Indian Statisti-
Hmilto
al Institut.In his famous pap, ..Th ost of Natural Sltion'' (1957),
Haldan argud that thr is a ap on th rat of adaptiv volutiorri.tus
of sustitutionl<lad,that is, usa sustitution (y mr.rtationor hang
in nvironmnt)ould not start mor frquntly than vry 300 gnrations.
Haldan'slaim was latr alld Haldan's dilmma.
Haldan was awrdd th Drwin-.!llaMdal of th Royal Soit in
1 9 5 8 , a n d i n 1 9 6 | r i v dt h K i m r d a l o f t h L I . S .N t i o n a l A d -
my of Sins.Aftr a prodigious arr)ldn did of olon an in
Bhubanswar' Orissa, India, on Dmr 1,,|964' Srious illnss, howvr'
did not dampn l-lldan's spirit. In his por, ..anr Is a Funny Thing'''
h wrot' ,.I know that anr oftn kills / But so do ars and slping pills /
And it an hurt on till on swats / So an ad tth and unpaid dts / A
spot of laughtr, I am sur / oftn lratson's ur / So lt us patints
do our it / To hlp th surgons mak us fit'' (lark 1968). Haldan has n
dsridas larg than lif du to his srvi i th Blak Wath, his famous
wit, nd his politial viws. Haldan Was' tO sur, iologist anl thn
som.
BIBLIOGRAPHY
lark, R. \).1968..|BS:h Lif d Worh rf .|.B. S, Hldna.Lodon: oward-
Mann.
Hldan,J. B. S. 192-3.Ddlus,rlr Sid the l-utur.Lonlon:KgarrPul,
Trnh,and Trunr.
1924'^ mtlrmatial thovof naturaland artifiials|tion, I.
rstionsof th mbridgcPhilosopiISlict23: 19_41'
1927.PossihlWorldst| othr ss1,5. London: h:ltto and Win]us.
\929.The <lriginof IiI.Rtiolistttl:.
190.zms.London:Longmans.
192.Th ssof uolution.London:Longmns.
1934. matlrmatial thoof naturalad rtifiials|tion' X' Geetis
1.9:,41.2429.
|97. Te fftsof variationon fitnss.AmrinNtrlist71 37_49.
1957. Te ostof naturalsltion. Jourlof Getis.5.5: 511-.524.
H a l d a n J,. B . s . , A . D . S p r u n t, n d N . . a l d a n 1
. 9 1 5 .R d u p l i a t i oinl l m i .
.|rlrl of Gntis5: 133-135. -R.'s.
Hamilton,W. D. (1936-2000I
!7illiam Donald Hamilton was on of th major volutiorraryiologists of
th twntith ntuy. any of his ontmpor:risompard lrim to harls
Darwin' partly ausoth rrrnshard dp lov of natural history and
partly aus both attemptd to solv som of iology's iggst mystris.
Prhaps th largst of ths mystrisis th qustion of altruism, prforn-ring
ats that nfitoths at a ost to th altruist. Strprfiilly,volutionary th-
ory suggeststh21tmutants hilliting mor ..slfish''hzrviorshould zrlto
wip out altruisrn, aus y dfinition th altruists hv rdud fitnss
whil th slfishindividuals prospr at thir pns.Yt altruism aounds in
natu. Afrian kingfishrs fd and prott young that ar not thir own;
60 Htilto

lronyesommit suiid whn thy sting intrudrs, and vn mor notaly sltir.prssursmust fl
from an volutionary prsptiv' workr s ar stril, sarifiing thir prodution still wins in th
own rprodution to hlp th hiv flurish. i n d e i l L r s i n gp p u l a t i o n
Harnilton's ir-rsightfor solvir.rgtlr puzzl rvirs dptivl1,simpl: slf- vid suh flututing sl
sarifiial vior an volv and prsist if it nfitsrlativsof th altruist attak as thir hosts vol
austhy shar th altruist'sgns.Thus aiding a siling, ousin, or othr th ontinual arrag of i
rlation an still favord y sltior-r' vn if it nrans forgoirrg th pro- ual rprodution advanta
dution of th altruist's own offspring. Hanriltn xprssdthis in a qua- tlrn turnd his :rtt
tion that lras om to alld Hrilton's rul: ltruism is favord whn ing tlrat thr, too, prsst
lc>1'lr, whr is th nfit, is th ost of th at, and r is th dgrof tion of showy sxual orn
rlatdnsstwn th dnor and th ripint of tlr altruisr. on should malsrsistant rl plrrlsit
thrfor b ror likly to s th most s.lrififor thos most losly r- flashy tr:ritsas indiirtors
latd, sulr as silings, nd lss sarififlr thos lss losly rlatd, suh as as w,ll.Parasits lrr of
third ousins. Th pross is alld kirr sltion,arrd th ida has n usd u d p r s s u r t r l m a i n i
lvith grat sussto plain a varity of prosssin animals, inluding th hang thir dfnss,tht
ooprativ irds nd suiidl smntiond rlir, although it dos not al- ass. Ftnalsthus s
ways provid a omplt answr. T s,it turns out' ar though a gnti th spifi gns onfr
puliarity of thir inst group vry lt>slyrlatd to thir hiv mts, mor n tstdin nurrlrous
so than thy rvould to thir own offspring, and h.nhlping thir mothr ata1,parasitsdl s
(th qun) to ris thir sistrsand rotlrrs,who rvill om fr.rtr-tr rpro- vlopnrnt of som su
dutivs,will favord undr manv irumstns. Ntul histor-v,part
In addition to his work on altruisr, amilton wrot aout th volution arilton, and his thor
of animal groups in
..Gomtry for
th Slfish Hrd'' (|971), aout sx ra- organisms havdin th
tios, aout aging and snsn'and :rout th origins and funtion of o- of 6, following orpli
opration and fightir-rg.uh of lris work is mllrkd y a r,villingnssto ongo to follow up tln
ntrtain nw idas and viw old inforrtion witlr a frsh y' and h was a polio vain. H won
rathr solitary sintistthroughout most of his lif. Th ida of kin sltion Kyoto Priz, th rafo
wnt rlativly unappritd for svrirlyars ftr its initial pr"rlitionin mdial iology), tlr Di
7964,ut vntully it ron f th ornstonsof mdrn volution- ml Bhvior Slit1,
ary ilogy' His ids almost always am ky omponnts of thir fild, Soity of Natuarlists.
whthr immdiatly or aftr a lag whil othr sintistsaught up to him. iplirrs,from ntomol
Hamilton vntually .rmintrigu1with on oI th gratstprolms of furthr tstimon to thr
vollttion' th xistnof sual rprllr.trion. This ld to an intrstin p- B|LIoGRAPl{Y
ilsits and thir ffts or-rhost olog,v,havior, nd volutiln that lastd
for th rst of his arr. S is a onundrum aus,just as th altruist H a m i l t o n ,W . D . 1 9 7 1 G'
1:29.5-311'
slrould los in a ra with a slfish onlptitor' an :rsualspis (on that 197.5.Gmblrss
simply rpodr'rsitslf rnaking gtltiallyidntil opis) shltrldout- rf BIogl'50:l7.5-1
ompt a sual spis tht rquirs two individuals to produ a singl 1 9 8 0 .S v r s u
offspring. rtainly som organisms' ir-rludingsorrustaans!lizards, in- 1995a.NrroF
sts, and plants' do rprlduasully. ost, howvr' pay th pri of Nw York: \(. H. Fr
1 9 9 5 .N r r o I
s, with all of its onsquns.As Harrrilton askd in a 1975 ook rviw, P
..Why all .Wy Oird Llnivrsitr'
tlris silly rigmtrrolof s? Why this gavottof hrot-l.losoms? 2005. N.lrr Rr
:rll ths uslssmls, this striving nl wastful loodshd, ths grotsqu Hmilto.Vol.3, La
horns, olors, . . . and why' in th nd, novls . . . ahout lov?'' (p. 175).
Hamilton, long with a trv othr sintists,suggstdthat sltionmust
favor th shuff1ingof gnotypstrvt-t gnrirtiol.ts
and hrrsx. But th

g intrudrs,and vn mor notaly
r s ar stril' sarifiing thir
h.
tzzl wils dptivly simpl: slf-
if it nfitsrltivsof th altruist
us aiding a siling' ousin, or othr
vrrif it mns forgoing th pro_
l : r r n i l r on p r s s dt h i s i n n q u a _
ln's rul: altruism is favord whn
lst of th at, an1r is th dgre of
ipintlf th altrr'rism.On sould
sarififor tlros most losly r-
)r thoslss losly rlatd, stth as
ltion,nd th id has n usd
prosssin ninrals,inluding th
lndarlir,altlroughit dos not al-
;, it turns out' ar through a gnti
,ly rlatdto thir hiv mats, mor
ing,nd hnhlping thir mothr
| r s \' ^ ' I 1r ov i | |l l r l l n f u t u r r p o -
l l]1stns.
arr.riltonwrtlt aout th volution
lfishrd'' (1977), aout s-ra-
out th lriginsand funtion t;f o-
lrk is markd y a willingrrss to
tion with a frslr y' and h was a
f his lif. Th ida of kin sltion
yars aftr its initial pulition in
ornst()ns of nrodrn volution-
rrn ky omponnts of thir fild,
thrsintistsaught up to him.
vit on of th gratstprolrns of
- . t i t l T' h i s I dt t l n i n t r s ti n p
lhavior,and volution tlrirt lastd
rdrum aus,just as th altruist
titor,an asul spis (one that
illr' idntil opis) should out.
vo individuls to produ a singl
rdingsorrustaans'lizards, in-
. ost, howvr, pay th pri of
ilton askd i a 7L)75 ook rviw,
hisgvottof hromosoms?
astfulloodshd,hsgrorsqu
v l s. . . o u t l o v ? ' ' ( p . 1 7 5 ) .
t i s t s .s u g g s t dt h t s | t i o nm u s
]ntionsand lrn sx. But th
Hmilton 61
sltivprssursmust flutuat in ithr spa or tim, or ls asxual r-
produtin still wins in th long trm. For a varity of rasons st plaind
irrdtail sing.loulationgnti modls, prasitsar idaliy suitd to pro-
vid suh flutuating sltionausthy ontinually volv nw mans of
attak as thir hosts volv dfnssagainst thm. Hamilton outlind how
thontinual rrg of insults y pathognslirrgand small ould kp s-
uaIrprodution zrdvntgousirr long_livdmultillular organisms.
H thn turnd his attntirrt<lsual sltionand rrrathoi, suggst.
ing that thr, too' prssurfrom parasitssrvsa ruial rol in th volu-
tion of showy sr.ralornamnts lik th til of th paok. If only healthy
malsrsistantto pi1rilsitsan produ ths Ornmnts'trnalsn us th
flashytraits as indiators of gr-rs
that will hlp mak thir offspring rsistant
as r,r,ll.
Parasits a of partiular signifianausthy provid Ontin-
ud prssur to intain gnti variaility; just as with sx, unlss hosts
hangthir dfnss,thy will ovrrun y th rapid volution of thir dis-
ass.Fmals tlrus us th orlllnts to 6rrd rsistant mts, evn though
th spifi gns onfrring this rsistan hang ovr tirri. This ida has
.shil
ntstdin numrous animal spis. parts of his thory rmain d-
batabl'parasitsdo sm to play a signifiantrol in mt hoi and th d-
vlopmntof srlm sual signals.
Natual history, prtiul:rrly of insts, was always <lf kn intrst to
Harnilton' and hrs thory was lways groundd in a sund knowldg of how
organisms havd in th ral world. H did in 2000 at th rathr ear| age
of 63, following onrpliationsof malari ontatdon an pdition to th
ongo to follow up On a ontrovrsial tlroy that AIDS aos from faulty
polio vain. H won many lf iology's prizs and awards, inluding th
Kyoto Priz, th rafoord Priz (Swdn'sNol Priz quivalnt for non-
mdialiology), t DistinguislrdAniml Bhaviorist Award from th Ani-
ml Bhavior S;ity,nd th Swall Wright Award fronr th Amrin
Soityof Naturalists. Baushis work inflund so many filds,many dis-
iplins,from ntomology to parasitology, laim him as on of thir own,
furtlrrtstimony to th road rah of volution in sin.
-
BIBLIOGRAPHY
Hamilton'!. D. 1971.Gomtrl'for th slfishhrd'Jlurllf horetilBiolog
3 1 :2 9 5 - 3 11 .
1975.Gmlrssinlif bgan:Barnls'aphids,e|ms.QrtrlRet'itl.l
tli Billrlg50: 17.5_180.
1980.S vrsusnon-svrsusrilsit.oi&s 3.5:282._29.
|995a.Nrro Rodsof GLd. oI, 1''uolutiltlof SoiIBehuior.
Nw York: W. H. Frman.
199.5b.Nrro Rodsof G Ld.o|' 2, uolutioof Se.oford:
.Wlry oford Univrsitl.Prss.
2005.NtrrlRods tlf G Ld: h olltdP"lprsof V. D.
Hmilto.Vol. 3, Lasl V(/ortls.
ark Ridlv,d. Oxford: oford UnivrsityPrss.
-.Z.
62 ennig

Long for Hnni1

Hnnig, Willi (913-1976)
nss of two spisW;
In omptition with arl Linnaus, \Willi Hnnig is prhaps th rnost influntial havior, and othr l
systmatist of all tim. Aftr harls Darwin's presntation of th thor of voivd simiiarit, t
volution, many biologists, inluding Darwin himslf, xprssd th ida that nalogial rlationsh
organisms should lassifid asd on thir volutionary rlationships-thir hritd from a omm'
phylogny. Flowvr, it was Hnnig who first prsntd a omplt thory for ritria; only th shar'
phylogny ronstrution and dvloped th prinipls fo phylogny-basd nti rlationship. B
lassifiation. tor shard y rrly
As a tnager, nnig Was an ardnt amatur zoologist. H olltd - volutionary unit (m.
tls and uttrflis, startd an hrarium' and workd s a vlunt in th (plsiomorphis)1o r
Drsdn usum. took an intrst in th sin of systmatis and, in a himp, and man, tht
high shool ssay writtn at th ag of 18, h larly idntifid th ronstru- that thy shar a tail
tion of volutionary rlationshipsas th main task of the disiplin.Biologists tor of all four spi
at th tim lakd a c|ear undrstanding of how organismal rlationships shard divd fatu
ould b rlialy ronstrutd, ut Hnnig disussd th mrits of diffrnt To Hnnig, asin1
typs of vidn of phylognti affinity and psntd original idas that namd taa must b
forshadowd his latr work. Howvr, it would tak him lmost tw dads th tim by rsarl-r
to dvlop a rigorous thory of phylognti infn. must b a omPromi
Hnnig studid zoo|ogy, botany, and gology at th Univrsity of Lipzig, ity (..volutionary s
whr h dfndd his thsis on th opulatory apparatus of Diptra (flis, ( 1 9 6 ) , w h oa r g u dt
m o s q u i t o s ,a n d m i d g s )t n t 9 3 6 , a t t h a g o f 2 . | n | 9 3 7 , h j o i n d t h phylogntirlatitln
Grman ntomologial Institut, whr h spnt most of his arly profs- nig's prinipls ar a
sional arr. Although omparativ morphology and taonomy of diffrnt dads for a on
groups of Diptra rmaind th maio fous of his work, Hnnig Was a pro- aus of th rsilirr
..p
lifi writr, pulishing mor than 9,000 pags and thousnds of drawings nosprms,',arrd
during his lifetim. In addition to th Diptra monographs, his writings in- Th ruls for namin
ludd outstanding thortial and mpirial ontriutions as wll as txt- ida of rognizing f.
books spanning th ntir fild of zoologial systmatis. that ranks tid t
.War
Hnnig was onsriptd as an infantryman whn Norld II rok out dat of origin. Aul
in 199. Aftr rovringfrom a srious injury h suffrdin Russia tn |942, tim, ut th rfin
h was ordrd to work with malaria ontrol. Shortly fo th nd of th molular data my
war he was snt to Gr and northn Italy' whr, in British aptivity' h Hrrnig disttss
ontinud with malaria work until th fall of 1945,It was during this priod sortd from honrolt
that h wrot th draft of his most important work, Grudzge ier Tborie inludd dtaild ar
der phlogeetisch Sstemtik (Pricipls of Theor of Phlogeneti Ss- atrs in th fssil r
tmtis' 1950). and th gographi
Hnnig finishd th manusript aftr th war, and th ook appard in rsolvd using h
1950 in a small dition and without an ind aus of th shortag o{ against ah othr. ,
papr at the tim. Th txt ws widly disussdmong Gmn-radingzo- asnof vidn
ologists, inluding Profssor Lars Brundin at th Swdish usum of Nat- iary prinipl. Henl
ural History in Stokholm' who am a powrful proponnt of th nw onflits' suh as p:
idas. owvr, Hnnig's thoris wr not introdud to the nglish sin- point toward thrn
tifi litratur until a rvisd vrsion of rh book was printd in th Unitd ida to Hnnig. H
Stats in 1966; severalrlatd paprs appard in nglish-languagjournals indpndntly dis
around th samtim. molular iologist

g is prhapsth most influntial
s prsntationof th thory of
himslf' prssd th ida that
volutionary lationships-thir
prsntda omplt thory for
pinipls for phylogny-asd
]ur zoologist. H olltd -
d workd as a voluntr in th
sinof systmatis and, in a
:larly idntifid th ronstru.
task of th disiplin.Biologists
how ognismal rlationships
lisussdth mrits of diffrnt
d prsntd original idas that
rld tak him almost two dads
irrfrn.
lgy at th Univrsity of Lipzig,
ory apparatus of Diptra (flis,
: of 2.In 797, he joind th
spnt most of his arly prlfs-
logy and taxonomy of diffrnt
of his work, Hnnig was pro-
3s and thousands of drawings
. a m o n o g r p h s ,h i s w i t i n g s i n -
,l ontriutions as wll aS tt-
systmatis.
.!7orld.sar
:rwhn II rok out
:y h suffrd in Russia in 1'942,
ll' Shotly for th nd of th
y, whr, in British aptivity, h
1945.It was during this period
work, Grudzge eier horie
lf Theor of Ph|ogeti Ss-
war, and th ook appard in
d aus of th shortag of
;sdamong Grman-rading zo-
t th Swdish usum of Nat-
owrful proponnt of th nw
introdud to th nglish sin-
ook was printd in th Unitd
.d in nglish-larrguag journals
Heig 6
Long for Hnnig, it was gnrally assumd that th volutionary rlatd-
nss of two spiswas orrlatd with thir ovrall similarity in morphology,
havio,nd otr traits. It ws also widly rognizd tat indpndntly
evolvd similarity, ommonly alld onvrgne' was fals vidn of g-
rralogialrlationship. But Hnnig ralizd that many of th sirrrilaritisin-
hritd from a on-lmon anstor' alld homologis, wr uslss as grouping
ritria; only th shard drivd traits (synapomorphis) indiat tru phylog-
nti rlationship. Baus a synapomorphy originatd in a ommon ans-
t o r s h a r d y o n l y s o m o r g a l ) i s m s .i t g r o u p s t h o s o r g a n i s m s i n t o n
volutionary unit (rnonophyltigroup or lade). Th shd primitiv traits
(plsiomorphis)do not dfin lds. For instan, in a sampl of dog, Itzard,
himp, and man, th dog and tlr lizrd annot b groupd on th ground
that thy shar a tail aus this trait was inhritd from th ommon ans-
tor of ll four spis;it is a plsiomorphy. Th lak of a tail, howvr, is a
shardderivd faturthat n usd to grou th himp and man.
To Hnnig, asing iologial lassifiation on phylogny mant that all
namd taa must b monophylti groups. This id was strongly opposd t
th tim y rsarhrslik rnst gIar (1969), who insistd that lassifiation
rust a ompromis twn volutionary rlationship and ovrall similar-
ity (..volutionary systmtis,'),and y Rot Sokal and Ptr Snath
(196),who argud that grouping y ovrall similarity was prfralaus
phylognti rlationships Wr so diffiult to rOnstrllt (..phntis'').n-
nig's prinipls ar almost univrsally aptd now' vn though it will tak
dds for a onsistnt Hnnigian lassifiation is univrsally adoptd _
aus of th rsilin of old group onpts-suh as ..invrtrats,''..gyn-'-
nosprms,,' nd ..protozoansD-and tlr lak of knowldg aout phylogny.
Tlr ruls for nanring lads ar still mr'rh datd, spially th Linnaan
ida of rognizing formal ranks (gnus,family, odr, t.). Hnnig proposd
that ranks tid to ag; ah rnk would srvdfor groups of a spifi
dat of origin. Aurat dating of lads smd ompltly unralisti at th
tim, u th rfinmnt of phylognti mthods and th growing amount of
molular data may yt lad to a ronsidration of this ida.
Hnnig disussd at lngth th prinipls y whih onvrgn ould
sotd from homology and plsiomorphy from apomorphy. His thniqus
inludd dtaild analysis of similarity, th squn of apparan of har-
atrs in th fossil rord or irr ontogrry (th dvelpmnt from egg to adult),
and th gographi distriution of harats.onflits Wto manually
rsolvd using hking and rhking, arfully wighting th argumnts
againstah othr' A asi ida was that homology should asstrmdin th
asnof vidn to th ontrry' ommonly rfrrd to as Hnnig's auxil-
iary prinipl. rrnig nv disussd quantittiv thniqus for rsolving
onflits, suh as parsimny and ompatiility analysis, ut his tts larly
point toward tlrm, and som arly parsimony paprs attriutd th asi
ida to Hnnig. owvr, th mathmatisof phylogny ronstrution Was
indpndntly disovrd at aout th sam tim y othr systmatists and
molular iologists unwar of Hnnig's work.
64 Hobbit

Th dat following th prsnttion of phylognti systmatis ws long ft, a short ig

plagud y outragous ontrovrsy (Hull 1988). Som of nnig's nmis ratio.
lald him ..ladist'' aus of his phasis on rognizing monophylti To dat, th rmai
groups (lads),and Hnnig's mor zalous followrs quikly adoptd th found in assoiation r
trm. Aftr Hnnig's dath, svral prominnt ladists argud that th - gint Komodo drago
sis of lassifiation should not volutionary rlationship ut th most dnd, fir-rakd r.
pasimonious arrangmnr of th availal dta (Ridly 1'986). othrs ston tools wr
(phylognti systmatists)maintaind that parsimony was only on of sv- tools' and flaking dl:
ral tools that ould usd to find th volutionary rlationships on omo florsisis su1
whih propr lassifiation should st' Dats inspird by Hnnig's rvo- r.rthrdarrd ookt
lutionary book still ontinu today, mor than 50 yars aftr its original ailitis in suh a smz
puliation. y an analysis of th.
LB1's rainas.A
|LloGRAPHY
th rain was higlrly r
Anonymous.2005' !illi Hnrrig.Das BiologiWiki, http://www.iologi.d/ r of advand ft
iowiki/!7illi-nnig. pandd in a rgion t
Dupuis, . 984.!illi Hnnig'simpaton tonomithought.ul Reuitulf
thought and for plr
olog d Sstmtis|5: |-24.
Flsnstin, ontradit th sugg
J.200. A digrssionon historynd philosoph.Inlferring
Phlogeies,hap. 10. Sundrland,MA: SinurAssoiats. human fflitd with
Hnnig, !. 1950. Grudziigir Thorider phIogetishen Sstmtik.Berlin
DutshrZntra|vr\ag.
1966. PbIogntiSstmtis'Urana:Ur-rivrsity
of illinois Prss.
Hull, D. L. 1988'Sies rlcss.
Chigo:Univrsitof hiagoPrss.
ayr, . 1'969.Principlsof SstemtiZotllog. Nw York: MGraw-ill.
Ridly, . 1986. uolutio d lssifiti:Th Rformtioof ldism.
Nw York: Longman.
Shl'D. 1978' Villi nnig 191-1976:in iographishSkizz.ntomollgi
()rmi4:377-391.
Sokal, R. R., nd P. H. A. Snth.1963.Priiplesof Numril T<lnom.San
.W. -F.R.
Franiso: H. Frmn.

Hobit (Homo floresiensis)

..Hoit''
is th niknam (aftrJ. R. R. Tolkin's 1937 novl, Tbe l7obbit)
for a tiny sklton (atalogd as LB1)that rprsntsa nw human spis
(omo floresiesis),whih lived from 95,000 to 12,000 yars ago. This
rmarkl find ws annound tn 2004 y Mihal orwood and othr
sintists from Australia and Indonsia. LB1 was avatd from 18'000-
yar-old dposits in Liang Bua, a av on th Indonsian island of Flors.
Th narly omplt sklton stands a littl ovr a mtr tall, and is b-
livd to rprsnt n adult woman. Th sklton and fragmnts of lvn
othr individuals show tat Homo floresisis manifstd a mitur o{
faturs found in very arly rlativs of humans (.g., ...LuC,',datd to
ovr 3 million yars ago), ear\ Homo' mor rnt Homo r,t||s, and liv-
ing popl. LB].'s omination of faturs of th rain, tth, shouldr, Virtual skullof LB1
wrist, plvis, lgs, and ft has not n found in any oth hominins, liv_ omputrtomgrap
ing or xtint. Homo florsisishad vry thik bons, rlatively short lgs' ndoast that rprod
 obbi 6.1.i

phylognti systmatis was
3 8 ) .S o m o f H n n i g . s n m i s
s on ognizingmonophylti
follows quikly adoptd th
nt ladists argud that th a-
ary rlationship ut th most
: data (Ridle 1986). othrs
,arsimonywas only on of sv-
vlutionry rlationships on
rtesinspird y nnig's rvo-
ran 50 yars aftr its original
<i,http://www.ilogi.d/
mi thought. ul Ruielu of
rilosophy. In Iferrig
rr Assoits.
logtish Sstemtik. Blin:
long ft, a short ig to, a tiny rainas, and an ap-sizd rain-to-ody
ratlo.
To dat, th rmains of twlv Homo florsiensisindividuals hav n
found in assoiationwith faunal rmins of dwarfd lphant-likeStegodo,
giant Komodo dragons, rts. and ats. harrd ons nd lustrs of rd-
dnd, fir.akd roks wr also found in Liang Bua. High dnsitis of
ston tools wr avatd from th av' inluding ors, anvils, rtouhd
tools, and flaking dris. Baus of ths assoiatd finds, th disovrrs of
Ilomo florsiesissuggstthat th spismad tools, hul-rtd,usd fir, and
uthrd and ookd mat. Although th suggstion of highr ognitiv
ailitisin suh a small-raindspisrmins ontrovrsial,it is sr'rpportd
y an analysis of th impssions of tlr rain that wr imprintd insid
LB1's rainas.A ast of th rainas (or ndoast;s figur) rvals that
th rain was highly onvolutd and uniquly organized (wird),with a num-
br of advand faturs.For ampl, th tips of th frontal los wr -
pandd in a gion that is known to impotant for ronitoing intrrral
thought and for planning ahad. Ths and othr dtails of LB1's ndoast
ontradit th suggstion y a fw rsrhrs that it may from a modrn
human afflitd with a pathology known as mirophaly.

lnivrsityof Illinois Prss.

nivrsityof higtl Prss.
\w York: Graw-Hill
Rformtion of ldism.

grphish Sk\zz' totologi

: of Numril onom. San

-F.R.

in's l937 novl, The Hobbit)

: p r s n t sa n w h u m a n s p i s
00 to 12,000 yars ago. This
ihal orwood and oth
I was avatd fom 18'000-
r Indonsian island of Flors.
) ovr a mtr tall. and is -
:lton nd fragmnts of lvn
zsis manifstd a mixtur of
tmans (.g., ..Luy,,' datd to
fecerftl-Iomo ertus, and liv-
of th rain, tth, shouldr, Virtual skull of LB1 (..Hoit'')that Was ronstrutdusing thr-dimnsionl
nd in any othr hominins, liv- omputrtomograplry.Th skull has bn flood-filldto rv:rla virtual
ik ons,rlativly short lgs' endoastthat rpodusth shaoand othr dtailsof ..Hit's''rin.
66 omolog

Th disovr of Homo fllresisis Was tmly surprising aus it twn dissimilar hal
showd tat Homo spis Was not th only hlminin arund 18,000 yars of intrmdiat forr
go (and for),as had n widly livd.Sintistsar mixd on thir trirrtionof sinrilr
intrptationsaout whr rhis nw spism frm. Som tlrink that it is ogy ritria aptur t
a dsndant from iggr.odid Homo rtusallstors that lramdwarfd asd on pattrns of '
aftr migrating to th island of Flors. This hypthsis is in kping with th lotid sustitutiorri
wll-known island rul, whry larg nimals onrsmallr (lik th aov- squns.Belttso
mntiond Stegodo\ and small animals lmlargr (.g.'th giilnt Kornodo volurion, fr.rrtlr
dragons) on islands whr rsours ar limitd. othr workrs liv that whih divrg as a r
Homo florsisismay lrav n dsnddfrom a small anstor' suh as paralogous gns,wh
thos in Luy's genus (tlstrlopithus), nd that its nstors simply r- Th onpt of hor
rnaind small aftr arriving On th islirrrdof Flors. Tim will tll. tirns. Aristotl fotrn
irl' and vn ftlr
|BLloGRAPl{Y
analogy, th idsa
Brown,P.' T. Sutikna,tr4.J. orwod, R. P. Sojono,.|tikmo, FJ.Y/. Saptlmo, and dat tw G
R. . Du..2004.^ nrvsmall-did hominirrfrllth Lt Pliston lf a r l v 1 8 3 0 s ( s A p t
F l o r s I. n d o n s i aN. t ' 4 . 3 1 1
: 0 5 5 - t 0 6l .
st undrstoldin tL
Flk, D., . ildolt,K. Srnith'..|. tlrwood,. Srrtikn, P. Bror,vn,Jatmiko,
. W. Saptml, B. Brutlsdn,and F. Piro.200.5.Th rainof LB|, Homl ments) nd within th
floresisis'Si8:2"42_24 5. Goffroy argud fr
lv{orwood,.J.. P. Brown'Jatikmo,T. Strtikn'. W. Saptomo,. . wstaway' plarr of all ninrals,
R. A. Dr"r, R. G. Rlbts, T. da' S. Wsisttl'nd . Diubintonl. 200.5. rognizd thirt idn
Furthrvidnfor small-bodid homininsfrm tll l,tlistln f Flors.
-D.F. arranging diftrnt ft
I n d o n s iN . I r 4 3 7 : | | 2 _ | 0 | 7 .
vids a rational syst
th rrotion of th ar
similritis 1m()ng al
Homology
In th ontt lf t]
Homology is on of th ntral onpts in iology. It rfrs to th similarity tnth ntury' th h
twnmorphologial haratrs'gtls'and othr iologial rraits, suh as th osrvd pttrn
haviors,al-nongdiffrrrtspis.Srn of th st-krrownnrplsof ho- in thir ommolt vr>
mology ar (1)th wings of irds, th arms lf htlnlans, and tlr forlims of for th ronstrutio
othr ttrapods and (2) th ony lmntsin th gill rhs of sharks and th phologial ritia,
thr middl ar ons (mallus,inus, nd staps)in humans. Ir-roth ass, asrtain homology r
th similaity btwn haratrs is stlishd indpndntly of thir siz, ws sd on th ssl
shap, and funtion. This fat is rfltd in Rihirrd own's dfinition of a si.rilarmryolrian
homologu as ..th sam orgn in diffrnt animls und vry varity of ogy would soon run i
form and funtion''' to distinguishdfrom an analogu as ..a part Or Organ gl ll, it ws not lt
in on animal whih has tlr sam funti<lnas anothr part or lrganin a dif- samnsstwntrv
frnt amphibian lss,lf
What is alady lar trom Own's dfinition is that in ordr to stalish th sion that th original l
istn of homology, w nd to hav a st of ritria tht allow us to dtr- prolemati.Anothr
nrin th samnss btwn diffrnt lraratrs. Irr th past' omparativ Th rognition that l
anatomy arrd mryology providd th tools for lromology assssmnts;tody ph ologial strtlturs
w also us molular and sequndata. Th morphologial homology ritriir tu.n haviors ws
inlud (1) position-homology is mor likly if two haratrs ar found in Today, th homol
sirnilar positiorr within two organisms; (2) strutur-lromology is mor likly if disr.rssions withirr i<
two haratrs shar many individital faturs; (3) trar-rsition-homology - irls nd dfinitionsof ]

]mlysurprising aus tt
lminin arrrund 18'000 ars
Silrtistsr rixd on thir
l f r m .S o m t h i n k t h a t i t i s
[storstht arndwarfd
othsisis in kping with th
:omsmallr(lik th aov-
Largr(.g.,th giant Komodo
l. othr wlrkrs bliv that
om a small allstor' suh as
tht its anstots simply r_
rs. llm wlll tll.
.rt tt
, Jtikmo' . W. Sptomo. and
i r o n l t I l l . t l ' l i s t o
. Sutikn,P. Brwn, Jatmiko,
5. Th rin of L'B|, Homo
W. Sirpttlrno,K. ..Wstwur'
l, nlT. Djuiantono. 200-5.
'm th Lt Plistonof Flors,
llogy. It rfrsto th similarity
othr iologial traits' suh as
ampls of ho-
l st-kr.rown
and th forlims of
lrurans,
h gill arhsof sharks nd th
lps) in humans. In oth ass,
d indpndntlyof thir siz,
R.ihardown's dfinition of a
rnimals undr vry varity of
.'a part o organ
tn anloguas
nothrpart or rgan in a dif-
r is tlrat in ordr to stalish th
that allow us t dtr-
lf ritri:-r
[n th pest, omprativ
trs.
assssmnts; today
lr homology
ritria
rrorphologiallromology
if two haratrs ar found in a
tur-homology is mor likly if
s; (3) transition-homlogy -
oologv 6.7
twlr dissilrrilarlrartsan stalishdif thy ar linkd through a sris
of intrldiat fornrs; and (4) svrrlongrun ritria tht apttlr th dis-
tiuti f simila hrats in losly rlatd spis.Tlr rnolular homol-
ogy ritria aptur th similarity rwn moluls, suh s DNA squns'
asd on pattrns of ommon dsnt.Hr th nmbr nd th position of nu.
lotid sustitution ar oftn usd as n approimation fr th losnssof rwo
squns.Baus of th high frquny of gn dupliations in th ours of
volution, a furthr distintion has to mad btwn orthologous gns,
whilr divrg as a rsult of indpndnt volution aft a spiation vt' nd
paralogous gns,whih divrg within ;.rspisafter a gn dupliatiorr vnt.
Th orrptof homology dats ak to pr-Dawinian and pr_volutionary
tirns' Aristotl found similarity twn orgnisms to somthing noti-
al, nd vn bfr Own fornrlly dfind th npts of hmlogy and
anallgy,th idas apturd y thseonptswr at th hart of th famous
dat twn Gorgs uvir and tienn Goffroy Saint-Hilir in th
n f arly 1830s (sAppl 1987I. Whil uvir hld that th divrsity of lif is
st undrstoldin th ontt of distint highr groups (so-alldmbrh-
mets) and within thos groups asd on th funtional nds of orgnisms,
Gffroy agud for th istnof an undriyirrgrmmon organizational
plan of all al-rimals,whih lr alld univrsal analogy or unity of ryp. Own
rognizd tlrat idntifying orspnding parts in diffrnt spimns and
arrangir-rg differnt forms aording to a nstd hirarhy of similaritis pro-
_D.F.
vids a rtional systm for ordring th divrsity of lif' H also introdud
th notion of th arhtyp as an abstrat rprsnttionof th undrlying
simiiaritis among e group f organisms.
In th ontt of th Darwinian rvolution of th sond half of th nin-
tnth ntury, th homology onpt ',vas historiizd. Th xplanation for
t osrvd pattrns of similarity twn organisms Was l1ow to found
in thir omlnon volutionar history, and lromologis nr th main tool
for th ronstrution of ths phylogrrtirlatinships. Along wit or-
phologial ritria, mbryologilvidnwas also inr:rsinglymployd to
.his
asrtain homology rlationships twn diffrnt struturs. prti
ws based on th ssumption tht sin-rilarstrutursshould also dvlop from
similar rnryoni nlagn. Howvr, this program of volutionary morphol-
ogy would soon run into diffiultis.First, if ll strutursdvlop from a sin-
gl ll, it was ntlt lar whilr mryoni stagsshould b usd to estalish
saftlnsstwll twl adtrlt struturs. Sond, rgnrating tissus, suh as
amphiin lnss'oftn dvlop fro a diffrnt souf of lls. Th onlu-
sion that th originIand tlr rgnrtdlns a nt hmollgotlswas larly
prolmti.Anothr appliation of th l-romologyonpt was mo fruitfl.
Th rognition that haviorl pttrns an tratd th sam wy as mor-
phologial strutursnd that on an thrforstalishhomologis also -
twn lrviors was th foundation of th sin of thology.
Today, th homlogy Onpt is again at th ntr of svral thortial
disussionswithin iology' This as rsultd in a varity of dif{rntpropos-
ls nd dfinitions of orrrologythat an ll groupd' aording to th rsarh
68 Hookr

qustions thy addrss,into (1) historial homology onpts nd (2) iolog- y a x p l o a t o r y! o y a

ial homology onpts. istorial homology onpts ar prominnt in susquntlyprdud
phylogny ronstrution. Th most widly usd approah, ladistis or 1 8 4 7 , h t r a v l dr S i l
phylgntisystmatis,haratrizsgroups of spis(lads)y th prs. oti plants suh as r!
n of shard drivd haratrs (synapomorphis).Ths synapomorphis p a n g a r d n s .H p u l
ar historial h<lmologus-thir samnssis a onsqrrnof thir phylog- am assistant to his
nti (anstor/dsndant) rlatiorrships.Th rsulting hlognti systl dirtor, ontinuing (
rprsnts a nstd hirarhy of groups, ah haratrized y rr uniqu st of of th British mpir. I
shard drived haratrsor homologus. ln \844, Darwin inf
. . s s a y , 'i
Th iologil homology onpt' on th othr hand, also mphasizes,in snr him th
addition to th ommon phylognti history, th dvlopmntal origin of two argud for many y
homologous struturs.Th similaritisof phnotypi strutursar snas a distriution of plants.
'sfagnr
onsqun of a shard dvlopmr-rtalprogram. As Giintr dfind anint lnd onntio
it: ,.Strutursfrom two individuals or from th sam individual ar honrolo- q u n t l y ns u n l r g
gous if thy shar a st of dvlopmntalonstraints,ausd y loally ating dfnding th viw th
slf-gulatorymhanisms of . . . diffrntition'' (Y/agner 1989, 62|. he rrr hmisphr, of wh
iologial homology onpt thus not only apturs th distriution of ho- sattrdrmnants. D:
mologus among groups of spis'ut also intnds to xplain th mhanis- th oans y flotatiol
ti aussfor th osrvd similaritis' whih ar to b found in th dtails of v o | u t i o n i s m , u t v r
th dvlopnrntalsystr-n. Th iologial omology onpt is trfor at th thory.
or of volutionary dvlopmntal iology' th rnt synthsis of volution- Along with harls
ary and dvlopmntaliology. h should tk ftr
i n r r o d u t o r y s s vt o h
lBLIoGRAPHY
p u l i n d o r s m n t os
Appl, . 1987. h uuir-GeoffroDebte:Frnb Biollgin t Dcdsbefore t h f a m o u s d a t w i t t
Dttui.oford: Oford UnivrsityPrss. th British Assoiation
Bok, G. R., and G. ardw,eds,1999.omolog.Novatis}.oundeltion
Was mor prsuasiv.
Symposium222. htestr,U.K.: I/ily.
all, B. K., ed. 1994. Ilomolog: Th HirrhiIBsisof omprtiueBiolog. Darwin and Hook
Sn Dig:AadmiPrss. dads.In 1878' Ho
Lauihlr'M. D. 2000.Homologyin dvlopment and th dvlopmnt
of th of London to dfr-rdh
hmologyonpt.mri Zoologist 4:777_788. Sphr. At th sam tin
own, R. 1.843.I'turso th omprtiuAtom d Phsiolog1'of t
o t h r st h r t h m i n s <
Iuertbrtnimls' DIiuerdt th Rol ollgof Surgrs.Londr:
Longman,Brown' Grn,and Longmans. g i o n s f u a s i n
!agnr,G. P. 1989. Th iologilhorlgyollpt.ul Rui,of colog away from th thory
d Sstmtis2:5 |-69 , -.D.L. Wlla, anl othr su
and otls.

Hookr, Josph Dalton (181'7-1911I BIBLIOGRAPH

Josph Daltorr L{ookr was a lading otanist of Vitorian Britain and on of Alan, NI. 1967.Th Hl
harls Drwin's staunhst supportrs. H was on of th first sintists to I{ookr,J. D. |845, he
Disot,rShipsre
informd of Darwin's thory and playd a lading rol in hlping Darwin Bros.
to formuiat his idas, spially in th appliation of volution thory to | 8 5 5 .F l o r l d
ioggraphy. Idi. I-ondon:\/ . 1
Hookr ws orn on Jun 30, 1817, th son of Sir \WilliamJakson F{ookr, 1860. FIor ,ls
who was dirto of th Royal Botnial Gardns at Kw. otaind an
MD at Glasgow in 189 and latr that yar joind }i.S rbus on a four-
 Hrloker 6L)
I

mology onptsand (2) iolog- yar ploratory voyag to Nw Zea|and, Tasmania, and th Antrti. H
O g y o n p t s r p r o m i n n i n susquntlyprodud a six-volum survy of th otany of ths rgions. In
[y usd approah, ladistis or 1847,e travldto Sikkim, Npal, and th astrn Himalayas, snding ak
ls of spis(lads)y th prs_ xoti plants suh as rhododndrons' whih soon am favorits in uro-
orphis).Ths synapomorphis pan gardns. H pulisd Flor Idi in 1855, th yar in whih h -
i onsqunof thir phylog- am assistant to his fathr t Kw. In 1865, h sudd his fathr as
l r s u l t i n gp h y l o g n t i s y s t m diretor, ontinuing Kw's rol as a lading entr for th onomi otany
haratrizdy a uniqu st of of th British mpir. Hook did in 1911.
|n 1'844, Darwin informd Hookr of his idas on volution and in 1'847
o t h r h a n d , I s o m p h a s i z s ,i n ..ssay'' in whih his thory was first dsried in dtail. Th
snt him th
ry' th dvlopmntal origin of two argud for many yars ovr th thory's impliations for th gographi
rnotypistrutursar sn as a distribution of plants. ookr livd that disprsal had takn pla ovr
1ram.As Girntr.!7agnr dfind anint land onntions twn th isting ontinnts' whih had sus-
th sam individual ar homolo- quntly n sumrgdy gologial ativity. He was partiularly ativ in
straints,ausd y loally ating dfnding th viw that thr had on n a grat ontinnt in th south-
ation'' (Wag'er 1989, 62). Te rn hmisphr, of whih Antartia, Australia, and Nw Zea|and ar ut
aptursth distriution of h_ sattrdrmnants. Darwin argud instd for th disprsal of sds aross
intndsto plain th mhanis- th oans y flotation, winds, and irds. Hookr was at first skptial of
l ar to found in th dtails of volutionism, ut vntually h am on of th arlist onvrts to th
nology onpt is thrfor at th thory.
th rnt synthsis of volution. Alng with harls Lyll, Hookr advisd Darwin on th ours of ation
h should tak aftr reiving Alfrd Russl Iil7alla'spapr in 1858. Hookr,s
introdutory ssay to is Flor Tsmnie of 1860 was on of th first major
puli ndorsmnts of th thory. H spok aftr Thomas Hnry Huly in
ch Biolog i tb Deds befor th famous dat with Bishop Samul Wilrfor at th oxford mting of
th British Assoiation in 1860, and som obsrvrsthought his ontriutin
gy. Novartis F-oundation
was mor prsuaslv.
l Bsis of omprtiu Biolog. Darwin and Hookr ontinud to dat iogography in th following
dads.In 1878, Hookr usd his presidntial addss to th Royal Soity
:nt and th dvlopmnt of th of London to dfnd his thory of an anint ontinnt in th southrn hmi-
777-788.
sphr. At th sam tim' h ndorsd th viw favord y Darwin and many
,tom d Phsbklg of the
othrs that th main sour of mor highly volvd spis Was th northrn
| ollg of Srgeos. London:
rgions of urasia and North Amria. Ltr in his lif' he gan to ak
nept.
nlReuietu away from th thory of sunkn land ridgs' undr prssrrrfrom Darwin,
"'\i}'.il',. !alla, and othr supportrs spousing th prmann of th ontinnts
and oans.

r.) BlLIoGRAPHY

lt of Vitorian Britain and on of Alan, M. 1967.h Hookrsof K' London:ihalJosph.

Was on of th first sintists to Hookr,J. D. 1845. Th rptogmiBltof the trti Vogof H..
DisourSlpsrusderror i th Yrs18']9_4.6vols.London:Rv
a lading rol in hlping Darwin
Bros.
pliation of volution thory to 1855.Flor Indi:Big Sstmticout f th Plts of British
.!.
Irtdi. London: Pamplin.
n of Sir \WilliamJakson Hookr' 1'860. Flor Tsmie. Londn: L. Rv. -P.J.B.
ardns at Kw. H otaind an
. joind T1S rbus on a four-
640 ost Prsituolution

and thir ombination

Host paasitevolution hst thn indntifis pa
Parasits an rt natural sltion on host populations, and vi vrsa' r- up natural sltionfoI
sulting in hangs in alll frqunisovr tim and hn volution. ara- host gnotyps for sl{
sits and lrosts an also rt riproal' similtanous natural sltion on om ommon' th}
ah othr, rsulting in lll frquny hangs in oth partnrs and hn typs, driving thm do.
ovolution. th rnost ommon gn(
T.hra svral ativ rsarh programs in th ara of prasit volution. frnt multilorrs paras
On of th most ativ aras is on th volution of prasit viruln. Th r- lation in oth host nd
sah program is nrainly ngagd in answring th qustion: !0hy do som g n r i d i v r s i t i n b o r
parasits mak thi hosts vry sik, whil othr parasits hav littl fft on now known as th R
thir hosts?Th onvntionalwisdom in this ara pior to th 1980s' spially oth host and parasit;
in th mdial on-lmunity' was that prasits should volv to inrasingly sam pla. (Th nam
..good.of.th- Ali in Lwis arroll'r
nign so as not to harm th host population. But this kind of
spis'' rsoning was ritd in tlr 1980s y thortil biologists.Thy tks all th running 1'
showd that parasits should in fat undr natural sltion to maimiz hypothsis urrntly s
thir own transmission to th nt host, without rgrd to th ffts on th m a i n t n a n o f g n t i
host population. In som ass' transmission is rrraimizdby pid rpliatin to b highly polymorpt
within th host, lading to high rats of host mortality (and/or moridity). t h a t a t t m P tt o p l a i r
Sulr an outom might xptd, for ampl, in parasits that a asily not mor ommonly r
movd twn hosts y vtors' suh as nrosquitos(.g.,malaria). Hn, basi ida her is that p
vn if th host is inapaitatd y disas, th inftious propaguls an as thy om loally
movd twn hosts y th vtors (or, similrly' y watr). Irr gnral' if dis- u a l p o p u l t i o ni n t h s
as transmission dos not quir a halthy, moil host, sltion may favor Parasits hav also
parasits that rprodu rapidly at th host's pns. rproduti<rnin som
Studis on th volution of parasit viruln hav important impliations parasit-mdiatdniltu
for human halth. For arnpl, mdial patis that inras th as of ag in populations tha
disas sprad not only inras th numr of inftd hosts, ut thy an asitsthan population
lso rsult in slion fr nror virlnt strains' Similarly, prasit stains is tht dlaying rprod
tht ar most rsistant to antiiotis will favord y sltion. If, for - dution of fwr offsp
ampl, antibiotis ar admistd in low doss that ar not suffiint to kill turing.
th ntir parasit population witin ah host, thn th antiioti rsistant Finally, parasitsh
strains will inras in frquny. Th volution of antiioti rsistanis a havior in ordr to nh
major prolm in human hlth and in th hlth of domstiatdanimals; it nw hosts. For nr
rPsntsa lassi as of rpid volutionary hang. d u h v i o r si n t h i
Parasits are also undr natural sltion to vad dttion and limina- atn y prdators.Bu
tion y thir hosts. onvsly, hosts ar undr natururlsltionto find and prdator is also anoth
kill thir parasits.utatiolrs that inrasth inftion Sussof parasits t h p r a s i t s u v i v s;
will favord y natural sltion,whil mutations that inras dttion sumption. Thus it wou
Sussof prasits y hosts will sinrilaly favord. This an lad to two ra1sltionto nhn
..arms ra,'' n- on of th hosts in th
typs of ovolutionary intrations.In on typ, alld an
fiil mutations in th parasit population swp to fixation; maning that
thy rpla th altrnativirlllsin the population, followd y similar su- BIBLIOGRAPH
sssfulswps of mutations for dfnsin th host population. Bll' G. !982. h st
In anothr gnral typ of ovolutionary intration, nfiialllls do Brkly:Univsit
not go to fixation. r hlst allls intrat with urlllsat othr host loi' w a l d ,P . W . \ 9 9 4 . u o

ulations,and vi vrsa' r-
and hn volution. Par-
lnous natural sltion ln
in oth partnrs and hn
] r o f p r a s i t v o l u t i o n .
lf parasitviruln. Th r.
th qustion: !hy do som
parasitshav littl fft on
prior to th 1980s' spially
uld volv to inrasingly
rt this kind of
..good-of-th-
thortial iologists. Thy
-rturalsltionto maximiz
. rgard to th ffts on th
rximizdy rapid rplition
rortality (and/or morbidity).
, in parasits that ar asily
ritos(.g.'malaria). Fln,
inftious propaguls an
, y watr). In gnral, if dis-
il host' sltion may favor
Ins.
h a v i m p o r t a n ti m p l i a t i o n s
s that inras th as of
inftd hosts, but thy an
s . S i m i l a r l y .p a r s i s t i n s
ord y sltion. If, for x-
hat ar not suffiint to kill
t h n t h a n t i i o t i s i s t n t
l o f a n t i i o t i r s i s t a n i s a
h of domstiatdanimals; it
lang.
: v d d r t i o na n d l i m i n -
natural sltion to find and
inftion sussof parsirs
rtionstht inras dttion
rvord. This an lad to rwo
:,alld an ..arms ra,'' n-
:p to fiation, maning that
ion, followd y similar su-
rost population.
ration' nfiialallls do
ith allls at othr host loi,
Hlst Prsite uolutio 641
..slf.''
and thir omination yilds a multilous gnotyp tht dfins Th
host thn indntifis parasits that do not math its ..slf'' gnotyp. This sts
up natural sltion for parasit gnotyps that an inft th most ommon
host gnotyps for slf. Howvr, on thse sussful parasit gnotyps
om ommon' thy impos sltion against th most ommon host gno-
typs, driving thm down in frquny. Thn a nw host gnotyp oms
th most ommon gnotyp' and th sltion on parasits hanges to a dif-
frnt multilous parasit gnotyp. In thory, this sts up a ontinous osil-
lation in oth host and parsit gnotyps' whih would funtion to maintain
gnti divrsity in oth th host and th parasit populatins. Th thory is
now known as th Rd Qun hypothsis, aus it would sm as though
oth host and parasit gnotyps ar running as fast as thy an to stay in th
sam pla. (Th nam oms from th famous rmark y th Rd Qun to
Ali in Lwis arrolr|'sThrogh the Looking G/ass: ..Now, ber, you se, it
taks all th running oxl an do, to kp in th sam pla.'') Th Rd Qun
hypothsis urrntly stands as on of th st-supportd thoris for th
maintnanof gnti divrsity t disas-rsistan loi, whih ar known
to b highly polymorphi. It also stands as on of th st-supportd thoris
that attmpt to plain why asual prodution of all-fmal offspring dos
not mor ommonly rpla sual rprodution in natural populations. Th
basi ida hr is that parasits would impos sltion against asual lones
as thy om loally ommon' and thus prvnt th limination of th s-
ual population in th short trm.
Parsits hav also n impliatd as a sltiv for in th timing of
rprodution in som animals. Th ida is that nimals would undr
parasit.mdiatd natural sltion to rprodu at a smallr siz and youngr
ag in populations tht ar sujt to a high risk of inftion y virulnt par-
asitsthan populations that ar t rlativly low risk of inftion. Th rason
is that dlaying rprodution in high.risk populations might rsult in th pro-
dution of fwr offspring' if, in fat' th host oms inftd for ma-
turing.
Finally' parasits have also n impliatd in th manipulation of host .
havir in ordr to nhan th transmission of thir inftiv propaguls to
nw hosts. For xampl, svral studis hav now shown that parasits in-
du haviors in thir hosts that inrs th liklihood that th host will
atn y prdators. But this is only known to our in parasits for whih th
prdator is also anothr host in th parasit's lif yl; nd it is known that
th parasit survivs and stalishsitslf in th nw host following on-
sumption. Thus it would ppar that som parasitshav rsponddto natu-
ral sltion to nhane thir rats of transmission y altring th havior of
on of th hosts in thir multihost lif yls.
BIBLIOGRAPHY
Bll, G. 1982. h strpiof Ntur: Tb uoltio d Gtisof Sulit.
Brkly:Univrsityof alifornia Prss.
wald' P. w. |994. uolutioof IftiosDiss.Oxford: Oford UnivrsiryPrss.
612 rd

Garrtt, L. 1994. Th omig Plgu:Ntul mrgigDisssi World ut of ontrovrsil, inspiin1

BI.Nw Yok:Farrar,Strausnd Giroux. lowd lnftiid:
Ns,R. -N4., Wh \WGt Sik.Nw York: Randlm
and G. . Willials.,|'994.
o d i t dw i t h G l n r l I '
ous.
Zimmr, . 2000. PrsitR: Isidth Bizrr \|y'orld
of Ntur'sNItlst t r s t di n d i l m m s
Dngerousrtures.Nw York: FPrss. -.L' dp into prhistoy. S
options that mothrs 1
m o t h r s . ' r o n g l . .
Hdy, SaahBlaffr (.1946)
group who assist a m
Sarah Blaffr rdy is on of Amri's most minnt anthropologists and a pardin 1999,l,ibrt
mmr of the National Aadry of Sins.Arr anthropologist and prima- Inrsinglv drar.nttr
tologist y training, sh works within th rod framwork of soiobiologi- Dahlm.!orkshop rp
al thory, to whih sh has mad signifint ontributions. Hrdy is most Hrdy's olks hv l
wll knowrr for alling attntion to th various volutionarY stratgisusd idas of infantiid w
by fmal primats, whih hs rought a radial shift away from primatolo- ogy ommunity, in 20
gists' traditional fous on mls as th prim soial tors. Mor rntly, rdy pologial Assoiation
has rvluatd th rol of mothrs and thir lationsip to thir hildrn. Al- otrtstanding ontri ut
though sh rlis havily on th ..otparativ mthod'' ntral to soioiol- Hrdy hld th positr
ogy, whn sh writs aout sx diffrnsrdy is an qul-opportunity riti Cliforni :rt Dvis fro
of oth fminists fo ignoring iology and of volutionary psylrologistsfor ig. nrrita, slr ontinus
noring soial histry. itorial oardsof uol
rdy was orn in 1946 to a walthy Southrn family. Slr graduatd from hr husnd liv ir.r
Radliff with a BA in 1969. It was in a Harvard anthropology ours that and sstainlagriu
sh rt hr futur husand, Danil Hrdy (a zh nm), whom sh mar'
BIBLIOGRAPHY
rid in 197\ e ltr ar a mdial dotor. Whil raring thir thr
hildrn, sh am inrasingly fousd on th nd to updat Attahmnt a t r , . s . ' l , . A h n t
and N. S.lhsr, 1
Thory with a viw to th volution of arly hominids as oorativ rd-
Prss.
rs. (Aording to this thory, dvlopd y Iohn Bowly, arly attahmnt
H r u s t r(,i . . l l d \ . H r .
to argivers is ruial for infants' normal soial and motional dvlop- uolutiorPrsp
ment.) H d y ,S a r a hB h f f r ' 1 9 7
Hrdy entrd graduat shool in nthropology in 1970, out th sam of Rprodutiolt. ()
tim that dward o. !ilson was ginning to writ Soiobiolog: Th Ne ditir-r rvithnw prt
198| [1999l.ll
Stbesis;s ompltd lrr PhD in 1975, te yar it was pulishd' Irvn
UnivsityPrss'( l9
DVlr's first fmal studnt, sh was havily inflund y Wilson and y 1 9 9 9 . o t h c rN
Robrt Trivrs. All thr suprvisd hr fildwork on th anuman langurs Nw York: Pnth
of India, pulishd as Th Lngurs rlf bu: Feml nd Ie Strtegies of W g.ttp://wrr w.
Reprodution ln 1977. Sh had initially undrtakn hr rsarh on infanti-
id y adult mals in langurs, living it was a patlrologial rspons to
Hutton, Jams(1
rowding. During hr first fild sason sh had to disard that hypothsis.In-
fantiidal havior was highly sltiv:mals targtd infants orn to fmals Jams Hutton, Sot
thy hd not matd with. Rgnizing tht fnralswr fr from pirssiv, th fath of gology
Hrdy gan to study fmal ountrstratgis. For xampl, y mating with nd was a mjor figur
multipl mals bfor thy usurpd ontrol of th toop, frialsstrov to rar1,of David Hum a
{orstallinfantiid latr l. ory of gology as lvll l
.Wom
In hr nt ook, The Tht Nuer uolud ( 198 1 ), Hrdy provoa. h argud-against th
tivly kpt fmals ntr stag and inorporatd th primat origins of p- bn formd y prr
triahy in hr analysis. Manwhil, th work on infantiid rmaind hot and tl-rat roks a
 uttott 64

lisess
i World ut of ontrovrsil, inspirirrg th first intrnational onfrn on th topi, fol-
lowd y lfticid: omprtiu d uolutior\' Perspctius (1984),
A. Nw York: Random
oditd with Glnn ausfatr. By thn, Hrdy had om inrasingly in-
of Ntur'sost trstdin dilmmas onfronting working mothrs thrrlughout history and
-.L. dp into prhistory. Sh st out to amin aross spis and ulturs th
options that mothrs hav -rployd,from sking th assistnof ..allo-
mothrs'' to nglt and ;rndonmnt. (Allomothrs ar mmrs of a
goup who assist a mothr in ring hr infant') rJ?hrrorr Ntur ap-
:lt nthr()pologistsnd a pard in1999, I-illrr Jlurl namd it on of tlr st ooks of th yar.
rtlrropolgistand prima- lnrasingly drawrr to th qustion of infant nds, irr 2005 sh oditd
m w o r ko f s o i o i o l o g i - Dhlm !orkshop rpot on attahmnt and ondirrg.
:riutions.Hrdy is most Hrdy's ooks hav n translatd into svral lnguags.Although hr
lutionary stratgisusd idas of irrfantiidwr lng rgardd with suspiion within th anthropol-
ift away from primatolo- ogy ommunity, in 202, at th annual mting of the Amrian Anthro-
ors.Mo rntly' Flrdy pologial Assoiation, other Nture was awardd th Howll Priz for
ship to thir hildrn. Al- outstanding ontriution ir-rBiologial Anthropology.
od'' ntrirl to soioiol. Hrdy hld th positiorr of Profssor of Anthropolog ilt th Univrsity of
r q u l - r l p p o t u n i tyr i t i aliforia at Davis from l984 to lrr arly rtirmntln I996' As Profssor
nr1,pshologistsfo ig_ mrit, sir ontinus loirrgrsrh,writing ooks, ad svil-rgorr th d-
itorial boards of t,llutilr Athropolog anl ut Ntur. Hrdy and
nily. Sh graduatd from hr husand liv in lionia, whr thy ar involr,din hitat rstoration
'nthropology ours that and sustainalagritrltur.
n a m ) ,w h o m s h m a r -
BIBLloGRAPY
Vhil raring thir thr
d to updt Attahmnt artr'. s.' L. Ahnrt,K. . Grossrann,S. B. Hrdy, ' . Lam,S..W.Porgs,
ids as ooprativ rd- and N. Sahsr,ds.2006.ttbmtd Bodig.ambridg,MA: IT
Prss.
iowly, arly ttahmnt
Hausfatr,G., and S. Hrd' ds.[1984]208.Iftiide: tplt'tttiue
d
and motionir1dvlop_ ulutior Prspti us, Nrv Brunswik'N.|: Aldin Transatiol-l.
dy, SaralrBlff.1977 L1980|.b Lltgttrstlf btt: Fnlld NIlStrtgies
1'970,about th sam lf Rprldtttilll. rrrridg, Prss.(1980,aprak
A: Hrvrd Univsit1.
Soiobiolog: Th Ntu d i t i o nw i t h r l r rp' r i ; r . )
r it was pulishd. Irvn 1981 [1999].T Womht Nueruolud.arnidg, N{A:Harvard
.Wilson UnivrsityPrss.(l999, papraklitiln with nrvpr[')
nd y and by
1'999.l>tbrNdtut : Histlrof otrs,Iftsd Nturl SItio.
rn th anuman langurs Nw York: Pantlrrr.
d le Strtegies of \h ag. h ttp://www'itrna.m/srhhrdy.htm. -u.s.
t h r r s ho n i n f a n t i -
pathologial rSpons t)
sardthat hypothsis. ln-
Hutton, Jams(|726-1797)
d infnts bon to fmals JrrrsHrrttln, a Sottish-orrr]otr and gntlmn firrtnr' is onsidrd
s wr f from pssiv. th fathr of gology irr Britain. H livd ruh of his latr lif in dinurgh
xampl,y mating with and was a major figur in th so-alldSottislrrrlightnmnt.A ontmpo-
trop, fmals strov to rar of David Hum nd Adam Smith, h hampiond h th Plutonist th-
ory of gologyas wll s th mthodologyof uniformirrianism.As a Plutonist,
i (198\), Hrdy provoa- h argud-against th so-alld Nptunists, who thor"rghtthat th world had
h prinrat origins of pa- n formd y pripittiln from Wat-that th ntr of th arth is vry
on infantiid rmaind hot and that roks ar formd y moltn lava pushirrg up to th surfa,
644 Hutto

whr it solidifis. At th sam tim' as a uniformitarian, h argud tht th half of th nintnth

way to undrstand gology is y ssuming that auss today ar th sam as Spis to trn suh ida
thos of ystrday,nd tht givn nough tim' gologial formations an
lLloGRAPHY
ratd natuall.
Hutton pulishd ths idas irr a massiv two-volum work titld Theor uvir,G. 1813. ssln
!. Blakwood.
of the rth (1795I, a tratiswhos lngth was ddonly y its osu-
H u t t o n 'J . 1 7 s s .T h r yo f
rity. Fortunatiy a followr, Jol.rnPlyfair, pulishd a mo radily assi-
th omposition,disso
bl vrsion in 1802, and Hutton's gologial position am rognizdand Trstiosof the R
stablishd. Howvr, i ws opposd y th nror popular thorizing of | 7 9 4 , I u s t i
Gorgs uvir (181)' who supposd that thr a many onvulsions in of Rsl,|rom Ss
arth histoy, and it was not until 180 that Hutton's uniformitrianism T. adll.
|795 . bor o|.t|
found a major hampion against th Frnhmn,satastrophism.T'his am .!illim
rh.
in th rhr-volum Priiples of Golog y hrls Lyll' probly th L l l .C ' l 8 3 0 - l 8 3 j . r l l l
work that was th gratstinflunon th young harls Darwin. Formr hgsi th'
Fronr th viwpoint of volutionary thinking, Hutton's importane lis in prtio. vols.Lol
th wy in whih h tndd a[th history. famusly spok of th wold Plafair,J. 1802.]Ilustrt
.!7illiam
rh.
s showing ..no vstig of a ginning' no prosPt of an nd'' (1788, 304),
t 8 0 5 .B i o g r p h i
and to Plafair laimd that on looking at stata' ..th mind smdto grow rltsctiosof th R'
giddy by looking so far into th ayss of tim.'' (Playfair 1805) oviously, Rphk,J. 200. h ,
tlris kind of thinking ws going against ilial histry rd litlly. Al- th rth,stiqit
though Hutton was ausd of athism, mor proaly h was (like many Rudwik,N4.J.S. 2005.B
lat'ightenth-nturythinkrs) mor of a dist, on who ss th glory of hizrgoPrss.
God in his urrrokrrlaws.
Although historially insignifiant in th snsthat this lif h hld was
Huley,Julian S. (1
not an influlon ltr thinkers, it is proal rhat FIutton shard with oth-
rs of his day (notaly rasmus Darwin, th grandfathrof Charls) n inli- Jlian Sorll uly w
nation to son] kind of trilnsformism in th ogani world. Indd,thr r Britain's intlligntsi.l
Passagsin his writing that sound almost lik an antiipation of natural s- t h 1 9 3 0 s a n d 1 9 4 0 s .l
ltin' H wrt ln is Priipls of Koldg that ..if an organisd ody th puliat\on o| t'ol
is not in th situation and irumstansst adaptd to its sustnnand tratd attntiorr onto I
propagation, thn' in oniving n indfinit varity mong tlr individuls ly promotd volutin
of that spis,w must assurd,that, on th on hand' thos whih d- of spiation, whil al
part most from the bst adaptd onstitlltion, will th most lial to pr- Huxly's ontriutions
ish, whil' on th othr hand' thos organisd odis, whih most approah ftious nthusiasm a1
to th bst onstitution for th prsntirumstans,will bst adaptd to sttrdonptsor i
ontinu' in presving thmslvsand multiplying th individuals of tlrir Born in Lndon, Hu
ra'' (Hutton 1794, )' l g ,o x f o r d ( 1 9 0 6 ) 'w
larly muh inflund y th work that h and otlrrswr doing on an- (zoology).H spnt tht
imal and plant rding in th advanmnt of ttr agriultur, Hutton ook, Th ldiuidul il
notd that dogs suviv thanks to ..swiftnessof f<rtand quikrrssoi sight'' sintifi aorrnt of (
..the most rtie y th Frnh
and hn dftivin rspt of thos nssaryqualitis, would
b th most sujt to prislr,and that thos who mpllydthm in gratst fondnss for th vitalis
perftion . . ' would thos who would rmain, to prsrv thmslvs, sin).In th ours
and to ontinu th r'' (utton 7794). Howvr, on should not rad too tlr tim twn th B
..arms
muh into suh passags,austhy wr almost olnmonple in th fist of a iologial

mitarian.he argud that th
ausstoday ar th san-las
qologialformations an
l-volunr work titld Thor
; ddonly y its osu-
i s h d m o r r a d i l y a
ition arl rognizd and
mor ppular thorizir-rg of
:rar lnny onvuIsionsin
H u t r o n ' s u n if o m i t r i n i s m
r's atastrophism.This am
harls Lyll, proly th
rg h:rrlsDarwin.
Hutton's importan lis in
famously spok of th world
) to f a n d , ' ( 1 7 8 8 , 3 0 4 ) ,
l, ..thmind smdto grow
, (Plyfir
1805) oviusly,
al history rad litrally. Al-
proaly h was (like many
t, on who ss th glory of
that this lif h hld was
hatHr.rttonshard with oth_
ndfathrof harls) an inli-
ani world. Indd, there ar
Lnantiipatin of natural s-
..if
e tht an organisd ody
daptd to its sustnanand
arity among th individuals
on hand, thos whih d-
zill b tlr most lial to pr-
'odis,whih nrost appah
rns,will b st adaptd to
ying th individals of thir
rnd othswr doing on an-
'f ttr agriultur, utton
foot and quiknss of sight''
s nssaryqualitis' would
,o mploydthm in gratSt
ain, to prsv thmslvs,
|vr'on slrould not rad too
.ostommonpla in th first
Hll', Juli S. 64.'
half of th nintnth ntury. It took harles Darwin in tbe origi of
Spiesto turn suh idas into full tlrory of volution.
|BLIoGRAPHY
uvir,G. 1813. ss\'o th Theoro|.th rth.R. Krr' trans.dinurgh:
.W.
Blakwood.
Huttoll'J. 1788.Thoryof th arth;Or an invstigation of th laws osrval
in
s s i -
th omposition'dissolution,and rstorationof lnd upon th glo.
T'rstios of th Rol Soitof dibrgb 1, no. 2: 209_304.
|794. luestigtiooi th Priiplsof Kou,ldgd of t|lProgrss
of Reso,from Sesto Sid Philosophy.dinurgh:A. Strlranand
T. Cadll.
1795. hor of th rth, ith Proofs d ]llstrtiozs.dinurgh:
willim rh.
Lyll, . 1830_1833.Priiplsof Golog:Big n Atte?ttrl plin tb
Formr hgesi th rth's Surf,lb Rfrto ussNott''ilt
()prtion.3vols.London:John urray.
Playfair'J. \802. Illustrtionsof th Huttoi Thor of th rth. dinurgh:
\filliam rh.
1805.Biographialaountof th latDr. JamsLluttonF. R. S. din.
.]-rstios
of th RrlISoitof diburgh 5, no. 3: 39_99.
Rphk,J. 2003. Th Vlho Ftlud Time:Jnls uttld th Disourof
the rth,sAtiquit'London:Simonnd Shustr.
Rudwik,M.J.S. 25.Brstigthe Limits of Time.Chiago:Univrsityof
hiagPrss. -.R.
Huxly,Julians. (1887_|975)
Julian Sorll Huxly Was a iologist, a popular writr, and a mmr of
Britain's intlligntsia.In volutionary studis,h promotd synthsisduring
th 1930s and 1940s. H ontriutd to th rvival of sltion thory with
th puliation of uolutio: Th odez Snthsis (1942) and h onn-
tratd attntion onto harls Darwin s th sujt'sthortialntr.Hu-
ly promotd volutionary taonoly, whih linkd lassifiation to th study
of spiation, whil also sking ojtiv mthods for taxononri work.
Huly's ontriutions of nw knowldg wr far lss important than his in-
ftious rrthusiasmand nourgmnt'as wll as his iliy to orrrin
sattrdonpts or idas into gnral prinipls and maningful visions.
Born in London, Huxly attndd ton Collg (1900), thn Balliol ol.
1g,oxford (1906), whr h rivd a first-lass dgr in natural sin
(zoology). H spnt th nxt yar at th Napls Zoo\ogla| Station. His first
ook,Tl Idiuidul i the niml Kigdom (7912), Was an attmpt to giv
a sintifiaount of volution inspird y th rntly pulislrd uolutio
retie by th Frnh philosophr nri Brgson (Huly always rtaind a
fondnss for th vitalism of Brgson whil ralizing that it ould nvr tru
sin).In th ours of th ook, r'rsingas an xalpl th omptition at
the tinr twnthe British and Grman navis, Huly introdud th ida
of iologial ,.arms ras'' twn ompting lirrsof organisrns' an ida
646 Hl,.|uliS.

tht has n muh ndorsd y suh thinkrs as Rihard Dawkins (1986).In ian intrvntion in url1
1910, Huxly rivd his first taulty ppointmnr, irr zoology rtoxford. mmrs of th Zoolgia
Whii in Oford h undrtook his first majrr rsarh suss, study of soity. Thy ford his
ourtship rituals and sxual sltion in th grat rstd gr. H also d- Aftr sveral ontriuti
vlopd skill in primntal mbryology. onstrution, in 1'946 l
Huxly's arr irrludd many mployrs. In 1,9|2, aptd an offr Nations duational, Si
from th Ri Institut in Houston to dvlop a nw dpartmnt of iology, rol suitd uxly prf
.sfhn .W.ar
whih provd diffiult. World I rupted, h rturnd to ngland. l i t i a l a s p t so f t h d i r
During th war, Huxly srvd mainly in rmy intllign.In 1919, h r- gav UNSCO an agndz
turnd to Oxford's zoology dpartmntnd onntrtedhis attntion on d- vision h had advotdf
vlopmntal iology and physiologial gntis.In 1920, h rportd on th s l f f o r i t s f u t u r v o l u t l
aility of hormons to provok mtamorphosis in axolotls. This rivd W hav to tak rsponsih
onsidrabl prss attntion and srvd as zratlyst for his liflong involv- family planning Programs
mnt in th nrdi. uly also invstigatdth rgulation of rats of gn t i s i n d v l o p i n g r r n t r
xpression and prolms of rlativ growth' suh as allomtry. In |925, sions, spiallyrgarding
movd to King's ollg London' apting th position of profssor in zool- md a vision of progrs
ogy and laoratory dirtor. Two yars lar,uly rdud his position to Aftr laving UNS
an honoary on and lft King's to pursu a arr as a frlanwritr. His sintist-lrity.H wr
ontriutions to his sin ar summarizd in Problms of Reltiu Groruth Lysnkoism), travld, at
(Huly \932) and The lemets of perimtI mbrology (Huly and Progrss and ultual d
d Br 194). Ths arnd Huxly ltiorrto th Royal Soity in 1938. ampaigns (Huxly 1954
Huly's most important frlanprojt was a ollaoration with . G. vious work and noua
.slls
and G. . to survy iology and th history of volution, titld Th Si- rsarh again. uly di
e of I'if (1929). During th 1930s, Huxly ontriutd to nwspaprs' As a sintist, Huxly l-
magazins,radio, and film. took on book and ditorial proits,as wll as bhavior,dvlopmnt,
spakirrgtours (sfor mpl uly |94). Smir-rgly vrythinginvolv- howvr, Huxly norml
ing puli sinin Britain in th 190s inluded Huly in som way. H is fousd attntion on iol,
st known for his rol on th BB radio program ..Th Brains Trust'' during thory and xplanation.
\forld \/arIl. Huxly's aility to ass th mdia provd ssntillatr in and his intrsts shiftd il
his lif. w o r k k t o t h s p i n r
In 1935, Huxly rturnd to institutional mploymnt, oming srtry topis am lss impl
of th Zoologial Soity of London. This ntaild marragingth soity and gav ohrn and ml
.Whil
its zoologil gardns in Rgrrt'sPark and \il/hipsnad. srtary,u- ply fat gathring.
ly kpt up his puli ontriutions. H turnd his rsarh intrsrsto sys- Huly's prfrn fo
tmatis and volution. Huly was partiulrly intrstdin volution lading tndd for nonspialist
to adptation and to th formation of nw spis.Polytypi spisnd go. ( 1 9 6 ) ,a n d i n h i s f i l m P l
graphial varitis (suspis)wr of intrst as thy suggstd..volution a m p l st o i l l u s r r a t g
in ation,'' a phras Huly oftn rpatd. He oind th trm line for car- providd a as for th
atrs that show gradual hang ovr a gogaphigradint.FI lso nour- mating havior' flight,
agd sarh into natllral nd sxual sltiorr'Dr"ringthis piod, uxly sud admiraly in pr<
organizd he N Sstemtis (1940) irnd wrot uolutio: The odr mation and th trati
Sthsis (1942). a supr xampl. This r
.!Var
Th outrak of world II in urop radially hngd Huly's om- stad, th ook's aom
mitmnts. H wrot xtnsivlyagainst fasism and against Nazi raist iol- ordination ot informt
ogy, nd h strssd th importan of intrnationalism and dmoray mriad fornls of slti
(Huly 1941,194). Aftr 1939, Huly ampaignd vigorously for Amr- Britain for a gnration

s R i h a r dD a w k i n s ( 1 9 8 6 ) .I n
tmnt' in zoology at Oxford.
rsrhsuss' a study of
at rstdgr. H also d-
1912, h aptd an offr
l nw dpartmnt of iology,
td, h rrurnd to ngland.
l intllign . In 1919, h r-
tntratdhis attntion on d.
.In 1'920,h rportd on th
;is in aolotls. This rivd
talyst for his liflong involv-
r rgulation of rates of gn
lh as allomtry. In 1925,he
position of profssor in zol.
I u x I y d u dh i s p o s i t i o n t o
ras a frlanwritr. His
Problms of Rltiue Groth
tl mbrology (Huly and
th Royal Soity in 1938.
as a ollaoration with H. G.
:y of volution' titld Th Sci-
y ontributd to nwspaprs'
d ditorialprojts,as wll as
Smingly vrything involv-
d Huly in som way. is
..Th
tm Brains Trust'' during
d i p r o v d s s n t i |I t
loymnt, oming srtry
ld managingth soity and
.!hil
lsnad. srtary, u-
l his rsarhintrsts to sys-
intrstdin volution lading
.s.Polytypi spisand go-
as thy suggstd ..volution
:oindth term lin for har-
'hi gradint.H also nour-
L.During this priod, uxly
rote uolution: Tb oder
ially hangdHuly's om-
and againstNazi raist iol-
:nationalism and dmoray
paigndvigorously for Amr-
Hle' Juli S. 647
ian intrvntion in urop. With so muh of his attntion lswhr, som
mmrsof th Zoologial Soity of London flt Huxly had aandnd th
soity.Thy ford his rsignationtn \942.
Aftr svral ontriutions to th war ffort and to planning postwar r-
onstrution, in 1946 Huly am first ditor-gnral of th Unitd
Nations duational, Sintifi, and ultural Organisation (UNSO). This
rol suitd Huxley prftly, although h frquntly fild to manag th po-
litial aspts of th dirtorship. He stayd only two yars in offi. Huly
gav UNSCO an agnda basd on sintifi humanism, xtnding a road
vision he had advoatd for many yars, that humanity an rly onl upon it-
slf for its futur volution (Huly 1948). Our dstiny is in our own hands;
w hv to tak rsponsiility for nsuring progrss. Huxly ampaignd for
family planning programs' intrntional onsrvation of natur, and univrsi.
tis in dvloping ountris. H lso usd sin to lvrag politial disus-
sions, spially rgarding ra and inequality. If nothing ls' Huly trid t
md a vision of progrss that was basd on dlirat human ation.
Aftr laving UNSO in 1948, Huly livd as a frlan writr and
sintist-lrity. H Wrot xtnsivly (inluding influntial work against
Lysnkoism), travld, and onsultd for govrnmnt and many othr groups.
Progrss and ltural dvlopmnt wr two main thms of his ongoing
ampaigns (Huly 1'954a,1'954, 1959, 1964). Whil h rvisd som pr.
vious work and nouragd many iologists, h did not undrtak original
rsarhagain. Huly did in 1975,
As a sintist,Huly had spialistprtisin svralsujts,spially
bhavior, dvlopmnt, nd volutionary iology. Throughout his arr,
howvr, Huxly normally prsntdhimslf as a ..gnal''iologist. This
fousd attntion on iologial prosss and prinipls, and gav priority to
.Whil
thory and planation. his arr shiftd from on pla to anothr
i n and his intrsts shiftd from spialty to spialty, Huxley always linkd his
work bak to th spin of ..gnral'' iology. Th partiulars of individul
topis am lss important; th undrlying gnral prinipls wr wht
gav ohrnand maning to partiulrs. Without thm, iology was sim-
ply fat gathering.
uxly's prfrn for genral prinipls is st illstratd in work h in-
tndd for nonspialistaudins.In ooks |lke Ats (1930) or t the Zoo
(1'96),and in his film Priute Lif of tbe Gnts (t94), h usd partiular
ampls to illustrat gnral iologial phnomna. Gannts, for instan,
providd a as for th study of dvlopmnt' parntal ar' adaptations,
mating havior, flight, ology, and so on. This approah hlpd Huxly
sud admiraly in projets rquiring th nylopdi olltion of infor-
mation and th xtration of ky thms. uoltio: he oder Sthesis is
a supr ampl. This rportd no original rsarhof Huly's making. In-
stad, th ook's aomplishmnt-and a landmark on-was Huxly's o-
ordination of information into an ovall vision aout volution and th
myriad forms of sltion in ntur. It hlpd shap volutionary studis in
Britain for a gnration.
648 Hule' lli S.
Hul,v, J. S., and G. d B
I n h i s p o l i t i s , H u x l y w a s a p o g r s s i v a n d h u m a r r i s t( H u x l y 1 9 2 7 \ .
m r i d g : a l r i d g
livd hunranity was rsPonsil for shaping its own futur, nd h Rus, . 1996. od tl
turnd to sien and rational planning to solv soial prolms. Th Tn. amridg' MA: arl
.Watrs,
nss Vally Authority's ltrifiation projt, modrnist arhittur' . K., and A. vn l
and uran planning wr among his favorit xampls. H frvntly sup- of Si. oustln: F
!7lls, H. G.' J. s' uly,
portd Population and onomi Planning' a soial poliy group. Huxly
Amlgirmtd rss.
w a s n o t a u g n i i s ti n i t s m m o n m e a n i n g ( i . . ' a t i v l y r m o v i n g u n d -
s i r a l s ) ,b u t h s u p p o r t d p o s i t i v i n n t i v st o i m p r o v d s i r a b l q u a l i -
tis as wll as birth ontrol for population managmnt. At th sam Huly, Thomas H
tim, h loid for improvmnts to puli halth and poor living ondi-
tions. Huxly argud strongly against th us of r as a sintifi or po- Thoms Hnry uly
litial onpt and ittrly opposd fursism. supportd osting aid to ogy duato' and popt
dvloping ountris' spially aid toward improving infrastrutur) du- Bulldog,'' onals mor
ation' and agriultur. l\4r'rhof his agnda in UNSCO supportd ths sintifi profssion and
aims. rival pulpits.
| 1919, Huly marrid Mari Julitt Baillot (|895_7994). Thy had Traind as a sur]
(1841'-1842), and at
two sons. Lik his grandfathr Thoms Hnry Huly' Julian uly's lif
was puntuatd y priods of svrmarri dpssion,whih somtimsd- o n I I S R t t l s k t
ilittd him or ld to major transitions in his lif (suh s laving admia). 1846_1850. His papr
wrot dtaild and informativ mmoirs and is now th sujt of in- m l l - o t - W a r )w, hih st
rasing intrst :rusof th wid sop of his thinking (l/atrsand van tion mmrans (tod
Hldn 1992; Rus 1996), Royal dal in 1852.
Th jolsstyro dnot
lLloGRAPY tbe Nturl Histor'of ,
Dawkins'R. 1986.Th Blind Wthmkr. Nw York: Norton. life upward. harls Da:
Huly, J' S. 1912.,Th ldiuidul i th iml Kigdom, amridg:amridg Down Hous in 1856 t<
UnivrsityPrss. onvrtd. Now th lt
1927. Rligiltt,ithoutRultil.London: rnstBnn. H u I y i m m d i t l a r
19.Ats. London:Jonthanap.
pushd farthr thn D
|93I. Wbt Dre l hik! h hIlngL'lfldrSieceto Htll1
tild Blif. Nw York: Harpr and Brothrs. from aps taonomiall1
\932, Problemslf RltiuGrlluth.London: tlrun. g o r i l l a a n a t o m y .L t u r
1934. lf I WerDittor. Nw York: Hrpr nd Brothrs. humns had animal an
196.t th Zto.London:Alln and Unwin. fumd th vanglial \,
1940. The Nu.,S1'51n*ofis. Oford: larndonPrss.
ws playing to nw audi
|941. Stadslo.Nw York: rpr.
.he tion wr familir in str
7942.uoltion: odr S'thesis.London: Alln and Unwin.
1943. TV: duenturi Plig,London: SintifiBok lu. image of nol-futu.fr
1948. UNso: ItsPurposd ItsPhilosrlph. Washington, D: Puli glian aristoray, ut th
Affairs Prss. Th puliation of Dl
1'954a.h volutionary pross.In J. Huly,A. . Hrdy, and . B. Ford, nw sin profssio
eds.,uolutio s d Prlss'1_23.L<lndon:Alln nd Unwin.
mnkind .rndwhlt es
1954.Sintilihumanism,volltion, nd lrunladstiny.Unpulishd
lturgivnir-rLlsngls,Otor16. HP 70.9,uxly Paprs,Ri dplod wht h alld
Univrsity,Houston. ism.'' His Manihan pt
1959.Introdution. In P. Tilhardd hardin,Th Phelml of ' today h is ralld for
11-28. London:ollins. Assoiation for th Ad'
1964. sssof Humnisl.London:hattond Win]us.
had plad on Vitorii
|97.mori,s. Lorrdon:Alln nlUnwin.
197.aories -l1.London:Alln and Unwin. inquiring whtlrth a

a humanist (Huxly 1'927).
ping its own futur' and h
soial prolms. Th Tn.
t, modrnist arhittur,
:ampls.H frvntly sup-
;oial poliy group. Huxly
.., ativly rmoving und-
to improv dsiral quali.
managmnt. At th sam
a l t ha n d p o o r l i v i n g o n d i _
of ra s a sintifi o po-
' supprtd oosting aid to
proving infrastrutur, du-
i UNSO supportd thes
l l o t ( 1 8 9 5 _ 1 9 9 4 ) .T h y h a d
Huly, Julian Huxly's lif
ssion,whih somtims d-
:(suhas laving aadmia).
nd is now th sujt of in-
ris thinking (.!flatrsand van
:k: Nortor-r.
rgdom. anlridg: amidg
rnst Bnn.
( odr S'iento I{unllt
trs.
-r:Mthun'
: nd Brothrs.
l.
'don Prss.
r.
ndon: Alln and Unwin.
: Sintifi Book lu.
;py. \/ashington,D: Publi
v, A. . ardy, and . B. Ford,
n and Unwin.
human dstiny.Unpulishd
).9, Huxly Paprs, Ri
tn,Th Phutoto of ,
o andWindus.
1n.
Hul, Tblms Hr 649
Huly' J. S.' and G' d Br.1934',helemtslf primtlmbrolog.
amridg:CmridgUnivrsityPrss.
Rus,. 1996.od to : The optlf Progrssi uolutiottrBioklg.
ambridg,\4A: Harvard UnivrsityPrss.
Watrs,. K., and A. vn ldn,ds. l992. lli ul1,:Bilklqistt! Sttesm
of Si'Houston:Ri UnivrsityPrss.
Wlls,H. G.' J. s. Huly,and G. P. Wlls.1929.h Siof Lif. London:
-l..
Amalgamatd Prss.
Huxly,Thomas Hnry (1825-1895)
Thomas Hnry Huly was an invrtratanatomist' plontologist, iol-
ogy duator, and popularizr of volution. Huxly's niknam, ..Darwin's
Bulldog,'' onals mor than it rvals:h ws a puliist for th mrging
sintifiprofssion and usd Darwinism's naturalisti idology to undrut
ival pulpits.
Traind as a surgon at an anatomy shool, Sydnham ollg
(184\_1842), and at Charing ross Hospita| (1842-1845), uly saild
on MS Rttlestthe to Australia's Grat Barrir Rf during th yars
1846-1850. His pirprs on th siphonophors (similar to th Portugus
nran-of-war),whih stalishdtlrat all olntratsomprisd two founda-
tion memrans (todrm and ndodrm), won him th Royal Soiety's
Royal dal in 1852.
Th jolss tyro dnound Robrt hams's transmutatio.ist Vstigesof
the Ntrl Histlr of rtil (1854) for its mnnikin-lik divin laws pulling
lif upward. harls Darwin's natulism, howvr' was appirling. alld to
Down Hous in i856 to air his ojtions to volution, Huly m away
onvrtd. Now th lturr in natural history ar London's Shool of ins,
Huly immdiatl ariaturd sllprnatural ..ration'' in lass. Publily h
pushd farthr than Dzrrwin:wh Rihard own in 1857 distndhumans
from aps taonomially' Huly in 1858 dw parallls twn humn and
gorilla anatomy. Lturing to working mn in 1859, Huxly said outright that
humans had animl anstors-th ..vilstand astlistparadox vr vntd;''
fumd th vanglialr'Witesson haring Huly in dinbugh. But Huly
ws playing to nw audins;pistraft-pikingassrtionsof human volu-
tion wr familiar in strt prints. A slf-profssd..plian,''lr onjurd an
imag of nol-futur-from-loworn-ginningsthat mant nothing to th An-
glian aristoray' but th frthinking mhanis lppd it up.
Th puliation of Darwin's the rigi li Spies (1859) rraldth
.Whn
nw sin profssionl to usurp old thologial qustions:
marrkind and what asis morality? Against th onsrvativlrgy, Huxly
dployd wht h alld Darwin's ..Whitworth gun in th armoury of liral.
ism.'' His anihan portrayal of sinvrsus thology mad nws. vn
tody h is ralld for upraiding Bishp Samul \/ilrfot th British
.Wilrfor
Assoiation fo th Advanmnt of Sin mting in l860.
had playd on Vitorian snsibilitisaout th santity of womanhood y
inquiring whthr th aps wr on Huly,s grandfathr's or grandmothr's
6''0 Hle, Tblms nr

sid. In rfrring an ap anstor ovr a ishop who prostitutd his talnts' fid to lad industrial so
Huxly, th Puritn, oozd moral sority to sham th worldly Nilrfor. him. As Dissntrs (out
Suh onfrontations only xaratd th Vitorians' ..risis of faith.'' In taking Oford and ar
1861, uxly denid own's laim that aps lak th human's third rral tion'' lgitimizing thir g
lo' whos avity has a postrior horn with a projting ..hippoampus Huly mad th wor.
minor.'' hrls Kingsly tnTh'Wter Bbis laughd aout ..hippopotamus a naturalisti maning, .
majors,'' and th affray rought human volution squarly to th puli's at- spiritulist A1frd Russl
tntion. Huxly mad his nwly aquired Nturl Histor Ruieta an anti. Huly oind th word
Own organ and st th trnd with his artil ..On th Zoo\ogtca| Rlations to ..agnosti'' aout
of an with the Lowr Animals'' (1861). Huly's wrking-lass talks, ou- with lean hands to tr
pld with a ltur on fossil humans (fousing on th rntly unarthd N- aS th nutral profssion
andrthal Man). formd th ornrston of his uidece s to 's Plc i his lst yars rguing h
Nture (1863). shaping a ntury of vo
uxly studid Dvonian fishswith lo fins and similar-looking amphi. Although volution w
ians, alld lyrinthodonts, from th strata surrounding oal sams.Rathr form, Darwinism's wal
than draw volutionary onntions' h mphasizd that layrinthodonts ltion had raisd manki
had prsistd unaltrd for immns priods. Initially, Huly livd that ..volution and this'' (
..moral indiffrn.''
th major groups had volvd for Silurian tims and had hangd littl t
sin. Only whn Darwin's Grman supportr' rnst Hakl, pulishd on tur within natur'' som
phylogny' or raial anstry' did Huly ronsidr. In 1'867,Huly rlas- Ultimatly, th supr
sifid birds gnealogially,and in 1868' h famously suggstdthat th small wht onrnd uly z
dinosaur ompsogtbus oti.d hav n anstral to irds (and may hav rd. Whras his Siet
orn fathrs). H dramatizd ths rlationships and promotd o. . tiv' his nin volums o]
Marsh's hors fossils from Nrask on his Amrian tour of 1876. on il- showing the Vitorian a1
lustration in Amri ddresss (1877), showing th inrasing lg and H. L. nkn alld H
tth siz from th small thr-tod on rohipps to th tiptoing who has vr livd.''
quus, was itd as th dfinitiv fossil vidn of volution for gnrations.
BILIoGRAPHY
Latr h dsrid th volution of th sondary palat in roodils.
.!hr Barr, A. ., ed. 1997.ho
Huxly's attitud toward Darwin's mhanism rmaind quivoal.
Darwin nvisagd variation ourring in minusul stps, Huly in 1859 eterssays.Ath
Dsmond,^. 1997.ul
piturd an innr for produing nw spis at on. His ntry on ..volu-
Rading,A: Addiso
tion'' in the cclopedi Britic (1878) mootd natural sltion only Di Grgorio,' 1984.. I
on' and Huxly's admi work ypassd it ompltly. uxly rsrvd Y | U n i v r s i tPy r s s
his disussion for popular talks: hr natural sltion was liknd to Lyons, S. L. 1999. homs
Napolon's troops rtreating from Mosow and rossing the Brsina Rivr, NY: PromthusBk
..vryon Paradis,J. G., and G. ' !(
hding only himslf,'' th fittst alon rahing th far ank.
PrintonUnivrsityP
ostly, Darwin's althusian mhanis srvd uxly in omating soial- Y/hit,P. 2003. ThlmsHl
ism or dunking rationism.Also, h avoidd volution in lass aush ambridgUnivrsity
Was training sinshoolmastrsaftr th duation At (1870). stalish-
ing his first propr laoratory in South Knsington in1'871', h fashiond a
pratial ..biology'' asd on th ommon plant and animal ..Typs.''This in.
flund shool urriula until th mid-twntith ntury. But inulating a
disrt ..Typ'' systm lft littl room for volutionary disussions. opning
Birmingham's nw Sin ollg, Huxly ontrstd th ryling of th
lassis at th Anglian univrsitis with sin's primntal approah.
Th undrvalud sintifi rsarhr' h ontndd, would ttr quali-

p who prostitutd his talents,
.!ilrfor.
ham th worldly
/itorins'
..risis of faith.'' In
ak th human's third rral
..hippoampus
h a pojting
..lrippopotamus
laughdaout
ion squarlyto th puli's at-
Iurl Histor Ruitu an nri-
..on the Zoologial Rlations
<ly'sworking-lasstalks, ou-
on th rntly unarthd N-
; uide s to is Plce itl
rs and similar-looking amphib-
urroundirrgoal sams. Rathr
phasizd that lyrinthodonrs
Initially, uly livd that
1 tims and had hangd littl
r, nst Hkl, pulishd on
nsidr.\n 1867, Huxly rlas-
nously suggstdthat th small
strlro irds (nd may hav
onships and promotd o. C.
tmriantour of 1876. on il.
rowing th inrsing lg and
e hippus o th tipoing
of volution fo gnrations.
|arypalat in roodils.
.!hr
sm rmaindquivoal.
nusulstps, Huxly in 1859
..volu-
s at on. is ntry on
mootd natural sltion only
it ompltly.uxl rsved
rral sltion Was liknd to
nd rossing th Brsina River,
alon rahing th far bank.
d uxly in onrbating soial-
:d volution in lass us h
d u a t i o nA t ( 1 8 7 0 ) . s t a l i s h -
ington in \871,e fashiond a
..Typs.'' This in-
nt and animal
'ith ntuy. But inulating a
'lutionaydisussions.opning
ontrastd th ryling of th
n's primntal approh.
.tndd,would tt quali-
Htll, Thoms Hr )/
fid to ld industrial soity.This dlightdth stl magnatswho finnd
him. As Dissntrs (outsid th Anglian huh), thy wr ludd from
taking Oxford nd amidg dges and saw Huxley's ..nw Rforma-
tion'' lgitimizing thir growing ivi authority.
Huxly mad th word euoltloz fashionalaround 1870. Fo him it had
a natllrlisti mning, wlrras for th atholi St. Gorg ivart nd th
.slla
spiritualist Alfrd Russl it onvyd th fiat of a highr powr. n,
Huly oind th word gosis in 1869. Biologists, as ultural ladrs, had
to ..agnosti'' aout suprnatual issus. They had to appoah sin
with lan hands to trustd.Huxly hlpd mold th imag of th sintist
as th nutral profssional,nd of volution as dmystifidsin.H spnt
his last yars arguing his antimiraulous as in thologial ontrovsis,
shaping a ntury of volution-rationdat.
Althlugh volution was fin for dmolishing Anglian doors losed to r-
form, Drwinism's wak-to.tlr-wall r"rpshotprturbd Huxly. A rural s.
ltion had raisd mankind, ut it ould not plain our ar of th wk. In
..volution
and this', (189) h portrayd thial man rnouning natur's
..moral indiffrn.''
this wr fort-ndy volution' yt mankinl's ..na-
tur within natu'' somhorvworkd gainst th primitiv sltivfors.
Ultin-latly, th suprmay of sin' naturlism' and agnostiism wr
what onrnd Huly as a populist. This ws vidnt as his works wr gath.
rd. Whras his Siet moirs (fiv volums, 7898_|902) wr dsrip-
tiv, his nin volums of ssays (789-7894) wr sintillating pros works
showing th Vitorian ag aommodting itslf to volution. Rading thm,
H. L. nkn lld Huxly ..prhaps th gratst virtuoso of plain nglish
who has vr livd.''
BIBLIOGRAPHY
Barr, A. P., d' 1997. hr.lmsI7rull',sPl irt Scid Lttrs:
tenrssys.Athns:Univrsit,vof Gorgia Prss.
Dsmond,^. 1997. Hule: From Deuil's Disipl tl uolutio'sHig Prist'
Rading,A: Addison.IWsly.
Di Grgorio'N4. 1984. T. H, Hul's Pl i Nturl Sie.N'"vvn' T:
Y l U n i v r s i tPr 'r s s .
Lyons' S. L. |999. I'homsHr Hl: he uolutilof Sitisf.Amhst,
NY: PromthusBooks.
Paradis,J. G.' and G. . Willirs,ds.19ti9.uolutild thi<:s.Printrr,NJ:
PrintonUnivrsityPrss.
Whit,P. 20.Tboms Hul: kig the ,, of Sin'''amridg:
CambridgUnivrsityPrss. -A.D.
Industrial mlanism
Industrial mlanism rfrs t th volution of dark ody olors in animal
spis that liv in haitats blaknd y industrial soot. Of pproximatly
1 0 0 r p o r t d a m p l s 'm o s t a r i n s s p i s .a n d o f t h s t h e v a s t m j o r -
ity ar moths' Th most thoroughly doumntd ampl is th ppprd
mot|t, B i ston b etlr i.
Th ommon name dsristh apparan(or phnotyp) of th typial
adult, a pal moth ..ppprd'' with lak-and-whit sals (s figur).
Pal and dark (mlni)f
Mlani phnotyps' namd aronaria, ar ssntiallysolid blak. Intrm-
diats, or insularia, also our ut riv rlativly littl attntion in gnral
summaris. xtnsiv gneti analysis onfirms that th phenotyps ar d-
trmind y multipl allls at a singl gn lous. akgrounds; (2) dirt
Th mlani phnotyp Was first disovrd nar anhstr, ngland, in onto tr trunks; and (3
1848. By 1895' about 98' of th spimnsnar anhstr wr mlani; in pollutd and unpollr
this on rar phnotyp had sprad aross th industrial rgions of Grat tlwll's pionring pt
Britain. With only on gnration pr ar' th narly omplt rvrsal in th yrs. ritiisms a
.!7ll
phnotyp frquny was astonishingly rapid. away from industrial r- usd' th positions on tr
gions, mlani spimns rmind rar. moths wr rlasd,an<
British lpidoptrist J. \/.Tutt advand th following Darwinian xplana- inspird nin additiona
tion in 1896: Ppprd moths ar ativ at night and rst during daylight was a svn-yar study I
hours' hiding on th surfas of trs. ost instivorous irds hunt y day N4ajrusrlasdliving
and loat pry primarily y vision. oths that ar wll onald against thir own 'hiding plas
akgrounds upon whih thy stay motionlss ar mor likly to sap d. propotions to math l
ttion y irds than ar onspiuous moths. Tutt proposd that typial pp- inoulars h mad dir
prd moths in undisturd environmnts gain prottion from visual ppprd moths. By ol
prdators y thir rsmlanto lihns on tr ark. In manufaturing r- usd in ths prdatin
gions whr lihns hav n dstroyd y pollution and th surfas of virtually mathd th s
trs hav n laknd y soot, th typil phnotyps los thir amou- t h m l a n i p h n o t p i
flag and fall vitim to ird prdators' whras th mlani phnotyps in Th wight of vidn l
suh pollutd habitats sapdttion long nough to pass thir gnson to prd moth phnotyps
progny. y xprimnt.
Tutt's idas wr not tstd until th 1950s whn Brnard Kttlwll Baus th rfltar
(197) initiatd a srisof xprimntsto dtrminwhthr or not irds a- ngativly orrlatd w
tually at mlani and typial ppprd moths sltivly. His xprimnts tstal prdition is th
inludd thr main omponnts: (1) quantitativ rankings of onspiuous- s o o t y ' p o I I u t dr g i o n s
nss (to th human y) of typial and mlani phnotypsplad on various ws lrly mt y th
6.t2

of dark ody olors in animal
lustrial soot. of approimatly
s , n d o f t h s t h v a s t m a j o -
ntd ampl is th ppprd
(or phenotyp) of th typial
:-and-whitsals (s figur).
ssntillysolid blak. Intm-
rtivly lil attntion in gnral
ms that th phenotyps ar d-
cus.
d nar anhstr' ngland, in
nar Manhstr wr mlani;
th industrial rgions of Grat
th narly omplt rvrsal in
l. \7ll away from industrial r-
following Darwinian xplana-
night and rst during daylight
instivorous irds hunt by day
that ar wll onld against
ss ar mor likly to sap d-
Tutt proposd that typial pp-
; gain prottion from visual
t r a k . I n m a n u f a t u i n gr -
r pollution and th surfas of
rl phnotypslos thir amou-
r a sr h m l n i p h n o t y p s i n
noughto pass thir gns on to
950s whn Brnard Kttlewll
rmitrwhthr or not irds a-
h s s l t i v I yH. i s x r i m n t s
t a t i v a n k i n g so f o n s p i u o s -
i p h n o r y p p s ldn various
]ndustril elnism 65
Paland drk (mlni)foms of th ppprdmtlr,isto btulri.
akgror.rnds; (2) dirt obsrvationsf prdation y irds on moths plad
onto tr trunks; and (3) apturrats of markd moths rlasdonto trs
in pollutd and unpollutd woodlands. Th dsign and xution of Kt-
tlwll'spionring xprimntshav n sujtdto intns srutiny ovr
th yars. ritiisms about his mthods (inluding th dnsitisof th moths
usd' th positions on tres whr th moths w plad, th tim of day th
moths wr rlasd,and th us of laboratory-rardand wild-aught moths)
inspird nin additional primnts y othr rsarhrs.Th most rnt
Was a Svn-yarstudy y ihai ajrus, ompltd in 2007. Spifially,
Majrus rlasd living moths into arnas at dusk so thy ould dtrmin
thir owrr hiding plas on trs for dawn. H adjustd th phnotypi
proportions to math lol frqunisnd kpt th dr-rsitis low. Through
inoulars he mad dirt osrvatior-rsof nin diffrnt ird spis ating
ppprd moths. By omparing th survival rats of th moth phnotyps
usd in ths prdation pimnts'e alulatd sltionoffiintsthat
vitually mathd th sltion offiints drivd from th rt of dlin of
th mlani phnotyp in th loal population ovr th orsof his study.
Th wight of vidnnow supporting Tutt's hypothsis hat irds at pp-
prd nroth phnotypssltivlyis sustantialand rmains unontaditd
y primnt.
Baus th rfltan of light fronr th surfa of tr bark is strongly
ngativly orltd with tmosphri lvls of suspndd partils, th
tstalprdition is that rnlani phnotyps should mor ommon in
sooty' pollutd rgions than thy ar irr unpollurd rgions. This prdition
was lirrly mt by th frqur-rylf nrelani ppprd moths distriutd
6.'4 Industril list

among 30,000 spirnsfrom populations survyd at 83 loations ass dlin in mlanism in

Britain twn 1952 and 1970. in atmosphri polluti
h gographi distriution of mlanism in ppprd moths is mor tions from southwst
strongly orlatd with atmosphri lvls of sulfur dioxid (So,) than with th sam tim intrval
smok. SO,, as gas, is mor widly disprsdfrorn point sours than pirr- tions of So, thn th
tiulat mattr that tnds to sttlloall as soot. Although sltitlnma op- mnts wr takn at
rat loally' gn flow from th migration of individuals among populations Amrian and Britis
has ontrir.rtdto mor gradual lins in mlanism in ppprdmoths, a rl- monomorPhism for th,
ativly moil spis,than in mor sdntarymoth speis('g., Goodo- dd lvls of tmosl
tis [=gtron'optrl bidtt) in whih frquny distriutions of mlani orrlations alon do n
phnotypsar mor sharply sudividd ovr thir rngs.The powr of grr suggStommon aus
flow t osur gnti daptation to lo:rl onditions rsulting from sl- volution ls dfind.
tion was littl appritd y arly rsahrs,lvho ntrtaind various quny in populatin
forms of nonvisual sletionin attmptsto aount for appantanmalis in to obsrv vtr now) tl
lins. olular thniqus using nonoding DNA sqrrnshv now isly. of th four pri
n dviopd for ppprd nroths y Ilik Sahri and his assoiats(Daly flow, nd sltion-ol
.Whth
et aL.2004)' Thir analysis of 12 mirosatllitloi from populations aross tionirl hangs.
a 125-kilomtr transt from north.s7'alsto Lds, ngland, indiats high is th sol, proximal a
lvls of disprsal and gn flor,.This and futrwrk should hlp qr'rantify tion in th volr-rtionf
.
th rol of migration in th volution of mlanism in this spis. urrd in populations.
In 1956 Rritin initiatd th lan Air Ats to stalishso-alldsmoklss eprimnts aimd at i
zons in havily pollutd rgions. Following signifiant rdutitltrsin etmos- ouflag from prdators
phri pollution' mlani ppprd moths av dlind in fquny as th ln all ras of tiv
typial form rovrd.Indd, th dlin in mlanism in th iattr half of ions and intrprrt
th twentith ntury is far ttr doumrrdthan rh inrasin mlanism portan of gn flow,
in th lattr half of th nintnthntury. An unintrruptd nnual rord th bhavior and olt
18 kilomtrs wst of Livrpool gun in 1959 y yril lark (lark t al. an outstanding amp
1985) inluds narly 19,000 spimnsshowing a drop in th frquny of st on rord dum
mlanis fonr a high of 9" to low.5ol, y 2003 (Grzrnt2005). The most populations.
rnt national survy' takn in 1996 (Grant t al. l998) shorvs markd d-
lins in mlanism hav ourrd vrywhr in Britain. Th prditd or- B|LIoGRAPHY
rlzrtiontwn hangs in th lvls of pollution rrd in th frqunisof C l k , . . , G . S . a n
-rlaniphnotyps in ppprd motlr populations as n fimly sta- th pppdmrh.
lishd. o o k , L . . 2 0 0 3 . T h r
Bru Gr:rnt and Dnis Own (Grant t al. 1996) have dou.rntdparal- Qurtrl Ruiluo1
Dly, D., K' !altham, J.
ll vlutionary hangs in oth dirtilnsirr th Nortlr Amian ppprd
S a h r ]i .0 0 4 . i n
mot, Biston betulri ogtri. Mlani phnotyps in th suspis r- btu!ri)'olI
sult from allls at th sam gn lous (Grnt 2004|' ut th ris and sprad Grnt,B. S. 1999,F.int r
of mlanism in Amrin populi.rtionsstartd aout 50 yrs ltr thn irr 2004.Alllim
Britain. arly musum olltions do not inlud any mlrnispimns rdit95:97-10
prior to 7929 in th viinity of Dtroit, Mihign, ut y 1959 th frquny 2005.lndustria
[ d o i : 1 0 . 1 083/ n p g .
of mlanis had rahd 90%. lan ir lgisltion Was thn inzruguratdin
G r a n t 'B . S . ,A . D . o k
th Unitd Stats in 1963. Rords ior sotrthastrllihigirn shtlw tlrt at_ tmporalviationt
mosphri SO, and suspndd partils susquntlydlind signifiantly. in Amriand Brit
B 1994 th frquny of mlani ppprd moths hd falln to 20o, drop- Grant,B. S.,D. F. own
ping to 5" 2001 (Grant and Wismn 2002). Th signifinof a sh:lrp ppprdmothsin ,

trv)rdat 83 loiltions aross
in ppprd moths is mor
;ulfurdilide (So:) tlran with
from point sours than par-
lt. Although sltionmay op-
ldividuals mong populations
rism in pprdmoths' a rl-
moth spis(.g'' Goodo-
ny distriutions of mlani
rir rngs.Th porvr of gn
nditions rsulting fronr sl-
- ' r s ,w h o n t r t i n dv a i o u s
,unttr apparnt nomalis in
4 DNA squns hav now
hri and his ssoiats(Daly
, loi fron-rpopulations aross
-ds'ngland, indiats high
Lrwork shuld hlp quantify
sm in this spis.
l stablishso_lldsroklss
4nifiantrdutions in atmos.
: dlind in frquny as th
mlanism in th lattr half of
than th irrrsirr rlarrism
unintrruptdnnual rord
y yril lark (lark t al.
ng a drop in th frqueny of
2003 (Grant2005). Tlr most
t a|, 1998) shows markd d-
in Britain. h prditd or-
ion and in th fr1unisof
ttions lrs n firnrly sta-
996) hav doumntd paral-
h North Amrian pppred
. ' n O t y p isn t h s u s p
1004),ut th ris and sprad
abotrt 50 yars latef than in
lud irny mlani spimns
Ln,ut 1959 th frquny
tio was thn inauguratd in
strn ihigan show that at-
luntly dlind signifiantly.
'ths had flln tl 20", drop-
:).Th sigrrifianof a shap
Idustril elism 65.'
dlin in mlanism in Amri:rn ppprd moths oinidnt with rdutions
in atmosphri pollution is omplmntd y omparisons of So. onntra-
tions from southwestrn Virginia, whr mlnism nvr dd 57" ovr
th sam tim intrvl. Th Mihigan lotion rodd highr onntra-
tions of So, than th Virginia loation in 23 of th 25 yars that masur-
mntsWr takn at oth plas.
Amian and British ppprd moth populations ar now approhing
monororphism for thir rsptivtypial plrnotyps,orrlating with r-
dud lvls of atmosphri pollution on oth sids of th Atlanti. Vhil
orrlationsalon d<lnot stalishausal rlationshios.ommon orrlations
suggstommon auss.
volution is dfind at th oprational lvl as a hang in gn (alll) fr-
qutlyin a population ovr tim. Th hangs w hav osvd (and ontinu
to obsrv evn now) in ppprd moth populations mt that dfinition pr.
isly. of th four primary volutionary fors-mutation, gnti drift, gn
flow, and sltion-only strong sltion an ount for suh rapid, dir.
tional lrangs.Whtlrr o not diffrntial ird prdation of moth Phnotypes
is th sol, proximl agnt of sltion' th ruial vidne for ntur:rl sl-
tion in th volution of mlanism is sd on th gnti hangs that hav o-
urrd in populations. This vidn is indisputabl and wholly indpndnt of
primnts aimd at idntifying partiular olnponnts of fitnss (e'g', am-
ouflg from prdtors and physiologial diffrnsamong gnotyps).
In all aras of ativ rsarh' partiipants routinly hallng assump.
.lfhil
tions and intprtations. w ontinu to invstigtth rlativ im-
portan of gn flow, lihns, prdators, potntil nonvisual sletion' and
th havior and ology of this moth spis,industrial mlanism rmains
an outstanding ampl of natural sltion asd on th most massiv data
st on reord doumnting long-trm alll frquny hangs in natural
populations.
BIBLIOGRAPHY
lark,. A., G. S. irni, and G. Wynn.1985.vlutionin rvrs: lan ir nd
th pprdmorh.Billogil.|<trl of th LitlttSoit1'26:1'89_199.
ook, L. . 200. Tlr ris and iIl of the rbrtriform of th ppprdoth.
Qurtrl Ruituof Biolog 78 99417.
.lfaltham, .!atts,
Dly, D., K. J. ully, P. . A. Rosin, S. J. Kmp' nd I. J.
Sahri.2004. Trinulotidmirosatllitloi fo th ppprdmoth (Bistrl
i s -
htulri) . llulrcologNo/,s[doi:10, | | | |li.147|-8286.2004.00607l.x].
Grant,B. S. 1999.Fin runingthppprdmoth paradigm.uolutio53: 980-984.
2004. Allli mlanismin Amrin nd British ppprdmoths.lournl lf
Hrdit95:97_|02.
2005. Industrilmlanism.Ilpedi rlf Lif Sies,www.ls.nt
I d o i l:0 . 1 0 3 8 / n p g . l s . 0 05004l1.
Grar-rt,l}.S.,A. D. ook, D. F. Own, and . A' lark.1998.Gogrphid
tmporalvriationin th inidnof mlanismin ppprdmoth populations
in Amria nd Britain.Jourl of Hrdit 89: 465-471.
Grant,IJ.S.' D. F. Own' and ' A. lark. 1996.Para||e| risrrdfall of mlni
ppprdmothsin Amrind Britain.llurlof Hrditl 87: 351_357.
656 Isets

Grant' B. S., and L. L. \/isman' 2002. Rnt history of mlanism in Amrian Thr ar striking pat
ppprd moths. Jourl of redit 93: 86_90. sity. To appriat ths
Kttlwll, H. B. D. \973' The uolutio of lnism: he Stud of Rurrig
txt. Insts,along witl
Ncessit. Oford: larndon Prss.
th four major groups t
ajrus, . . N. 1998. lism: uolution i tio. Oford: oxford Univrsity
Prss. a r t h s g m n t d n i n
27. The ppprd moth: Th proof of Darwinian volution. http://www alld suphyla, ar th.
.gn.am.a.uk/Rsarh/ajrus/SwdnPpprdmoth207.ppt;http://www tivs), and th rusta
.gn.am.a.uk/Rsarh/M jrus/Swdnta|k220 87.pdf . many othrs). Th inst
Tutt, J. W. 1896. British oths. London: Gorg Routldg and Sons.
th suphylum thy ol
-B.S.G.
suphyla has n quit
varlet of hpods ar
Th haratristi fa
Insts
organizd into thr ma
It is diffiult to argu that volution has produd any singl group mor lgs mrging from th r
sptaular than insts. Thir numrs alon ar staggring:ovr 820,000 nd outr shll that is r
spis hav en dsrid, and stimats for th atual numr of spis od; and a varity of d
xd 2 million. jor groups of insts ar still ing disov ered.|n 202, sts hv no lungs, r
for xampl' an ntirly nw ordr of mantidlik insts was dsrid, and th body that opn at tt
hundrds of nw spis ar dsrid from th tropis vry yar. molting' shdding thrr
Th divrsity of insts is no lss staggring.Insts oupy vry haital panding into a nw skin
rgion on arth. Thy hav invadd vry haitat pt th open sa, and thy On of th striking pl
hav radiatd into narly vry onival fding nih. any spis uild th 31 tant ordrs ar(
shltrs nd nsts, ranging from th asings of sand and dbris onstrutd by an found in vry ol
addis fly larva to the intrit mound and tunnl systms of tropial trmit vry othr major anim;
olonis. Whil th majority of insts lad solitary livs, thr is a sptrum of fossils or in whih ord
soil organization that inluds family aggrgationsin atrpillars, small fam- springtail that is stima
ily groups of wood roahs, olonial nsting in wasps, and highly rfind, om- mian, aout 250 millior
pliatd soitisof silings that form ,wasp' ant, and trmit olonis. Th and fossils of thm ar '
ants alon inlud spis that ar savngrs,prdators, and gardnrs;som ordrs that had not ap[
ant spis tnd aphids for thir sugary srtions and othrs ptur ants of and mammals, whih at
othr spis and nslav thm as workrs in thir own olonis. mals in th fossil rord
Insts provid som of natur's most sptaular phnomna and display olaanths, Ltimeri s
som of natur'S most sophistiatd faturs.Th ilial plagu of lousts r- ognizabl mmbrs of th
flts a sptaular natural ourrn' th ruption and organizd movmnt this standrd' w might
of millions of individuals of a speis namd Shistocr grgri. Priodial Th only inst ordr
idas mrg in th forsts of astrn North Amria y th millions on 13- dityoptra, whos m
and 17-ear yls, only to mat' rprodu, and di within wks. Monarh Unlik dragonflis' th
uttrflis mak sasonal migrations aross hundrds of mils twn North C r o n i f r o u s ( a l u rt
Amria and Mio. Many insts appar to dfy iologial onvntions: othr insts,and disap1
thr ar spis that onsist only of fmls, and many spis an produ of th Plaodityoptr
many individuals, from dozns to hundrds, from a singl gg. Instshav so- normously sussful,
phistiatd snsory systms' sing in wavelngths that humns annot' suh sts from thos priods
as th ultraviolt, nd haring at frqunis tht humans annot. Insts ar disappardso suddnl
not asy to poison; thy hav th apaity to assimilat a varity f toxi mol- !hil narly all of th
uls, somtims raking thm into harmlss omponnts and somtims hav n four prids l
squstringthm as dfnsiv ompounds that dtr prdators. dramatially. In th first

.y of mlarrismin Amrian
m: Tb Stud of Rurrig
fio. oxford: Oford Univrsity
.wirrianvolutio.http://www
rdmoth2007.ppt; http://www
:0807.df.
lutldgad Sons.
-B.s.G.
lud any singl group mor
ar staggring:ovr 820,000
t h a r u ln u m r o f s p
till bing disovrd.|n 22,
ik instswas dsribd, and
r trpis vry yar.
Instsoupy vry haital
,t xpt th opn sa, and thy
Jing nih. any spis uild
;and and dris onstruted by
tnl sysrmsof tropial trmit
ay livs,thris a sptrum of
tions in atrpillrs' small fam-
asps,and highly rfind, om-
r,ant, and trmit olonis. Th
rdators,nd gardnrs; som
ons and othrs aptur ants of
ir own olonis.
rular phnomna and display
h bilial plagu of lousts r-
tion and oganizd movemnt
:histlr gr gri. Priodial
\mria y th millions on 13-
nd di within wks. onarh
rdrds of mils twn Nrth
d f y i o l o g i a l o n v n t i o n s :
ad mny spis an produ
,m a singl gg. Insts hav so-
l r h s t h . l th u m a n s a n n t ' s u h
lat humans annot. Insts ar
similat a vait of toxi mol-
;s omponnrsand somtims
i dtr prdators.
lnsets 657
Thr ar striking pttrns in th organization and history of inset divr.
s i 1 'T. o p p r i a t h s p a t t n s i. t i s i m p o r t a n t f i r s t o p I i n s t si n l n .
txt' Insts' along with rnyriapods (ntipedsnd nrillipeds),ar on of
t four major groups that ompris th arthropod phylum. Th arthropods
ar th sgmntd aninrals, and th othr major arthropod groups, oftn
alld suphyla, ar th triloits, th hlirats (spidrs and thir rl-
tivs),and th rustaans(a divrs group that iniuds ras, llstrs,and
many othrs).Th insts ar a distint lass' th Hxapoda (six lggd)'within
.Whil
t suphylum thy oupy with th myriapods. ah of th arthropod
suphylahas n quit sussfulin numr of speis,th prolifration and
varity of hxapods ar approahd only by th rustaans.
Th haratristi faturs of insets inlud having thir sgmntd odis
orgaIrizdinto tr majo divisions (had,thorax' admn);thr pairs of
lgs mrging irom th ntral division (th thorax); an xoskiton (a hard-
nd outr shll that is not lways so hard, as in atrpillas);a vntral nrv
i s ord; and a varity of distint morphologial nd physiologial faturs.In-
stshav no lungs, rathing instad through spherils,whih ar tus in
th ody that open at th ody surf.Lik all arthopods, instsgrow by
molting' shdding thir trnl skin and xosklton priodilly and -
panding into nw skin and xoskieton.
on of th striking pattrns in th divrsity of insts is that narly all f
th 1 tant ordrs ar old; individuals that rprsntmost of ths ordrs
an b found in vry old fossil dposits. This is quit diffrnt from narly
vry othr major animal group, in whih many ordrs ar known only from
fossils or in whih ordrs rang widly in ag. Th arlist inst fossil is a
springtail that is stimatdto e 96 million yas old. By th nd of th Pr-
mian, aout 250 million years ago' most of th knwn ordrs had appard
and fossils of thm are widspred from that point onward. Th only major
ordrs that had not apprd y th Prmin ar th toparasitsof irds
and mammals, whih appard nly aftr th apparnof irds and mam-
mals in th fossil rord. o alirat ths osrvations'rmmbr that th
olaanths,Ltimri spp.' arc alld ..living fossils'' bausthr ar r-
ognizalmmrs of this group psrvdas 380 million-yar.old fossils.Bv
this standard, w might all most inst ordrs living fossils.
Th only inst order tht has gon xtint was a goup alld th Palo-
ditylptra, whos mmrs rsmld drgnflis in thir apparan.
Unlik dragonflis, ths insts wr hrbivrous' rrging in th mid-
aronifrous (aout 30 million yrs go), not muh latr than man
othr insts,and disapparingat th nd of th Prmian. Th striking aspt
of th Palaodityopter'sdisppaanis that thy appr to hav n
normously sussful,ornprising aout 50% of th known spis of in.
stsfrom thos priods. Rsarhrshav no ida why s sussfula goup
disappardso suddnly.
.shil
narly all of th ordrs and many f th familis ar vry old' thr
hav bn four priods whn th divrsity of spis within familis inasd
dramatially. In th fist of thspriods, th rly aronifrous (abotrt380
658 Ists

million yrs ago), wingd insts appard and gan to multiply. ln th s- ago), nd th arlistnl
ond, toward th nd of th Prmian, th arlist holomtaolous insts ap- t y o p t r i d a 'w h i h p p
pard, and this group gan divrsifying soon aftrward. Holomtaolous sil plant lvs from th
insts hv a omplx lif yl of gg, larva, pupa, and adult; th larva and damag from hrivors
adult ar dramatially diffrnt in apparan and lifstyl, and thy rpr- t h m a j o r r d i a r i o n so f J
snt almost two distint organisms. Holomtaolous insts inlud th famil- major radiations of flow
ir groups of flis, uttrflis,tls' ants' nd lawings. In ontrast, th This jint radiation
insts w all hmimtaolous, whih appar in th fossil rord for plants volve dfnssa1
holomtaolous insts, hav a simplr lif yl in whih th organism that om thos dfnss,an
mrgsfrom th gg, alld nymph, rsmlsa simplr and smallr vrsion mrge. Th vidn f
of th dult. ommon hmimtalous insts inlud grasshopprs, rikts, al studis. Instsat a
gnuin ugs lik stinkugs, and roahs. Holomtabolous insts hav n vidul platstht hav
normously sussful,rprsnting aout 9Yo of isting inst spis. hrivors or highr o:
Th third priod of inst xpansion ourrd in th rtaous,frm 130 al to surviv, grow lar
to 65 million yars ago, whn aout half of th modrn familis appard fnss at as agnts of t
and inst spisdivrsity inrasddramtially. This pansion gan im- om dfnsive strutur
mdiatly aftr th initial radiation of flowring plants. Th fourth priod of As a plant spisrfin
pansion, whih som sholars onsidr as a ontinuation of th rtaous tion, only spifi grou
radiations, was in th Trtiary, th priod that gan 6 million yars ago. through volution. As al
Th ontinud radiation of insts with phytophagous hits, maning that a s p i so r g n u s o f p l
thy fd on plants in one Way or anothr' was rtainly a ontinuation of th that mploy diffrnt <
rtaous pansion. As witnss to this, aoot 9 5" of th rnt Lpidoptr- p l n t s t h t p r o d u n w
ans (buttrflis,moths, and skipprs),narly all of whih ar phytophagous' p o p u l a r i o n s .o f t n i n
mrgd in th Trtiary. But th Trtiry radiation also inludd th mr- slvs ttling with diff
gn of th diddly nonphytophagous ants, s, wasps' and highr tr- twn ths plant pop
mits with thir sophistiatd soial systms as wll as th ordrs of li, pross of spiationha
whih ar toparasitsof irds and mammls. ra.
wo tis ind th volutionary historis of insts and flowring plants to Through this pross
ah othr. First, th ovrwhlming majority of flowring plants ar polli. othr riproally towar
natd y insts, and inst pollination srvi hs promotd divrsifiation rsult is that most plant
and spiation. Instshav n ating polln for quit som tim. Polln is phytophagous insts ar
rognizal in th guts of som fossil insets in th Prmian' ovr 250 mil- soiations t broad lv
lion yars ag. Th striking hang that oinidd with th mrgn of lis ar found only on
flowring plants was th apparan of spializd faturs of th had and finr lvls. onarh
mouth that failitatd not th ating of polln ut the transport of polln as a n d p i p v i n s w a l l o w t a
th insts fd on ntr. Ths first appar in som flis in th arly rta- tolohi); in ah s
ous. By th lattr half of th rtaous' 90 million yars ago' most sp- narly all oth insts.
ializd pollination systmsknown tday had volvd and ar larly sn in Th study of insts
th fossil ord. And should on doubt that inst pollination has on. volutionary iology. Th
triutd to th suss of flowring plants, it is worth rmmring that rgardd as on of th ;
groups of plants pollinatd by animals hav divrsifid at twi th rat of natural sltion, and sl
thos groups that ar pollinatd y wind or watr. anon of arly xprim
Sond, phytophagous inst divrsity has followd th divrsifiation of th fruit fly (or, mor
flowring plants. To sur, many insts hav n fding on plants for Thoms Hunt Mogn
most of thir xistn.Fossil plant tissu shows vidnof ing pird y ism that has n usd i
mits and ollmolans sin th lat Dvonian (aout 400 million yars tion and spiation. Th

gan to multiply. In th s-
holomtaolous insts ap-
rfterwrd. Holomtaolous
pa, and adult; th larva and
rd lifstyl'and thy rpr-
rusinstsinlud th famil-
lawings.In ontrast' th
in th fossil rord for
in whih th organism that
s i m p l r n d s m l I rv r s i o n
lud grasshopprs, rikts,
taolous insts hav n
f i s t i n gi n s ts p i s .
n th rtaous,from 130
modrn familis appard
. h i s p a n s i o n g n i m _
llants. Th flurth priod of
tinuation of th rtaous
lgan6 million yars ago.
agous habits, maning that
.tainly a ontinuation of th
'o
o{ th rnt Lpidoptr-
lf whih ar phytophagous'
on lso inludd th mr-
s' Wasps' and highr tr-
wll as th ordrs of li,
lts and flowring plants to
flowring plants ar polli_
a s p o m o t dd i v r s i f i a t i o n
r q u i t s o m t i m . P o | | ni s
th Prmian, ovr 250 mil-
ld with th mrgn of
:d fatursof th had and
t th transport of polln as
'm flis in th arly rta-
rillion yars ago' most sp-
' l v d n d a r | a r l ys n i n
nst pollination has on-
s Worth rmmring that
:rsifid at twi th rate of
owd th divrsifiation of
n fding on plants for
vidn of ing pird y
. (aout 400 million yars
Ists 659
ag),and th arlistntirly hrivorous ord of instsWas th Palaodi-
tyoptrida,whih appard aout 330 million yars ago. Aout 83% of fos-
sil plant lavsfrom th Prmian (whih ndd 250 million yars ago) show
damag from hrivors. owvr, it was in th rtaous and Trtiary whn
th major radiations of phytophagous spisourrd' following losly th
majo radiations of flowring plants.
This joint radiation rflts a ovolutionary pross through whih
plants volv dfnssagainst inst feding, insts volv faturs to ovr.
om thos dfnss,and spializdassoiations of inst and plant groups
mrg.Th vidn for this pross hs om from primntal ologi-
al studis.Instsat as agnts of natural sltion on plant dfnss;indi-
vidual plants that hav dfnsiv faturs, Suh as dnsr hairs tht dtr
hrivors or highr onntrations of toxi hmials in thir tissus, ar
al to surviv, grow largr, and lav mor offspring. onvrsly, plant d-
fnssat s agnts of sltion on inst traits; individuals that an ovr-
om dfnsiv struturs or dtoxify toxi ompounds leav mor offspring.
As a plant spisrfinsits dfnssthrough ontinued sltionand volu-
tion, only spifi groups of insts ontinu to ovrom thos dfnss
through volution. As an inst spisrfinsa spializdability to ploit
spis or gnus of plants, it appars to los its aility to xploit othrs
that mploy diffrnt dfnsiv ompounds. As a rsult, populations of
plants that prdu nw typs of fftivdfnssdivrg from thir siling
populations, often ing fr for a whil of inst psts but finding thm-
slvsbattling with diffrnt insts thn thir siling populations. Hyrids
twnthsplant populations far mor poorly thn ithr parnt and th
pross of spiation has gun, drivn y a ontinual and unforgiving arms
ra.
Through this pross, phytophagous insts and plants hv drivn ah
othr riproally toward mutual spializationsand divrsifiation.Th nd
rsult is that most plants ar ploitd y spifigroups of insts and most
.Whil
phytophagous insts ar spializd on partiula plants. thr ar as-
soiations at road lvls-for xampl, larva of individual uttrfly fami-
lis ar found only on a fw plant familis-th ar many assoiations t
finr lvls. onarh uttrflis fed only on milkwds (gnus sclpis)
and pipvine swallowtails fed only on pipvins and snakroots (gnusrls-
tolohi); in ah as th plants hror highly toxi ompounds that dtr
n r l ya I l o t h r i n s t s .
Th study f insts has ontriutd normously to th dvlopmnt of
volutionary ilogy. Th mimiry of noious uttrflisy palatal ons was
rgardd as on f th grat hallngs for harls Drwin's hypothsis f
natural sltion, and studis of mimiry oupid a ntral position in th
anon of arly xprimntal work on natural seltion. Th domstiation of
th fruit fly (or, mor proprly, vingar f|), Drosophil melogster,
Thomas Hunt Morgan in th arly twntith ntury produd a modl organ-
ism that has n usd around th world for primntal studis of adapta-
tion and spiation. Th hallng of xplaining how th sphistiatd soial
] 11sts

strutur of nts' Wasps' and s volvd' with its haratristi of stril f-

ml works raising thir sistrs,inspird.s7.D. amilton to dvlop a formal

J
thory of kin sltion.Kin sltion now undrlis muh of our undrstanding
of soiality and many pattrns of havioral volution throughut th animal
world. Studis of individual vriations in moth and uttrfly wing pattrns y
. B. Ford and his ollagus Wr on of th foundations of ologial gn-
tis, th xprimntal study of natural sltion and volutionary hang out-
sid of th laoatory.
Mor rntly, many diffrnt inst groups hav n usd as modl sys.
tms for volutionary studis. Th flour tls, Tribolium spp.' hav n
Jalonski, David (
usd in pathraking studis of group sltion. Studis of Watr stridrs
David Jlonski is amon
hav illuminatd th onflit twn th ss ovr mating, and how that
gists who, hving ntr
onflit has shapd th physiology nd havior of th ss, ttr than any
othr systm studid to dat. Th dmslfly spisthat sgrgaty typ of that took pl in th lat
larg-salpttrns nd
lk (with or without fish) hav n usd to illuminat how rapidly a ]roup
nring palontologists
n divrsify in novl haitats and how adapting to a novl ologial ir-
S t v nS r a n l y .J a l o n s k
umstan oftn oms with th ost of losing th aility to prsist in th
tintions and thir rsr
original onditions.
has approahd this su
|LIoGRAPY sis, whih uss omputr
Brrnaum, . R. 1995.Bgsitt thSstm:lstsd Thir Imptrl um of data drawn from th f
ffirs. Rading,A: Addison-\/sly. i n t i o n si n t h g o l o g i ;
isnr,T. 200. For Lluof Ists.amridg,A: BlknpPrssof Hrv:rrd undrstanding of th mo.
UnivrsitvPrss. iodivrsity risis.
Grimaldi, D., and . S. ngl. 205. uolutio of tb Ists.Nw York:
o n o f J a l o n s k i .ps r i
mridg Urrivrsity Prss. - l I

prov a numbr of tho

dynamis y using a vrr
ontologists. His rntv
thoy that volution tr
rror. Sintists tnd tl s
dus th misladingim
thus mor volutionaril
nations and tintions
yars,Jalonski argudt
advantg.In a similar s
iodivrsity ovr th pa
For a numr of yars' s
gratr marin animal di
t h i s s a m p l i n g r t i f a ri s
fossil anstry of hundr
of sintists who showed
btwn 87" and 95,,"
firm mpirially what p
major plosion in iodi
lion yars ago.
d,with its haratristi of stril f-
j \. D. Hamilton to dvlop a formal
.undrlismuh of our undrstanding
oral volution throughout th animal
l moth and uttrfly wing pattrns y
lf th foundtionsof ologial gn-
:ltionand volutionaryhang out-
grouPs hv bn usd as modl sys-
r tls,ribolium spp., hav en
sltion.Studis of watr stridrs
h ss ovr mating, nd how that
lhavior of th ss,ttr than any
lfly spistht sgrgaty typ of
d to illuminat how rapidly a group
v adapting to novl ologial ir-
lf losing th aility to prsist in th
: ]ltstsd Thir lmpt l Hum of data drawn from th fossil rord. In addition to important analyss of x-
'idg,A: Blknap Prss of Hrvrd
ri of th Ists,Nw Yok:
-1.r.
J
Jalonski,David (. 1953)
David Jalonski is among th most prominnt of gnration f palontol-
gistswho, having n traind in th midst of th rnaissanin paloiology
that took pla in th lat 197s and arly 1980s, hav tnddth study of
larg-saipattrns and prosssof volution and tintion gun y pio-
nering palontologists suh as Stphn Jay Gould, David . Raup, and
StvnStanly.Jalonski,s partiular ara of study has n th dynamis of
tintions and thir rsulting impt on pattrns in volutionary history. H
has apprahd this sujtprimarily through statistial,quantitativanaly.
sis' whih uss omputr modling and simulation to pross larg quantitis
tintions in th gologi past' Jalonski's work has md ontriutions to our
undrstanding of th modrn dynamis of xtintion and to th urrnt glol
iodivrsityrisis.
On of Jalonski's primry omplishmntshas n to hallngor dis.
prov a numr of thortial assumptions out volution nd xtintion
dynamis y using a varity of quantitativtols not availalto arlir pal-
ontolgists. His rnt work, for xampl' has shown that op's rul-th
thory that volution tnds to favor largr ody sizs-is asd on sampling
rror. Sintiststnd to study largr, mor ..iting'' organisms,whih pro-
dus th mislading imprssion that lrg organisms ar mor ommon (and
thus mor volutionarily sussful)than smll ons. In a study f th origi-
nations and xtintions of 190 linags of moliusks ovr many millions of
yars,Jalonski argud that, on th ontrary, ody siz onfrs no partiular
advantag.In a similar study, h showd that stimatsof a major inrasin
iodivrsity ovr th past 50 to 100 million yars ar not an aggrtion.
For a numr of yars, sintistslivdthat th rnt fossil rord shows
gratr marin animal divrsity aus th rnt rord is ttr prsrvd;
this sampling artifat is known as ..th pull of th rnt.''By studying th
fossil anstry of hundrds of gnra of living ivalvs,Jalonski ld a group
of sintistswho showed that th rnt fossil rord is rmarkaly aurat-
twn87o and 95o over th past .50 million yrs. This work hlps on-
firm mpirially what palontologists hav susptd for som tim: that a
major plosion in iodivrsityfollowd th Palozoi ra, rughly 250 mil-
lion yars ago.
661
662 Jeki

Prhaps Jalonski's most signifiant rsarh involvd studying th dynam- rviw is oftn suppos
is and pattrns of tintions. In this ara, Jalonski has ontinud to tst of hrdity, suh as
volutionary hypothsswith quantitativ analysis. On qustion h has ad- om a plausil thor
drssdis whthr survivorship during mass xtintions is pditalor ran- rviw, othr sltionr
dom, and whthr th volutionary trnds that follow major tintions populations, not indivr
display a uniform pattrn. A rlatd study askd whthr th survivors of nism of hrdity.
major tintion vnts (for ampl, th mass tintion that killd th di. Jnkin was orn in I
nosaurs 65 million yars ago) r in fat sussfulvolutionry omptitors family movd to Italy
aftrward. H found a numr of possil routs gnra may follow aftr Gnoa in 1851. H t
suh an vnt, ranging from thriving pansion and divrsity to stgnarion dsign and mnufatur
and vntual xtintion. H onludd tht whil thr wr som pr- h i m s l u p a s o n s u l t
dital fators involvd, th paths appar to vary asd on rgional nvi- som with his frind \
ronmntal diffrns, an osrvation that supports th hypothsis of Jnkin wrot ports fo
randomnss. n's ommitt on lt
Surpisingly, Jalonski onfirmd thr was oftn littl orrlation b- was aptdas an as
twn tintion survival and posttintion volutionary suss.In many Jnkin had wid si
ass, formrly struggling gnra thrivd, whil dominant organisms Wr him to ommnt on D
..Dad lad Walking,'' dsrisgn-
wipd out. A third altrnativ,alld rviw, pulishd in 1
ra that wr not immdiatly killd off ut wr doomd to vntual d- arth must hav old
..rippls'' tintion oftn follow millions
lin, suggstingthat sondary of dud th amount of p
of yars aft mass tintion vnts. ovrall, ths studis and othr work H also rpatd th tr
hav stalishdJalonski at th forfront of fforts to us nw mthodolo- r s t r i t dw i t h i n d f i n
gis (and oftn th latst thnology) to answr many of th qustions sltion ould aum
out th auray and usfulnssof th fossil rord that hav vxd pl- variation was a distin
ontologists sin harls Drwin's day. In gnral, Jalonski's work sup- would happn to th n(
ports th onvition that th fossil rord is rlial and thus an usd to sumption that hrdit
infr th rlativ importn of diffrnt vlutinary mhanisms pattrn- parnts. H ntd that
ing th history of lif on arth. By ttr undrstanding th dynamis of x- onntratd within a
tintion' Jalonski has also highlightd th ways xtintions, whih r ft. This had happn
normally sn as ngativ prosss' ontriut to th ovrall dynamis of th farmr had rd or
volution. isti. In a wild popula
with unhangd mmb
|BLIoGRAPHY
halvd in ah gn
Jalonski,D. 1999.Th futurof th fossilrord.Siece284 21,14-2116. Darwin had nvr tl
2001. Lssonsfrom th past:volutilnary impatsof massxtintions.
think of thm as indrv
l,rrledigsrlf th Ntirlnl Acdm lf SiesUS 98: 5393_.5398.
2004. tintin: Pst and prsnt.Ntr 427: 589. -D.Se. Jnkin,s rviw, modif
prosss Wr at work
pointd out' howvr' t
diffrns within a p
Jnkin,Hnry CharlsFlming(1833_1885) favord individuals ar
Hnry harls Flming (pronound Flming) Jenkin made major ontri-
butions to physis nd ltrial nginring in mid-nintnth-nturyn- BIBLIOGRAPHY
gland. Rmmrd y volutionists as th author of a rviw of harls
Bowlr,P. 2003. uolut
Darwin's o th rigi of Spies' h agud tht natural sltion would aliforniaPrss.
not wlrk if hrdity Ws a pross that lndd prntal haratristis. This D w i n , . l l 8 5 9 ] l 8 " 2 .

involvd studying th dynam-
lonski has ontinud to tst
ysis. on qustion h has ad-
:intionsis prdital or fan-
lat follow major xtintions
kd whthr th survivors of
i xtintion that killd th di-
' s f u l v lu t i o n a r y o m p t i t o r s
uts gnra may follow after
' n n d d i v r s i r yt l s t a g n a t i o n
whil thr wr som pr_
vary basd on rgional nvi-
supPots th hypothsis of
ls oftn littl orrlation b-
:volutionrysuss.In many
il dminant organisms Wr
lad !7alking''' dsris gn-
wr doomd to vntal d.
(tintionoftn follow millions
thsstudis and othr work
ffortsto us nW mthodolo.
sW mJny of th qustions
l rord that hav vd pal-
nral,Jablonski's work sup-
lial and thus an b usd t
Jtiony mhanisms pattrn-
rstanding th dynamis of x_
ways xtintions, whih ar
lt to th ovrall dynamis of
Sci284:.21|4_2776.
impatsof massxtintions.
: e sU S 9 8 : 5 9 3 - 5 3 9 8 .
427:589.
3-188s)
g) Jnkin mad major onti-
n mid-nintnth-nturyn-
Luthorof a rviw of hals
l that natural sltion would
d parntalharatristis.This
Jnkin 66
rviw is oftn supposd to hav shown that th mov to a partiulat modl
of hrdity' suh as ndlirr gntis'was ssntilif sltion was to -
om a plausil thory. In fat' although Drwin was imprssd y Jnkin's
rviw, othr sltionists ralizd that if variation Ws sn as a proprty of
populations, not individuals, sltion was plausil whatvr th mha-
nism of hrdity.
Jnkin was orn in Dungrnss,Knt, ngland, on arh 25, 1833. His
family movd to Italy and Jnkin rivd an MA from th Univrsity of
Gnoa in l85]. rturnd to ngland nd spnt 10 yars working on th
dsign and manufatur of sumarirr tlgraph als, aftr whih h st
himslf up s a onsulting nginr in London. H took out numrous patnts'
.sillianr
som with his frind Thomson (latr Lord Klvin). In th 1860s,
Jnkin wot rports for th Bitish Assoiation for th Advanmnt of Si-
n'sOmmitt on ltrial Stndards, s a onsqun of whih th ohm
was aptdas an asolut standard of rsistan.
Jnkin had wid sintifi intrsts,nd th Nort Britis Reui, asked,
him to ommnt on Darwin,s thory fom a matlrmatial viwpoint. In his
rviw, pulishd tn 1867, Jnkin ndorsd homson's argumnt that th
arth must hav oold down too rapidly for natural sltion to hav pro-
dud th amount of progrssiv volution osrvd in th history of lif.
H also rpatd th traditional viw that within ah spies,variation is
rstritd within dfinit limits. or originally, h ana|yzed how natural
sltion ould umulat favoral variations. Arguing that ah suh
variatiorr Was a distint salation, or sport of natur, h xplored what
would happn to th nW hratr as rh individual rprodud, on th as-
sumptiol1 that hrdity blnds th haratristis of th mal nd fmal
parnts. H notd that in artifiial sltion, suh nw haratrs ould
onntratd within a small popultiorr and ould thus hav signifiant f-
ft. This hd hppnd in th as of th shot-lggdAnon shp, whr
th farmr had rd only from th singl ram orn with th nw haatr.
isti. In a wild population, lrowvr, th favord individuals would rd
with unhangd mmrs of th popultion, and th hatristi would
halvd in h gnration until its fft was too small to ount'
Darwin had nvr thought of variations s larg-salsports, ut h did
think of thm as individual haratr hangs. was thus imprssd y
-D.Se. Jnkin's rviw, modifying ltr ditions of his b<lokto mphsizthat othr
prosssWr t work. othr Darwinists, inluding Alfrd Russl !7alla,
pointd out, howvr, that if on thought of variation as a rang of haratr
diffrns within a populatiorr, Jnkin's argumnt was infftiv baus
favord individuals ar oitiful rathr than r.
BIBLIoGRAPY
Bowlr,P'200 . t , o l u t i o : h i s t o r lnf l d . 3 r d d . B r k l y : U n i v r s i t y o f
California Prss.
Darwin,. [1859l 1872.o th rigi of Specis.6th d.London:John Murray.
664 Jnki

Hull, D. L', ed. 197. Dr,i d His ritis' amridg' A: rvard Univrsity
Prss.
Jnkin, F.1867. Th origin of spis.North British Ruie 46:277-3I8.
-h
Rus, . 1,999. Duii Rurlltio: Si Rd i Tllthd l.2nd d.
hiag: Univrsity of hiago Prss.
-h
Vorzimmr, P. J. 1970. hrles Dnui: Yrs of otrlurs. Philadlphia:
mpl Univrsity Prss. -P.J'B'

Kttlwll, H. B. f
Hnry Brnard David l
atd group oftn rfr
ntis' whih inludd s
S h p p a r d .W h i l r h r
markd y a ommn
populations, and, in pa
ural sltion as th m
was a giftd naturalist \
oyhood lov of ntor
s a r h i n . B . F o d . s
Kttlwll is st kn
mlanism, whih rfrs
in many moth spisin
of th fisr larg-salar
ngland and ontinnt
trnd ourrd in th p'
spiswas fist disov
50 yars it m so .
1 8 9 1 i s t h l s s i n i n
Kttlwll am fa
arly l950s doumnt
ommon. In pollutd a
s o o t - d a r k n dt r t r u I
mnt ondutd in th .
m k dp a l a n d d r k
tmptd to aptur as n
hind his invsrigti
for th two forms shot
Was at an advantag'o
p th rapturrt
vivd during th intrv
what Kttlwll found.
unpollutd wood in Do
ln this wood h ws a|
t I w l ls u p p l m n t d tI
amridg,NtA; arvard Univrsity

,itish Ruieu' 46: 277-3 18.

ee Rd i Ttlrltbd !.2nd d.

lrs of otror:ersy.Plriladihi:
-P.l.B.

Kttlwll,H.B. D. (1907_1979)
nry Brnard David Kttlwll ws a founding mmr of a loosly assoi-
atd goup oftn rfrd to s . B. Ford's Oxford Shooi of ologial G.
ntis,whih inludd suh luminris as A. J. ain, yril lark, and Philip
Shppard. !7hil thr was nvr any formal strutur' th mmrs wr
markd y a onrmon appoah to the study of gnti variatin in natural
populirtions,and, in partiular, y a ommitmnt to th prvasivrol of nat-
ural sltionas th most important agnt of volutionary hang. Kttlwll
was a giftd nturalist who lft mdial prati at th ag of 41 to pursu his
oyhood lov of ntomology. H gan working full tim as a snio r-
srhrin ' B. Ford's nwly foundd suunit of gntisat oxford in 1951'
Kttlwll is st known for his rsarh on th phnomnon of industrial
mlanism, whih rfrs to a rapid inras in th frquny of rar drk forms
in mny moth spisin th viinity of mantrfaturingntrs as a osqn
of th first larg-salair pollution assoiatdwith th Industrial Rvolution in
ngland nd Continntal urop. Th first and most famous ampl of this
trnd ourrd in th ppprd mot|7,isto betulri' Th dark trm in this
spiswas fist disovrd nar Manhstr in 1848. .$ithinth sp of only
50 yars it am so ommon that th pal frm was loally xtint. (Tutt
1891 is th lassi nintnth-nturydisussion of th topi.)
Kttlwll am famous f a sris of fild primnts ondutd in th
arly 1950s doumnting th rason why th dark forrn was oming mof
omlnon. In pollutd aras, dk oloration prottd th moths, rsting on
soot-darknd tr trunks, against ird prdation. In n lgant fild xpri-
mnt ondutd in th su1mrof 195, Klwll rlasdlarg nrrmrsof
markd pal and dark moths into a pollutd wood nar Birmingham and at-
tmptdto aptur as many as possil ovr th nt svral nights. Th logi
hind his invstigationwas that, all things ing qual' th raptur rats
for th two forms should th sam. If' on th othr hand, th dark form
was at an advirntag'owing to its rlativ inonspttlousnss'on would -
pt th raptur rt for it to highr aus mor of th drk form sur-
vivd during th intrvl twn rlas and rapture. And this is prisly
what Kttlwll found. Kttlwll ondud ompanion xprimnt in an
unpollutdwood in Dorst in th summr of 19S5 whr h found th rvrs.
In this wood h was a|to raptur mlr of th pal form thn th dark. Kt.
tlwll supplmntd ths primnts with a film rod doumnting that

66.t
666 Kimur

whn irds wr givn a hoi, thy passd ovr inonspiuous moths in DNA. DNA onsistsl
favor of th moth that was mor onspiuous in its akground rsting sit. sin, guanin, and thy
(S Kttlwll 1955 and 1956 for th original rports and Kttlwe|\ 197 of a fw thousand as
for a summation and ovrall disussion.) qun of ass in tha
Whil ttooks oftn portray Kttlwll's invstigations as unprolmati' th DNA has no knov
rsarhers (inluding Kttlwll himslf) rogniz that th initial invstiga- A mutation is a prma
tions involvd numrous assumptions tht ar opn o qusrion (.g.,th rSting to a T. If th hang l
sit of moths). This has ld thos ritial of volutionary thory to laim that th fft on th individua
phnomnon is no longr an xampl of natural sltion, and furthr that osrval fft. If a l
thre is a onspiray y ttbook writrs to hid this fat. (Hoopr 2002 js a frquny in th popuI
.Wlls
snsationalisti aount of th work y a journalist and 2000 is a svr' nti drift.
rligiously motivatd ritiqu by a iologist who is a followr of th Rvrnd ln t968 Kimura put
Sun Myung Moon, foundr of th Unifiation hurh.) At last ight studis di- ulk of hangs in th
rtly addrss privd problms in Kttlwll's initial invstigation.Th stud- random drift. Kimura
is hav indpndntly onfirmd that ird prdation is th most important th population and ulti
fator in th sprad of th dark form. Prhaps th most onvining vidn rsult of natural sl
that it is an ampl of naturl sltion' howver, oms from th prditd ssnof Kimur's id
rapid dlin of th dark form sin th introdution of ..lan Air'' lgislation ory of volution; Kimu
in th Unitd Kingdom and lswhr. (Majrus 1998 givs a full ovrviw of ovrd indpndntly
modrn rsarhand Rudg 2002,2005 addrssth lims of frud.) most of th susqu
narily attriutd to hin
|BLIoGRAPY
It oftn happns in t
Hoopr, J.2002. f oths d n: Th Untold Stlrlf Sicd th Ppprd its own sak' turns ou
of. Nw York: Norton. stantial ody of thor
Kttlwll,. B. D. 1955.Sltionprimnts on industrialmlanismir-rth
was fortunat that th
lpidoptra.rdit 9: 23_342.
1956.Furthrsltionprimnts on industrilmlanismin th nturis latr, ut wi
lpidoptra.Ilredit 10: 287-301. at studnt, Kimura s
1973. Th uolutilof lism,Oford: larndon. d o m p r o s s s ,s u h
Majrus,M. . N. |998. elism:uolutili Atio.Oford:Oford Univrsity inras in th popula
Prss. gnrations it would t
Rudg,D. !. 2002. rypti dsignson th ppprdmoth. Itrtiol lourl of individuals that woul
Tropil Biolog d oserutio(Ruistd Bklklgi Tropil).50,no. 1: 1-7.
2005. Did Kttlwllommit fraud? R-xaminingth idn'Pbli Kimura l'vorkd out t,
Udrstndig lfSi14, no. 3:249-268. ial, or dltrious.Su
Tutt, J. !. 1891. lism d elohrlismi British Lpidlptr.London: in gologial tims.
Swan Sonnnshin. It turns out tht th
.!lis'
J. 2.los of uolutio:Sior th?.Washington,D: Rgnry. for anothr is simply
-D.vR.
rasonaly wll know
an hkd agains
squnsof this dis
Kimura, otoo (|924_994) w know th mutatiol
Motoo Kimura is st known for th ida that most volution t th DNA tion. For studis of
lvl ours y random prosses'rthr than y Darwinin ntural sl- lok, whih depnds'
tion. Prior to this, volutionists had attriutd ssntiallyall of volution to It is now gnrallyt
natural sltion. Kimura argud tht for most volutionary hangs' han mals and othr vrt
is mor important. is still ontrovrsial, h
Within a ll, a hartristinumr of hromosoms an found (for It is lar that in som
xampl, 46 in humans). h hromosom is a long, tightly oild thrad of ltion. Th answrsv
 Kimur 667

ovr inonspiuous moths in D N A . D N A o n s i s t s f s q u n o f f o u r k i n d s o f a s s , d n i n , y t o -

i r l i t s a k g o u n dr s t i n g s i t . sirr,gunin. nd tlryrin,rviatd A, , G, and T' A gn is ir streth
l rprtsand Kttlwl'|197 of a fw thousand ass that is rsponsil for spifi funtion; th s-
qun of ass in tht rgion dtrmins th funtion. ost of th rst of
vstigationss unprolmati, th DNA has no known funtion and is oftn rfrrd to as ..junk DNA.''
; n i z t h t t h i n i i a l i l l v s t i g - A mutation is a prmannthang in th DNA; for ampl, a may hang
)nto tlstion(.B..th rsting to ;r T' If th hng ours within a ]n'it will oftn hv som visil
tionary thoyto lim that th fft n th irr.lividulinhritirrgit. hangs irr tlr ..junk'' usully hav nl
l sltion' and fr-rthrthat osrvalfft. If a partiular qn or other sqltnof DNA hangs its
l this fat. (Hoopr 202 is a frquny in th population y rlrdm prosss' this is alld random g-
Llistirnd Wlls 2000 is a svr, nti drift.
o is a followr of th Rvrnd ]n 1968 Kimura put forth th ida-vry daring at th tim-that th grat
lrurh.)At last iglrt studis di- ulk of hngs in th DNA ar pr-rlyndom, th rsult of mtltation :.rnd
; initial invstigtion.h stud_ random drift. Kimura argud tht whthr th nw mutation iIrrassin
dtion is th most important th population and ultimatly displsits prdssorDNA is usually not th
; th most onvining vidn rsult of natural sltion ut, muh mor oftn, simply han. This is th
vr' oms from th prditd ssnof Kimura's ida. This might alld th mutation-random drift th-
..lanAir'' lgislatin oy of volution; Kimura alld it h nutral thory' Tlr sam ida lvas dis-
tion of
s 1998 givs a fr"rllovviw of ovrd indprrdntlyy Jak King and Th<lmasJuks (1969), ut baus
s th laims of fraud') nrost of th susquntdvloprnntsar du to Kimura, th thory is ordi-
naily attriutd tl him.
It oftn happns in th history lf sin tht mathmatis, dvlopd for
itor1,of Sied th Ppprd its wn sak, turns out latr to hav pratial uss.]t turnd out that su-
stantil ody of thory lrad alrdy rl dvlopd lry Kimura himslf.
mlnisnrin th
r industrial
was fortunat that th nutral thoy found its ppliation, not dads or
nturis latr' ttt within his own liftim. Starting whn h was a gradu-
'ustrialmlanismin th
at studnt, Kimura solvd a numer of diffiult prolms involving ran-
: lndon. d o m p r o s s s ,s u h a s t h p r o a i l i t y t h a t m u t a t i o n w i l l u l t i m t l y
l . f r d :O f r d U n i v r s i r y
./ll,lo inrirsin th population and or-th prvailing form, th nr-rmrof
gnrations it would tk flr sr-th poss to our' and th numr of
d roth.Itrtiol lourl of individuals that wtluld afftd y th muttion during this pross.
' Billogiropil)50, no. 1' 1_7.
aminingte elde.Publi Kimura workd out th answrs' whthr th mutation is nutral, nfi-
ial, or dltrious.Sussfor th last is xtmlv rar, ut not ngligil
Britisb Lpi dopter. I'onrJon: in gologial tims.
It tuns ollt th:lt th rat of volutionary srrstittrtionof on nrrtrls
li \rVashington,
D: Rgny.
for anothr is sinrply qual to th mutation rt. Busmutation rats ar
-D.W:R.
rasonably wll known, this prrrritsa prdition of volutionary rats that
irn hkd against th histrial rord. On of th most important on-
squnsof this disovry is that it maks possil a ..molular lok.'' If
w know th muttion rat' W an dtrmin th at of molular volu-
t most volution at th DNA tin. For studis of volutiorr, th timsal is dtrmind v molular
'n y Darwinian ntural sl- lok, whih dpndson Kimura's nutral thly.
d ssntiallyll of volution to It is now gnrlly aptdthat th grt ulk of DNA hangs in mam-
it volutionaryhangs' han als and othr vtratsis nutal' following Kitnura's thory. Th thory
is still ontrovrsial, howvr' rgarding hangs in fr.rntionalrgions of DNA.
hromosomsn found (for It is lr that in sonl ss' han prdominats;in trtlrrs.it is ntul s-
i a long, tightly oild thrad of ltion. Th nsrvswill hv to om from as-by_asnalvsis.
668 KIn|r

otoo Kimur was orn in okzaki, Japan,on Novmr 1,7924.He

showd an early aptitud for mathmatis' ut his dpst intrst was
otany. H gratly njoyd a mirosop givn to hinr y his fthr. I-Invr
lost his intst in flowrs, and in his larr lif h was a sussfulorhid
rdr. H was in high shool during th arly days of .!7orld .War II' and in
L
1944 was admittd to th Kyoto lrnprial Univrsity. H was not invoivd in
military srvi, ut wartim lif in Japn was hard for vryon. Aftr th
war' h joind th lartoy ot Japn's prmir gntiist'Hitoshi Kihara.
ihar rognizd Kimura's talnts and prmittd him to spnd his tim
studying th litratur of mathratialpopulation gntis.
With th hlp of Amrin gntiists working for th Atomi Bomb asu- Lak, David Laml
alty ommission, Kimura Was al to otain sholarship to study in th
David Larnrt Lak v
Unitd Stats. Aftr a yar t lowa Sta Univrsity, h transfrrd to th Uni-
studis of Galfpagos f
vsity of Wisonsin and bam a studnt of Jams F. row. H had th op-
th wild. Born in Lo
portunity for daily ontat with Swall Wright, on of th foundrs of
shool in th Knt and
mathmtial popultion gntis.Aftr rivirrghis dotorat in 1956,he
prizs for natr.rallristo
rturnd to Japan and for th rst of his lif was mployd y th National
lg, amridg, in 19
lnstitut of Gntis in Mishima, Japan. H ontinud his ollaoration with
tion in his formal stud
row, and ah took vry opportuniry to spnd tim with th othr. To-
Cambridg ornitholo
gthr thy authrd a nurnr of paprs and a txtook. Kimura did of
srvations and onlus
amyotrophi ltral slrsis on Novnrr 13, 1994, on his 70tlr irthdy.
writtn a dozn papr
Although Kimura is st known for th nutral thory and th attndant
mbridgshir (I934
mathmatis, h was lso knowl-rfor pplying diffusion modls to othr pro-
Lak surprisdfrin
lms. Ths involvd tlr solution of partial diffrntial quations, at whih
living that fildwork
h was partiularly adpt. lnsidrd th ffts of random flutuattons
aptd a jo as iol
in th intnsity of naturl sltion,whih n lad to prosssthat mimi
Dvon. Lak was no
random gnti drift. H dvlod a modl of migration twn olonis,
dnts trap and and r
in whih migrants mov o on of th nighoing olonis. Altogthr, h
mak osrvationson
wrot six ooks and ovr 6.50 sintifi paprs' At th tim of his dath h
th asis for a four.1,
was Japan's st-known gntiistnd h had n th ripint of many in-
Suprfiially'Lak's
trntionl honors'
Llowvr' Lak writir
ILIoGRAPHY h apted an obsrv
row, J. F. 1997. otoo Kimura. Biogrphilmoirs of Felloso|-tb Rol rr sn on at lst s
Soit<l|. Lldo4: 255_265, thniqu of anding t
Kimura, M. 1983. he Nutrl Thor lf olulr uolutio.amridg: viduals, and it allowd
rridgUrrivrsiry Prss. behavior. Lak liev
King, J. L., nd T. H. Juks.1969.Non-Darwinianvolutiln:Rndomfiationof sintifi. latr ad
sltivlv nutrairuttions.Si|64: 788-798, ( 1 9 5 0 ) ' a o l l t i o no f
akahat,N., d. 1994. PopultioGtis'olulr uolutitld th Nutrl
Thlr:SltdPprs.hiago;Urrivrsityof hig Prss. -|.F'. this appoah to a sing
Julian uxly an
Lk visitd th Gal6p
study finhs. As had r
of prdators and <tm
trmly wak on th
with whom Lrk ons
Aadmy of Sins ;
L N o v m r1 3 , 1 9 2 4 . H e
his dpst intrst was
'im y his fathr. H nvr
was a sussful orhid
ys of !7orld !ar II, and in
ty. H was not involvd in
rd for vryon. Aftr th
gntiist,Hitoshi Kihara.
:d him to spnd his tim
l gntis.
or th Atomi Bom asu-
holarship to study in th
, h transfrrdto th Uni-
s F. row. H had th op_
, on of th foundrs of
; his dotorat in 1956, h
mploydy th National
rud his ollaoration with
j tim with th othr. To-
ttbook. Kimura did of
|94,on lris 70th irthdy.
Ll thory and th ttndant
-rsionmodls to othr pro-
rntial quations' at whih
: t so f r n d o m f l u t u a t i o n s
.ad to prossSthat mimi
n i g r a t i o n t w n o l o n i s .
ng olonis. Altogthr, h
At th tim of his dath h
n th ripintof many in-
irs of FIl'sof th Rol
i,uoluti-l
. amrid g:
lutio:Random fiation of
,8.
,r
uolutio tL th Nutrl
)hiagoPrss. -I.F..
L
Lak, David Lambrt (1910_|973)
David Lambrt Lak wrsa British ornithologist irnd ologist famous fo his
studisof Glipagos finlrs,n of th ky ampls of nturl sltionin
th wild. Born in London, Lak gan ird-wthing whil in oarding
shool in th Knt and Norfolk ountrysid. H ws additd, winning shool
prizs for natural history thr onsutiv yzrrs.H ntrd agdaln o1-
lg, amridg, in 7929,t::.'lik harls Darwin' Lak found littl inspira-
tion in his formal studis. Instad, h took to th fild with mmrs of th
amridg ornithologial lu. Lak also gan pulishing systmatio-
svationsand onlusions on bird havior. By graduation in 1933, h had
witt dozn paprs' and th lu sponsord him to writ Tb Birds of
m bridgesh ir (|9 4|.
Lak surprisd frinds and family y not pursuing an aadmi arr. B-
living that fildwork was unpopular in nrridg faultis, Lak instad
aptda jo as iology mastr at Dartington Hall' a progrssivshool in
Dvn. Lak was nouragdto us th fild as tahing spa. H had stu-
dnts trap and nd roins in th nary wlods, and h nouragdthm to
mak obsrvationson teritory and hvior. Tlrse studnt projts am
th irsisfor four-ya stlldy of roins.
Suprfiially,Lak's he Lif of the Robi (|943| was a typial lif history.
Howvr' Lak's writing hid volution in rnthod. mpirially mtiulous,
h aptd an osrvation as ral ..only whr th ation onrnd has
n sn on at last six oasions'' (Lak I94, 6). H also usd th nw
thniqu of anding irds. This prmittd long-trm studis of known indi-
viduals, and it allowd him to ask omplx qustions aout lif history and
havior.Lak livdthat suh mthods mad thology mor rigorous and
sintifi. H ltr ddd to his studis of roins wit Robin Redbrst
(1950),a olltion of rltd litrtur,musi, and tolk ultur. H rpatd
this approah to a singl ird with Sifts i Tor (1956).
Julin Huly am a mntor and pomotr. Undr his nourgmnt'
Lak visitd th Galipagos Islands (from Dmr 1938 to April 199) to
study finhs. As had many for him, Lak fist supposd that th asn
of prdators and omptitors for food mant that natural sltion was -
trmly wk on th islands. This was rinford y Amrian ollagus
with whom Lk orrsultdon th way hom to ngland. At th Califorrria
Aadmy of Sins and th Amian usum of Ntural istorv, L:rk
669
670 Lck
..gntidrift'' modls pro-
was shown how his osrvations fit wll into th adaptd to diffrnt hl
posd in th 1930s y Swll Wight and fashionalin Amri. A thnial vrgn) Lak suggs
monograph (Lak 1945, writtn in 194) dtaild his onlusions drawn tan; thy offrd a s
from this influn. wild. Inspird y this
Lk rvrsd his viws shortly aftr rturning to nglnd and to Huxly's (1944), and of th or
influn. ..Unptdly, a ronsidrtion of th original matrial ld to Dri's Fihs
markd hang in viwpoint rgarding omptition twn spis and th natural sltion into
ak diffrns twn th finhs'' (Lak 1947' 1).|n Dr,i's Fihs whn volutionists w
and many popular artils thraft, Lk mad two points aout natural ..artifiial'' (laorato
sltion in Gal6pagos finhs. prinipl of omptit
..
First, h onntd finh aks to dit, arguing tht thy wr wll Gaus in th 190s.
adaptd to spifi fding styls: larg sds, small sds, insts, and fruit nglish fforts to d
(s figur). Natural sltion within ah spis rinford spializatin . B. Ford nd rinfor
and ratd highly adaptd forms. Sond, Lak fousd on ass whr sv- land snail, p, a
ral spisoupid idntial ologis in th sam loation. In ths ass, Bisto. Although th
..Sin
h said, natural sltionwas spiallyintns: th han of thir - Rort Bowman and
ing qually wll adaptd is ngligil, on of thm should liminat th othr sity f omptition' i
ompltly'' (Lack 1947,62)' n small islands, Lak found only on spis undr way y Ptr
of ground finh, suggsting that omptition ld on spis to dominat and onlusions.
othrs to loal xtintion. on islands whr two or mor spiswr found, Io 1945, Lak b
Lak rlizd that ths spis W nvr found with atly th sam nithology' part of th
ologis.Natural sltionws ausing spisto highlight thir diffrns, institut had two go
thry rduing omptition. This ausd divrgn, as diffrnt spis data and ovrsaw n.
Trust for Ornitholog;
al ommunity. S
ornithology. Lak fo
lation of population I
ontrolld populatior
ators. H fousd t
took pla. Lak's th
..
dwards and thr
i Birds (1'966).Hts 1
sizd ologia1isola
Not a strng admi
in th dirtor's hait
too narrowly. Lak'r
his waknsssas ar
stdnts.
Lak onvrtd to
th Anglian Churh
uolutior hor1
and volution ould
han prosss's
Figur 1. ampls of Darwin's finhs.Not t vry diffrntak shaps, natural, not vil. v
adaptd to diffntfeeding stuffs-ig strong aks for hard nuts and th lik, squnt history. Th
fin aksfor smll sdsand th lik. an of th first h
 Lk 671

h ..gntidrift'' modls pro_ adaptd to diffrnt haitts. hanging dits was th usual avnu for this di-
lnalin Amria. A thnial vrgn' Lak suggstd.In this intrprtation, aks took on spial impor-
ltaild his onlusions drawn tan; thy offrd a sptaular, nuand ampl of natural sltion in th
wild. Inspird y this suss'Lak mad similar studis of passrine irds
rg to ngland and to Huxly's (1944), and of th ormorant and th shag (1945).
. th original matrial ld to a
Druin's Fibs mad Lak famous. H had transformd a as against
ition twnspis and th natural sltion int a prmir xampl for it. His rsrh am at a tim
947, vt). | Dri's Fibs ..natural'' rathr than
whn volutionists wr alling for mor studies of
d two points aout natural ..artifiial'' (laoratory) xampls of sltion.It did th sam for ology's
prinipl of omptitiv xlusion' mad thortially important y G. F.
arguing that thy wr wll ..Lak's finhs''' many ltr alld thm, also oostd
Gaus in th 1930s. as
small sds,insts' and fruit nglish fforts to dvlop volutionary olgy, hampiond y oxford's
is rinford spilization . B. Ford and rinford y Arthur ain and Philip Shppard'sstudisof th
:k fousdon asswhr sv- land snil, p, and Brnad Kttlwll's work on th ppprd moth'
sam loation. In ths ass, Biston, Although th finh as was ritiizd in th 1950s' spially y
..Sinth han of thir -
: Rort Bowman and Danil Simrloff, for ovrstating th tnt and intn-
rm should liminat th othr sity of omptition, it nvr lost its allur. By th 1970s, long-trm studis
;' Lak fundonIy on spis undr way y Ptr and Rosmary Grant onfirmd and tndd Lak,s
,d on spisto dominat and onlusions.
lo or mor spiswr found, In 1945, Lak am dirtor of th dward Gry Institut of Fild or-
found with xatly the sam nithology, part of the Dpartment of Zoologial Fild Studis at Oxford. Th
s to highlight thir diffrns' institut had two goals. First, it was a ntr for hang of ornithologial
ivgn.as diffnt spis data nd ovrsaw nationl nsus work in ollaoration with th British
Trust for Ornithology. This put Lak in a ntral position in th ornithologi-
al ommunity. Sond' th institut sponsord original rsarh in fild
ornithology. Lak fousd this rsarhon population ology and th rgu-
lation of population numrs (Lak 1954). H argud that only thr fators
ontrolld population sizs:disas,food availaility' and parasits or pred.
ators. H fousd th institut's rsarh on prisly how this rgulation
took pla. Lak's thoris w svrly ritiizd, spially by V. . Wynn-
..group
dwards and othr sltionists.''H plid wtth Plpultio Stdis
i Birds (1966). His final ook' ologil Isoltil i Birds (1971'),mpha-
sizd ologil isolation as th driving for of divrsity.
Not a strong administrator, Lak Was mor omfortl in th fild than
in th dirtor's hair. H was ritiizd for fusing th institut's attntion
too narrowly. Lak's strngths as n ornithologist, howvr, far dd
his waknsssas an dministrator. H was inspiring and mntord many
studnts.
.!Var
Lak onvrtd to hristianity aftr \World II. H was onfirmd in
th Anglian hurh in 1951, and h rmaind dvout throughout his lif. In
uolutionr Thor d hristi Belif (1957), Lak argud tht sin
and volution ould hlp to plain mny unrsolvd thms in rligion.
han prosss' suh as natural sltion, ould rativ. Dath was
vry diffrntak shaps, natural' not vil. volution did not spak to th origin of lif, only to its su-
..Th tru signifi-
ksfor hard nuts and th lik, squnt history. Th Bil and ntur did not ontradit:
an of th first haptr of Gnsis is to assrt that God mad th univrs
672 Lmrck

and all in it, that H saw that it was good, and that plad man in a sp-
Lamark initially ml
ial rlationship to Himslf'' (Lack 1957,34). Lak also sought to tterun-
te 177s with th ir
drstand aras of gnuin onflit. For instn'many asptsof man's moral
in Paris. His rr l
bing smdto him to unplaind y thorisaout th moral volution
was ronstitutd as th
of animals.
on of th profssors
Lak's ..y Lif as an Amatur ornithologist,, (797| is an autoiography
was givn instad th 1
to 1,945, writtn whn h am dirtor of th dward Gry Institut. For
..amtur''to ..profssional.'' mals.'' H soon hara
him this markd an important transition from
mals without vrtr
!. . Thorp (1974) provids th kind of knowldgal mmorial only a
initially to what h kn
los frind ould podu. Lak was awardd th Godman-Slvin Gold
am ovr th nt twr
dal of th British Ornithologists Union in 1958 and th Darwin Mdal of
zoology.
th Royal Soity i 1972.
lassifying th inv
BlLloGRAPHY thm wr not Lamar
onward, h had spu
Grant, P., and R. Grant. \986. olog d ullutilof Dri's Fihs. pls of physis, hmi
Pinton,NJ: PrintonUnivrsityPrss.
mbridg:amridgshirBird lu. think aout th histor
Lak' D. t934. The Birds of mbridgshir.
-.7943. The Life of tbRobi. London:Withry. rlatd to his studis
-. 1944.ologialasptsof spis-formation in passrinirds.Ibls 86: similaritis and diffr
260-286. of his ontmpraris
-. 1945.Th Gal6pagosfinhs(Gospizina): A studyof variation.sioI ampls of unprodutiv
Pprsof th lifori dm of Sis2T: 1-|59. ..small fats
-. only with
1947. Duin's Fihes:A ss o th GrlBiologil Thor of
uoltion.amridg:amridgUnivrsityPrss. livd) wr afraid t
-. 1950.Robi Redbresl.oford: Oxford UnivrsityPrss. Lamark first st fo:
-. 1954. Th Nturl Rgultioof iml Numbrs.oxford: lrndonPrss. trodutory ltur h
-. in our.London: Mthun.
1956. St,uifts lassin 1800. By 180
-. -hor
|957. ullutilr d hristiBlif:h Ursoludo|lit. arth, from th simpl
London:thun.
-, dud y natural aus
1966. PopultionStudiesof Birds. Oxford: larndonPrss.
-, 197I. ologil Isoltili Birds. London: Blakwll. mous xpositions of h
-' 1973.y lif as n amaturornithologist[withappriatins y si (Zoo\ogia| Philosop}
ollagus]. Ibis 115: 421431. svn-volum Work ol
Prrins,. .2004. Lk' David Lamrt(1910_1973).In H. . G. Matthwand sns urtbres (Naturt
B. Harrison, eds',ford Ditior of Ntiol Biogrph.oxford: Oxford
Lamark sought to
UnivrsityPrss.
Thorp, \/.|7, 1974. David LamrtLk, 19|0-197. Birlgrphilmlirsof volutionry thorizir
Fellos of th Rol Soitof Lodo 20z 271-29. th sussivprodu
.Winr,
J. 1994. Th Bk of the Fi: Storlf utllutioni ur im.Nvl th diffrnt faultis
York: Alfrd Knopf. -]..
maintaind that th v
to b' gnratdspon
sifid, h laimd, as t
,.po
Lamark, Jan-Baptiste (|7 4 4_|829) powr of lif'' or
omplx'' (Lamark 1
Jan-Baptist Lamark Was th first iologist to st forth road, ompr- partiular irumstan
hnsiv thory of organi volution. Othrs for him had raisd th possi-
ntural ation of flui
ility that nw organi forms had appard in th ours of th arth's
said, for th ordrly sr
history, but Lamark was th first to dvlop at lngth th ida that all th
ths auss, in ontr
diffrnt forms of lif on arth had n produd sussivly,ginning
th min sal.It invo
with th vry simplst forms and poding gradually to th most omplx.
most famous: th id
 Lmrk 67
that H plad man in a sp-
Lamark initially mad his sintifirputtion as a otanist, intratingin
L,kalso souglrt to ttr ur-
the 7770s with th irl f otanistsat th Jadin du Roi (th King's Gardn)
' many sptsof man's moral
in Paris. His arr hangd dramatially io 779 whn th Jardin du Roi
risaout th moral volutior-l
Was ronstitutd as th National usum of Natural istory. H was namd
on of th profssos of th nw institution, ut not to a hair of otany. H
t " ( 1 9 7 )i s a n a t o i o g r a p h y
Ws givn instad th profssorshipof .,insts,Worms' and mirosopi ani-
h dwzrrdGry Institut. Fo
..matur''to ..profssional.'' mals.'' H soon haratrizd this part of th nimal kingdom as th ..ani-
mals without vrtra''' Although his xprtis in this ara was limitd
owldgablmrnoril nly
initially to what h knw from having n an avid olltor of shlls, h b-
[d th Godman-Salvin Gold
am ovr th nt two dads th world's lading authority on invrtrat
958 nd th Darwin dal of
zoology.
lassifying th invrtrats and dvloping a ours of instrution aout
thm wr not Larark only onrns in th 1790s. Fror te Late 1770s
onward, h had spulatd roadly aout what h took to th asi prini-
toof Drtui'sFinhs.
pls of physis, hmistry, and mtorology. In th 1790s, h also gan to
ridg:mridgshir Bird lu. think aout th history of th arth. This nw gologial intrst was dirtly
y. rltd to his studis of shlls and to th mging prolm of plaining th
in passrin
irds.Ibls 86: .Whil
similaritis and diffrrrstwn living and fossil shll typs. many
of his ontmporaris wr inlind to s his road spulativ fforts as x-
t studof vrition.sionl
arnplsof r.rnprolutiv..systrrr_uilding,''
h omplaind of ritis who dalt
, 1 :1 - l 5 9 .
rcrIBilk;gilTbor of only with ..small fats,'' who faild to addrss important issus, and who (h
)rss. livd)w afraid tht thir rputationswould hrrrt y his idas.
rrsityPrss. Lamark first st forth his rvolutionary notion of spis hang in an in-
nbers.ford:lrndn Prss. trodutory ltu h delivrd to th studtrts in his invrtrat zoology
lass in 1800. By 1802, h was arguing that all th diffrnt forms of lif on
|'if:
Ursolt',d
o|lit.
arth, from th simplst to th most omplx, had n sussivlypro-
lrndonPrss. dud y rraturalaussopratingovr immnspriods of tim. is most 1-
BIakwll. mous positions of his idas appard in 1809 tr'hts Philosophie zoologique
ithappriations
y si (Zoolgil Philosophy), and thn in 1815 in the irrtrodution t<lhis grat,
svn-volum wok on invrtrat zoology, Histoir trell ds tm
973).|nH. . G. atthw and
sns urtbres(Natural istory of th ]nvrtebrats).
l Biogrph.oford: oxford
Lamark sought to aount for mor than hang at th spis lvl. His
973.Biogrphilemoirs of volr"rtionarythorizing addrssd in ddition th natur and origin of life,
1-29. th sussiv prodution of tlr diffrnt forms of animal organization' and
uolutiotti our Tle. Nw th diffrnt faultis to whilr ths forms of organization giv ris. H
-J., maintaind that th vry simplst fornrs of lif had bn,and wr ontinuing
to , gnratd spontanously. Ths forms thn am sussivly divr-
sifid,h laird' s th sult of two vry diffrnt sorts of auss:(1) ..th
powr of lif'' or ..powr that tnds unasingly to mak organization mor
ompl'' (Lamark I815-t82z 1: \4) and (2) th modifying influn of
to st forth a broad, ompr-
partiular irumstans. plaind th first of ths as a funtion of th
: f o r h i m h a d i s dt h p o s s i -
natllfal tion of fluids nrving through living tissus.It was rsponsil,h
l in tlr ours of th arth's
said, for th ordrly sal of progrssion of th animal lasss.Th sond of
at lngth th ida that all th
ths auss, in ontrast' Was rsponsil for th latral rmifiations from
' d u ds u s s i v I y. g i n n i n g
th main sl.It involvd th part of Lamark,s thoy that has sin om
'aduallyto th most omplx.
most famous: th ida of th inhritan of aquird haratristis.
674 Lmrck

Lamark proposd that animals wr indud to dvlop nw haits whn rpport,1t;prd

thy wr onfrontd with hanging or nw nvironmnts. hangs in haits esstils d l'il
efi, l'pllsitio d
invitaly ld to rtain organs ing usd mor and othr organs ing usd
D6trvill.
lss' thry strngthning th formr and wakning th lattr. Diffrns
thus aquird wr passd on to th nxt gnration, alit too gradually to
apparnt on a human timsal. Ovr grat priods of tim' howvr, this
Land, Russll (b
rsultd in signifiant organi hang. Lamark was onfidnt that th dis-
tintiv faturs of suh divrs raturs as giraffs, mols, storks, nd Russl1Land is widl
snaks ould undrstood as th umulativ rsult of th inhritan of notypi volution and
haratrs aquird aus of long-maintaind haits. Although th asi in Jakson, ississip
ida of ..th inhritan of aquird haratrs'' Was not novl with him, h Shrvport, Louisiana
stands out as th first sintist to argu at lngth that th opration of this Th son of a iology
pross on a gologial timsalould produ spishang. swamps of Louisiana u
It was not until nar th nd of th nintnth ntury, wll aftr Charls H playd Littl La
Darwin had advand his own thory of volution, that th ida of th inhr- shool, and h Was a r
itan of aquird haratristis, or ..us.inhritan,''am to widly r- Land's intrstsin t
gardd as th ..Lamarkian'' xplanation f spis hang in ontrast to at th Univrsity of
Darwin's ida of volution y naturl sltion. This took pla vn though and th physial sin
Lamark's idas aout us and disus onstitutdiust on part of his thoriz- approah in th biolog
ing and Darwin himslf strongly ndorsd us-inhritnas a mhanism rading lu organiz
sondingnatura1sltionin th volutionary pross.Th first srious hal- Thir influn nou
lng to th ida of us-inhritan am from Grman iologist August versity of hiago an<
.W.ismann
in th 1880s. Although th ida prsistd into th twntith n- C. Lwontin, minnt
tury, it am inrasingly disrditd whn it faild to gain primntal volutionay iology.
onfirmation and th mpirial vidn typially itd in its support was in- tin movd to arvrd
trprtdas ing amnal to othr planations. As a graduat stud
Lamark lft his mark on sin through his ontriutions to otanial for th inhritan of
and spially zoologial systmatis, his pionring work in invrtrat pa- typi volution. A ha
lontology, and his volutionary thorizing. Although h is primrily rmm- Whil lturing on
rd today for an ida that iologists n longr apt-th ida of th quippd, ..of ours
inhritan of aquird haratristis-h nonthlss stands out in th his- polygni traits.',Thr
tory of iology as th first sintist to st forth a omprhnsiv thory of or- in th filds of quantit
gani volution. dotoral thsis.
Land ontinud to
|BLIoGRAPHY
a postdotoral studn
.!.,
Burkhardt' R. Jr. \995. h Spirit of Sstm:Lmrk d uolutior Madison. H also had
Biolog. Nw d. amridg,x4A: arvard UnivrsityPrss. from th Univrsity o1
'1790_180.
orsi, P. 1'988.Th g of Lmrk: uolutit'lrhorisi Fr, \/right th draft of hi
J. andlaum'trans.Brkly:Univrsityof CaliforniPrss.
Lamark, J.-B. 1809. Phillsophiezlologiqu,o positilds osidrtios not ar for th appr
rltiuesd l'histoirenturelldes nimu; l diuersitde lur orgnistio gns that undrlay tl
et ds faultsqu'ils obtient;u ussphsiquesqui mitite that h might hav b
I ui t dotliu mouumsqu'ils utfi'd llsqi Land's dvlopm
produist,ls usl stimt,t ls trsl'itlligd ttqui c sot
joind th faulty at t
dous.2vols.Pris:Dntu.
\8\5-1822. Histoir turllds imu ssurtbrs, prsttles Land pplid his fra
rtresgruet prtiulirsde s im,lr distributio,lurs ual dimorphism and s
lsss,lursfmills'lursgres,t l ittirldspriciplsspsqui s' typi sltion, phno

to dvlopnw haits whn
ronmnts.hangs in haits
and oth organs ing usd
lnirrgth lattr. Diffns
ltion, lit too grdually to
'riods of tim, howvr, this
l was onfidnt that th dis-
giraffes,mols, storks, and
rsult of th inlrritan of
l haits. Although th asi
' was not novl with him' h
;rh that th pration of this
spishang.
,h ntury, wll aftr harls
on, that th ida of th inhr-
tan,''am tO widly r-
pis hang in ontrast to
This took pla vn though
'd
i u s to n p a t o f h i s t h o r i z -
- i n h r i t n a s a n r
)ross.Th first srious hal-
,m Grman iologist August
sistd into th twntith n-
it faild to gain primntal
l y i t d i n i t s s r r p p o r tw a s i n -
)ns.
ris ontriutions to botanial
ringwork in invtbratpa-
houglrh is primariiy rmm-
ngr apt-th ida of th
.thlssstands out in th his-
a omphnsivthory of or.
tmrhd ullutionr
riversity Prss.
horis i Fr,1790-180.
]aliforniPrss.
lo sition ds osi drti os
diursitd lur orgltistio
ohsiqs mintienl|tn
1u,i
s t,ttt; fi,h llsqui
itIlig d u 1ui sont
prsettls
ssuertbres,
ru,le.distributio'lurs
l dspriiplss|lsqui s'1'
Lnde 675
rpportt; prd d' ilttrodtio offrnt l dtrtitio ds rtres
ssentilsd l'iml, s distitkl du ugtl et ds utrs orps turls;
fi, l'positirl ds prinips fdmttl d l 7oologie.7 vols. Paris:
_R.vB.
DtrviIl.
Land,Russll(b. 1951)
Russll Ld is widly rognizdas th rhitt of modrn thorisof ph-
notypi volution nd as an innovator in onsrvation iology. Born in 1951
in Jakson, Mississippi, Land spnt th formativ yars of his youth in
Shrvport,Louisian; Roswll, Nw io; and Sal Bah, alifornia.
Th son of iology tahr nd a ptrolum gologist, h plord th
swamps of Louisiana and ody surfd on th bahsof southrn alifornia.
H playd Littl Leagu and rd hoodd rats as a studnt in grammar
shool, and h Was a memr of math and honor soitisin high shool.
Land's intrstsin ology w awaknd whn h was an undrgraduat
at the Univrsir1,f alifornia at lrvin. A tlntd studnt in mathmatis
and th physial sins,Land lookd for pportunitis to tak a quantitativ
approirh in th iologil sins.That opportunity was providd in a wkly
h a n i s m rading lub organizd y graduat studnts Td s and Mihal Gilpin.
hir influn llourgd L:rnd to apply to graduat shool t th Uni-
vrsity of hiago and to wrk with profssors Rihard Lvins and Rihard
. Lwontin, minnt pratitionrs of th modling appoah in ology and
volutionary iology. Shortl aftr his arrival at hiago, Lvins and Lwon-
tin movd to Hrvard' Land followd suit.
As a graduat studnt at avard, Land fousd on dvloping thory
for th inhritan of quantitativ haratrs nd using it to modl phno-
typi volution. A han rmak y Lwontin hlpd st him on this ours.
.Whil
lturing on ndlis-r,Lew<lntinturnd from th halkoad nd
quippd, ..Of osall ologially important haratrs ar quantitativ'
polygni traits.'' Thr paprs (1976a, \976, 1977| tat ar now lassis
in th filds of quantitativ gntisand phnotypi volution onstitutdhis
dotorl thsis.
Land ontinud to dvlop his novl approah to phnotypi volution as
a postdotoral studnt with Jams row at th Univrsity of lWisonsin at
.!right,
Madison. H also lrad rgular onvrsationswith Swall thn rtird
.Whn
from th Univrsit of hiago nd living in adison' Land showd
Wright th drafr of is 1"979papr' anothr lassi, Wright said that h did
not ar for th approah baus it did not modl th frqunis of th
gns that undrlay th quantitativ traits. A yar latr, !right voluntrd
that h might hav n hsty in his judgmnt'
Land's dvlopmnt of quantitativ gnti thory ontinud whn h
joind th faulty at th Univsity of hiago. During his Chiago priod'
Land applid his framwork to th volution of lif-history haratrs' sx-
ual dirnorphism and sually sltdharatrs,th masurmntof phno.
typi sltiolr,plrnotypiplilstiity,and nrany othr prolms in phnotypi
676 Leke

volution. ah of ths ontributions stands out aus novl theortial if thr wer qualifid
formulations ar ombind with insightful litratur rviws and appliations C a m r i d g d u l y g a v l
to atual data. thy disovr that h w
Bginning in th 1980s, Land's interstsshiftd to onsrvation iology, thn rqustdro tin
as h took th lad in haratrizing th dmographi plight of th Northrn qunt|y rivdtop h
Spottd owl by using a omination of lassial' modrn, and original d- ors in his major filds
mographi thory. During this ongoing priod, Lnd joind th faulty at ..d
C m r i d g a l l d
th Univrsity of orgon and latr movd to th Univrsity of alifornia at ship from St. John.s
San Digo. Land is urrntly a Royal Soity Rsarh Profssor at Imprial Laky had n a
ollg, London. th summr aftr his s
pursu work in th ar
B|LIoGRAPHY
tory. Th first two a
Land, R. 1976a.Th maintnanof gntivariailityy mutationin a polygni d o u m n t t h s i sf o r
haratrwith linkd|oi.GtilRsrcb26:22I_24.
tumn of l930, h wa
1976. Natural sltionand random gntidrift in phntypivolution.
uolutio:14_34. asis of The Ston g
1977. Stattstlaltstsfor natural sltionon quantitativharatrs' u n i v r s i t i st' h r w r
uolutio7:442444. t o a r .T h s o l r q u i
1979.Quantitativgntianalysisof multivarit volution,applidto p u l i s h d o o k o u n r
rain:bodysizallomtry.uoltion3: 42-416. -cI
His Afianarho
tools. Baus Laky
s i s t a n o f S i r A r t h u r
Laky,Louis SymourBazett (t903-I972)
th Hunrian usu
Louis (pronound Lwis) S. B. Laky, th son of an nglish missionry t was to hav a prfoun
Knya in ast Afria, Was orn at hurh mission station t Kat, north- dspit dlaring his L
wst of Nairoi in Kikuyu ountry. It was th poliy f th hurh ission- yars in a hous on th
ary Soity to rotat its missionaris ak to ngland vry fourth yar so (Kith l 9 l5 ). H ws
that thy would not forgt thir nglish roots. Th Lakys wr rought human volution in r
ak in 194 a.d stayd for two yars' rturning to Knya in 1906. Thir dnfor humn vol
nt nglish sojourn was twn 1910 and 191'3, at whih tim Louis I n l 9 l , L a k yl
bgan formal shooling. In twn thos tims, thy livd t th missin sta- visit to th sit that wo
tion at Kabt. Thr young Louis joind a pr group of Kikuyu boys and susqunttim: oldr
am flunt in thir languag. Adoptd into th group' h ws formally ini- stanof Laky'sin
tiatd as a Kikuyu. Although h oftn said that h thought of himslf as a vai for mor than a th
Kikuyu, h atd and soundd lik a good British olonialist for th rst of ston tools dating a<
his lif. v o l v di n o t u h u m
\World .VarI intrruptd plans to Snd him to shool in ngland. .sfhn th i a l f v i n a m p ' h i s
family rturnd thr in 1'919, Leakey was duly snt to a publi shool in thos tools (M. Lak1
Dorst. Th phnomnn rfrrdto in ngland as ..puli'' shool is rally tition' but ir lkd tl
a privat institution. \/ith th id of a govrnss' h was instrutd in math- walking bipd. l
matis' Latin, and Frnh. Although of a longtim nglish family, Laky's ustrlop ith cs' pre
fathr had n orn in Fran, so Frnh was oftn spokn at th Laky forms hat did volr.
.!7hil
dinnr tal. Louis ould spak th languag, h vidntly did so with a nw gnri nd s
a pronound nglish ant' as did many of his ountrymn. not stik' lhoughin
Laky ntrd St. John's ollg, Camridg' as a frshman in th au- first ast Afrian and
tmn of 1922. He askd if h ould offr Kikuyu as on of th two modrn tivs frrdto s hol
languags rquird, and th univrsity turnd to authoritis in London to ask had n dvlpdt

)ut usnovl thortial
t u r r v i w s n d a p p l i a t i o n s
ftd to onsrvation biology'
raphi plight of th Northrn
:al, modrn, and original d-
, Land joind tir fauity at
l U n i v s i t yo [ a l i f o r n i a t
lsarhPrfssorat Imprial
ilit by mutation in plygni
6:22|_24.
i drift in phnotypi volution.
n quatitativharatrs'
virritvlution,pplid to
t6. -s./..
.1e72)
n of an nglish missionary to
ision station at Kat, north-
poliy of th Churh N4ission-
ngland vry fourth yr so
s. Th Lakys wr rought
ning to Knya in 1906. Thir
I |9\3, at whih tim Louis
; , t h yI i v da t t h m i s s i o n s t a -
)r group of Kikuyu boys and
h group, h was fomally ini-
at thught of hirself as a
itish olonialist for th rst of
l shool in ngland. Whn th
ly snt to a publi shool in
..puli''shool is rally
d as
'ss' h was instrutd in math-
ryim nglish farrrily, Laky's
ts oftn spokn at th Laky
;uag,h vidntly did so with
lls ountrymn.
Jg, as a frshman in th au-
yllas ol1 of th two modrn
o authoritisin London to ask
Lke 677
if th wr qualifid Kikuyu spakrsin ngland. Tlr answr Was ys' so
ambridg duly gv Laky th permission h had rqustd.Onl lat did
thy disovr that h Was on of only two Kikuyu spakrs known, so h was
thn rqustd to train th prson who am his own xaminr. I I sus-
quntly rivd top honors in his languag xams. H also rivd top hon-
ors in his majo filds of anthropology and arhology' giving him what
amridg alld a ..doulfirst'' in 1926'This arnd him rsearlrfllor'i.
ship from St. John's ollg for tlrr yars' rnwd again for thr mor.
Laky had n ak to ast Afria as part of a dinosaur-hunting group in
th summr aftr his sophomor yar, ut with th fllowship h was al to
pursu work in th :rraof his training and intrsts,namly' Afrian prhis-
tory' Th first two ast Afrian arhologial pditions allowd him to
doumnt th asis for an arhologialsqunof ast A1ri. In th au-
tumn of 1930, h was awardd a PhD from amridg Univrsity on th
asis of Th Sto Age ulturs of Ke olo (1931). Unlike Amrin
univrsitis,thr wr no oufss or xams rquird in ordr to an do-
toat.Th sole rquimntwas n aptaldotoral dissrttion,nd a
publishd ook untd.
His Afrian arhaologialwork yildd human skltalmatril s wll as
tools. Baus Laky was not traind in skltal ilnalysis, h souglrt th as-
sistanof Sir Arthur Kith, a highly rsptdantomist and th urator of
th Huntrian usum at th Royal Collg of Surgons in Londorr' Kith
was to hav a profound fft on Laky's views on hurrrnvolution. Kith,
..Darwinism''
dspit dlarirrghis lif in and spnding narly th lst 25
yars in a hous on th Darwin stat'was in fat pofoundly anti-Darwinian
(Kith 1915). H was largly th rason why many profssional studnts of
human volution in ngland and Amria rfusd to apt th vilal vi-
d n f o r h u m a n v o ! u t i o n .
In 193i' Laky ld his tlrird st Afrian arh:rologialpdition tllr e
visit to th sit that would idntifid with Laky's impat on th fild for all
susqunt tim: olduvai Gorg in what is now alld Tanzania. Th su-
stan of Laky's intrntional rputation was groundd in his work at oldu-
vai for mor than ir third of a ntuy. In th first yar, h found quntitis of
ston tools dadng ak to th vry alist mad y t anstorsof w,hat
volvd into tr humans. In July 1959, whil Lor.risws suffring from a trop-
ial fvr in amp' his wif' ary Laky, found tlr first fossil of makr of
thostools (M. Laky 1984).It h:rdan ap-sizdrain and an normous dn-
tition, ut it lakd th projting anins of a tru ap and it was an rt-
walking bipd. larly it ws latd to thos rtrmrs of tlr gnus
Austrlopithl,ts, prviousiy known only frorn South Afriil and inluding
forms that did volv into th genus Homo. As h did so oftn, Lky gv it
a nw gnri and spifi name, Zinjthrops blisei' Th gnus nam did
,,Zinj.,,
not stik' although informally it is still known as Not only was it th
first ast Afrian and th most sptaulr mmr of th arly hurnn rla.
tivs rfrrd to as hominids, ut th thniqus of radioativ isotop irnalysis
had n dvlopd to th tl1ttht it ould givn an asolr.rtdat of ovr
678 Ldrbrg

1.5 million yars. Previously, th Australopithins known only from South ss in atri that r,
Afriir wr assumd to approximatly half a million yars old and muh that th prossLdr
too nt to hav playd r<>l irr l.luman volution. vn though Ztni was a or plasmid, that mdia
sidlirr,llot o th pth toward tu humans, its grar ntiquity nd primitiv othr' This pss,kr
hominid traits providd sr.tstan for Charls Drwin's (1s71) suggstionthat nbld sintiststo rr
Afria was th originl lol for th volution of hr-rnr:rn
form. soms' in a mannr lo
Laky took full advantag of th opportunity provid ed Zinj. H wrot y Thomas Hunt or
a sisof thnial as wll as popular dsriptiv artils. vn mor ff- Sond, working
'sisonsin,
tiv was his prsnon th ltur pltform. A harismati and olorful pu- Ldrrg
li spakr' his frqunt intrnirtionalapparansrought him arningsthat as vtors to mov g
supportd furthr fildwork, spially at Olduvai Gorg nd lswhr in this virus-mdiatdp
ast Afrir.This also nour:rglothrs to n.agin th sarh for arl ho- rol irr failitating ya
nrinids and thir tools lsrvhrin ast Afri. Hrdly a yar gos y now atil gntis.
without furthr disovris. Third, Ldrrg :
Laky also promotd fildwork on living grat aps-himpanzs and rplia-platingpri
gorillas in Afria and orangutans in SouthastAsia-y a sris of prot6g6s rial mutations-in par
who hav grtly pandd what w know aout t lif and ways of hu- ourrd prior to th
mans' losst nonhumn rlativs. mutations ould not
rtd mannr to th t
BIBLloGRAPY
at ando, with sl
ol' S. |975. Lk1,.sLk:.l'h l-if r'tfLouis S\'tlt<lr
tszttLk'Nrv York: nti dapttion. In o
HarourtBraJvaIrovih. winian mhanisms i
D:lrwin,. 1871.Th Dstttof , d Sltil i Rltitl/rl Se.Lor.rdln: Salvador Luria and tr
John Murra. sion, th rplia-plati
ith,A. 1975 . h tiquittlf . London:Williamsand Norgat.
Lky,L. s. B. 19I. Th StonAg ltrsof llon.amridg: it providd a striking
ambridgUnivrsityPrss. on a mastr ptri disl
1974.B thuide:loirs,19.]2_19.'1' Nll,Y.ok:arourtBra mutnt dsndantsi
Jovanovih. .!idnfld popultion, using a v
Laky'N,I'l984. Dislosingtha Pst:lt utobiogrD. Lodn: and sujt to sltionfo
Nilson.
lv{orll,V. 1995. Astrl Pssios:b Lk Flill'd th Qst for from th orrspond
Humnkid'sBgiiltgs. Nw York: Simlnand SlrLrstr. -.L.B. is<llatmutations witl
ditions that favord tl
At th tim that L
Joshua(1925_200s)
Ldrbrg, primnts, littl wa
Joshua LdrrgWas on of th lading miroiologistsand gntiistsof th ads, th fild of mi
twntith ntury. His primntsplord th natu of ftrndamntalgn. pralrs to undrst
ti nrhanismsin rria:rnd thus slrdlight on prosssthat hlpd shp attntion to volutio
th volution of th rrrostaundnt organisms on ilrth. For his ontriutions, work on mirobil v
lr rivd a Nol Priz irr 1958, at th ag of 3. Ldrrg also advisd prativs.First, th fi
poliymakrs on suh mattrs as mrging inftious disassand iologial trial spisto or
warfar, and h was awardd th Prsidntial edal of F.rdomin 2006. ry y arl !7os of
Ldrrg mad thr major ontriutions to undstanding atrial gn- girnismsthat ar dist
tis' First, as a graduat stulntworking with dwrd Tatum at Yal Univr_ of molular voluti
sity, Ldrrg showd that batria ould romin thir gnsto produ lrang within and a
nW gnotyps.This primnt dmonstratdth istnof a sual pro_ of th gntimhar
 Ldrberg 679

ss in atria that was not prviously known. Susqunt rsarh showd

ns known only from South
that th pross Ldrrg disovrd involvs an trahromosomal lmnt,
million yars old and muh
or plasmid' that mdiats th movmnt of atrial gns from on ll to an-
ion. vn though Zin1 was a
othr. This pross' known as onjugation, also providd a powrful tool that
g r te n t i q u i t ya n d p i m i t i v
nald sintists to map th physial psition of atrial gns on hromo.
rwin's(1871) suggstionthat
soms' in a mannr logially similar to th mapping studis pionerd arlir
human form.
y Thomas Hunt N4organ and ollagus with fruit flis.
providd Ztn1. F{ wrot
Sond, working with his studnt' Norton Zinder, at th Univrsity of
v artiis. ven mor ff- .Wisonsin,
Ldrrg disovrd that viruss that inft atria ould srv
harismatiand olorful pu-
as vtors to mov gnti matrial from on strain to anothr. Thy alld
s brought him arnings that
this virus-mdiatd pross transdution and it, too, playd an important
v i G o r g a n d l s w h r i n
rol in failitating yars of inrasingly dtaild and sophistiatdstudis of
g in th sarhfor arly ho-
atrialgntis.
Hadly a yar gos y now
Third, Ldrrg and his first wif, sthr Ldrrg' dvlopd th
rplia-platingprimnt.This lgantprimnt dmonstratdthat at-
at aps-himpanzs and
rial mutations-in partiular, thos that onfr rsistanto viral inftions-
\sia-y a sris of protgs
ourrd prior to th atria ing xposd to viruss. Therfor' th
)ut th lif and ways of hu-
mutations ould not hav n ausd y th atria rsponding in a di-
rtd mannr to th sltiv agnt. This work showd that mutations aros
at random, with sltion providing th dirtional for rsponsil for g-
ur Bztt Lk. Nw York: nti adaptation. In othr words, atria ar sujt to th Sam basi Dar-
.shil
winian mhanisms as highr organisms. an arlir xprimnt by
l in Rltitl,o Sex. l,ondon: Salvador Luria and a Dlriik (1943) had supportd th sam onlu-
sion, th rplia-plating xprimnt sttld th issu vn for skptis aus
ims nd Nrgat.
it providd a striking visual dmonstration. That is' rar mutants that aros
lo. ambridg:
on a mastr ptri dish, whih wnt undttd asnt any sltion, yildd
1. Nw York: Harourt Br mutant dsndants in th sam rltiv loations following transfr of th
population' using a vlvt pad, to svral rplia plats whr atria wr
lph. London: Widnfld and sujt to sltion for rsistan to viral inftion. In fat' y srning lls
from th orrsponding loation on th mastr plat, th Ldrrgs ould
il d tb Qust flr
-.L.B. isolat mutations without dirtly xposing th batria to th sltiv on-
1Shustr.
ditions that favord th mutant phnotyp.
At th tim that Ldrrg and his ollagusprformd ths pionring
primnts, littl was known aout atrial volution. For th nt fW d-
ads, th fild of miroiology mostly pursud gnti and molular ap-
iologists and gntiists of th
proahs to undrstanding llular strrrtur and funtion, paying littl
) natur of fundamntal gn-
,n prosssthat hlpd shap attntion to volution. Howvr, th past fw dads hav sn inrasing
work on miroial volution, whih has followd svral approahs and im-
n r t h .F o r h i s o n t r i u t i o n s ,
prativs. First, th fild of molular volution rvald th rlationships of
lf 33. Ldrrgalso advisd
atrial spis to on anothr and to othr organisms, inluding th disov-
tious disassand iologial
ry y arl \Wosof th Arha' a group of singl-lld' non-nulatd r-
Idalof Frdom tn206'
ganisms that ar distint in many rsptsfrom tru atria.Th approah
undrstandingatrial gn_
of molular volution has also n usd to amin th xtnt of gn .
idward Tatum at Yal Univr-
hang within and among atrial spis,thus tnding our undrstanding
lmin thir gns to produ
of th gnti mhanisms disovrd y Ldrrg in th laboratory to thir
th xistnf a sual pro-
Lssonsin omparativAnatomy

importan in natur. Sond, owing to thir rapid gnrations and othr ompr (Lssos il
tratabl faturs, atria ar now oftn usd in primnts that sk to thr mor volums'
undrstand th dynamis of phnotypi and gnomi volution as wll as p u l i s h d y 1 8 0 5 . V
to tst partiular hypothss aout volutionary phnomna. Finally, thr th organs of snsatio
is growing attntion to th rol that vlution plays in th mrgn of and th organs for pr
nw inftious disass,inluding th aquisition of viruln fators that and srtion. uvir
allow pathogns to inft thir hosts as wll as th sprad of antiioti r- ..th ono
tions on
sistan. Th mhanisms of gnti hang disovrd y Ldrrg prfa to th work, l
hav n found to vry important in th mrgn of many nw a- ing dstroyd''(uv
trial pathogns. uvir quotd Immar
..us'' of vry sing
ILIoGRAPHY
organism in itslf. Dr
lna, S. F., and R. . Lnski. 2003. volution xprimntswith miroorganisms: notably Antoin-Laur
Th dynamisand gntibassof adaptation.Ntre ReuitusGetis4 tion of parts" (1789\
457469.
a n a t o m y ( 1 8 0 5 ) ' u v
Ldrbrg, I. 1997.Inftious disasas an volutionaryparadigm.mrgmg
IfctiosDisss: 417423. tion, nutrition, vision
..laws of oxistn'
Ldrbrg, J., and . M. Ldrrg.|952. Rpliaplatingand indirtsltionof
btrialmutants.Jourl lf Btriolog63: 99406. tion of on organ or
Luria, S. .' and M. Dlriik. 1943. utations of atriafrom virus snsitivityto th whol organism a
virus rsistance.Getis28 491-51\.
funtions of intrnal
National Lirary of diin.N.d. Profiisin sin.Th JoshuaLdrrgpaprs.
http://profi ls.nlm.nih.gov/BB/. divisions within th l
Tatum, . L., and J. Ldrrg.|947. Genrminationin th trium tiv anatomy provd
shrihioli. Jourl of Bteriolog5 673-684. sifiation.
.Wos,
. R., O. Kandlr,and . L. !hlis.1990.Towardsa naturalsystmof uvir plad stron
organisms:Proposalfor th domainsArha,Btria,and uarya. tion y insisting that
Prodigsof th NtklI Adm of SinsUS 87 4576-4579.
isms. Lif, h agrd
Ztndr, N. D., and J. Ldrrg.1952. Gntihangi Slmoll.lourl of
Bteriolog 64: 67 9-699. -R..L. proprty to tmpora
agnts. H did not dt
tim unvil th intim
..animal mahin.'' Fi
Lessons i Comprtiue Antom (Gorgsuvir)
tn of a hain of
In his vr-usylif, th yars 1795 to 1802 wr partiularly hti ons for isms to man. Gradua
Gorgs Cuvir. on July 2, 1,795,th profssors' assmly at th Mus6um intrnal organs, and
national d'histoir naturll approvd his nomination s assistant to Jan- twn ah of th ma
laud N4rtrud, th anatomy profsso, and h startd in arnst tO arrang anatomy was disov
th omparativ anatomy olltion. H gav lturs at th museum and
rad sintifi paps to a varity of soitis (th Soit6 Philomatiqu and |BLIoGRAPH
th Natural istory Soity in partiular) and at th Institut d Fran (insti- C o l m a nW , . 1 9 6 4 .G
tutd in 7795 to rpla th old Aadmis, inluding th Aad6mi frangais ambridg,MA:
and th Aad6mi ds sins).uvir took full advantagof th musum li- uvir,G. \80.Leqo
rary that nfitd, as did th musum itslf, from important additions . Baudouin.
q u i r d f r o m t h o s u o p n o u n t r i s o n q u r d y t h F r n h a r m i s . 185.Lgosd
Baudouin.
During 1798 and 1799, assistd y Andr6.Mari-Constant Dum6ril
J u s s i uA, . L . d .1 7 8 9
(1774*1860), uvir workd on th nts Dum6ril had takn at his lturs O u t r a m ,D . 1 ' 9 8 4 .G
in th prvious four yars. Th first two volumS of t'eLos d'tomie ruolutiorFrn

apid gnrationsand othr
in pimntstht sk to
lnomi volution as wll as
/ phnomna.Finally, thr
plys in th mrgn of
on of viruln fators that
th spra i f antiioti -
disvrd y Ldrrg
nrgnof many nw a.
mrrtswith mirorgnisms:
,ttlr
Ruies Gtis 4:
rry paradigm. mrging
Ltingand indirt sltion of
9406.
tria from virus snsitivity to
. Th Jshua Ldrrg pprs.
ation in th batrium
-684.
fowrds nturl s,vstmof
rtria'and uary.
s US 87: 4576-4579.
'ngein SImoll.lourl of
-R..L'
)orgsuvir)
. partiularly hti ons for
rs' assmlyat th usum
ination s assistant to Jan-
startdin arnstto arrang
ltursat th musum and
h Soi6t6Philomatiqu and
th Institutd Fran (insti.
' u d i gt h A a d m i f l r n g i s
advntagof th musum ii-
.rom important additions a-
d y th Frnh arnris.
|r6-ari-onstant Dumril
r6rilhad takn at his lturs
s of th Lqons d.lttomie
Lssonsin omparativAnatomy 681
()|pLlr(Lessons i omprtiue tomy/ appard in arh 1800;
thr mor voluts, ditd y Gorgs Louis Duvrno (1777_1'85.5), wr
pulishd y 1805. Volum 1 dalt with th organs of movmnt; volum 2'
th organs of snsation;volums 3 and 4, digstion, irulation, rspirarion,
and th organs for produing sounds;and volum 5, th organs of gnration
and srtion.uvir dvotd his first ltur to mthodologial onsidra-
tilns on ..th onomy of lif.'' Living organisms, h had dlard in th
prf to th work, ar ..mahins. . . that annot takn apart without
ingdstroyd'' (Cuvir 1800, 1: V). Ltur 1 pandd upon this onpt.
uvir quotd Immanul Kant to stat that th raison d'tr, th intimat
..:lus''
of vry sirrglliving part Or organ' was in th ..whol,'' that is, th
organism in itslf. Drawing upon prvious otanil and anatomial work,
notaly Antoin-Lurnt d Jussiu'sonpt of ..orrlationand sr-rordina-
tin of parts'' (L789) and Fli Viq-d'Azyr's appliation of th prinipl to
anatomy (1805)' uvir showd tlrat systms of organs (digstion,loomo-
tion, nutrition, vision' t.) ar in vry animal lass rlatd to ah othr y
..lws
of oistn''(Cuvir 1800, 1: 57). Thus th prsnand onfigura-
ti<rnof on orgn Or Prt lgitimatdognt ddutions s to th strutuof
tlr whol organisrn and its way of lif. It ws in any as th strutur and
funtions of intrnal organs that onstitutd th ky to all wll-groundd
divisions within tlr animal kingdom. Thrfor th progrss of ompara-
tiv anatomy prov.d ssntial to th dvlopmnt of a natural systm of las-
sifiation.
uvir plad strong mph;rsison th impossiility of spontanosgnra-
tion y insisting that all known organisms ar dsndd from othr organ-
isms. Lif, h agrd with his frind and ollagu vir Bihat, is th
proprty to tmporarily rsist th dstrutiv ation of physiohmial
gnts.H did not dny, howvr, that th progrss of hmistry would in
tim unvil th intirrrtstrutur and poprtis of h omponnt of th
..animal
mahin.'' Finally, h dvotd a long stion to disprov th xis-
tn of a hain of ings,gradually asnding from th most simpl organ-
isms to man. Gradual variation is possibl only within th sam systm of
intrnal organs' and thr ar no known forms making th transitin -
twn ah of th main anatomil ar-rdfuntional animirl plans omprtiv
anatomy was disovringand dsriing.
BIBLIOGRAPHY
olman,w. 1964. Gorgstluir'Zoologist: Stud in th istor of uolutio.
ambridg,A: HrvardUnivrsityPrss.
uvir,G. 1800.Lglsd'tomi()t|7pL1r. Vols. 1-2. C. Dumril,d' Paris:
Baudouin.
185.I-gt-lsd'tlmiot|Iplr.
Vols. 3_5. G. L. Duvrnoy,d. Pris:
Baudouin.
Jussiu,. L. d. 1'789.rpltrm.Paris:Hrissant.
outram, D' |984. ()orgeuuir:Votil,Sincend uthorit i Post-
rolutkr\ F rul. Manhstr:ahstrUnivsityPrss.
682 Leonti

Rudwik, N{. 1997. Gorges uuier, Fossil Bons d Gologil tstrophs: Th findings of L
Nlu Trsltios d Itrprettils of th Primr xrs. Chiago: Univrsity
Huy, suggstd-gl
of hiago Prss.
Viq-d'Azyr, F. |794 [1805]. uurs reuillis t publies ue ds ots t u|1 vry gn lous is po
discours sur s ui t ss ouurgs |)r . Jqus L. oru d l Srth. 6 (Huby and Lwontil
vols. Paris: Baudouin. -P.. This might sm to
viw of volution (i..
ural populations, mai
Lwontin, Rihard (. 1929) and against th
..las
(i.., that natural sl
Rihard Lwontin has provd to vry it as influntial in population g- '
ntis as his mntor, Thodosius Dozhansky. Lik Dozhansky' Lwontin within populations).
is wll known for artiulating a vry dfinit prsptivof th fild and th th rsults wr parad
most important prolms to solvd-inluding, lik Dozhansky' an s- hr was just too
pially pozz|ing..paradox.'' Lik Dozhansky, Lwontin is wll known for stiv' Lwontin ar1
a highly original approh to masuring variation at th gnti lvl and Kimura and Jams
for his own viws of how st to plain that vriation. Lik Dozhansky, a htrozygot ross (
periority at a partiul
Lwontin twakd molular iologists for suggstingthat mlulr pro-
sssWr mor fundamntal than th volutionary prosssthat atually rior homozygots and
gav ris to thm. Lik Dozhansky' Lwontin has n dply onrnd of th population. Tht
with th roadr moral and politial signifian of gntis and volution. suprior' th gratr
population, until th I
ary biology. And, lik his tahr, h has mntord sors of graduat and
postdotoral studnts. But ths ar similaritis at a vry high lvl of Morovr, Lwontil
gnrality. othrwis, ths two influntial population gntiistsar vry ttr snsof doum
diffrnt! variation'' tht thr
Along with Dozhansky' rnst ayr, Gorg Simpson, and othr volution- tion, no mattr whth
ary iologists' Lwontin grtdth fanfar surrounding th advnt of molu- th alan thoristsl
lar gntis in th 1960s with a ationary rmindr to kp in mind th proposd y th lass
xplaind in trms of
distintion twn ffiint and final aus in iology. Th molular tion, a position that L
onfigurations of living organisms ar th ffiint auss of iologial What is/was to d
phnomna ut not thir final auss. That is, xpt in a trival sns' tial tt, he Geti
th laws of gntis ar not th rsult of th strutur of DNA, ut rathr suh a rih thorti
DNA has bn hosn y natural sltionfrom among an immns va- op with th ody oJ
rity of moluls prisly aus it fits th rquirmnts of an volvd ary insight that in a flr
gnti systm. DNA is only th tati adoptd in th ours of working tivity did for th ont
out an volutionary stratgy.That is why som orgnisms n gt on th prolm mor pr
without it. (Lwontin 1964, 566| n? I livit is th
on rason for th
But h saw onsidralpromis in th nw fild and was at th vanguard of not ..mpirially suffi<
applying molular gnti thniqus to volutionary issus. In prtiular, b masurd dirtly,
h hlpd to pionr th us of gl ltrophorsis to doumnt variation twn altrnativ at
los to th gnti lvl-namly, at th lvl of amino aid diffrns rn pistmologial on
protins (du to nulotid diffrns in th gns that od for thos pro- infamous-for prsist
tins).In so doing, h hlpd stalishth fild of molula population gn- iologists an possib
tis. His studnt, Martin Kritman, WaS th first population gntiist to know. His ritiqus o
study variation dirtly at th DNA nulotid lvl, using mor rnt DNA drls of Sn aro an
squningthniqus. Gould) and gnti d

d G llogicl tstrop h es :
m r t s . h i a g : U n i v r s i t y
'ublis u des otas t Im
s L. oru d l Srth. 6
; influntialin population g-
L i k D o z h a n s k y ,L w o n t i n
rsptivof th fild and th
ling, lik Dozhansky' an s-
., Lwontin is wll known for
ltion at th gnti lvl and
variation. Lik Dozhansky,
lggstingthat molular pro-
ionary prosssthat atua1ly
n hs n dply onrnd
r of gntisand volution-
ttordsors of graduat and
itis at vy high lvl of
pulation gntiistsar vry
Simpson,and othr volution_
oundingth advnt of moiu-
ndr to kp irr mind th
: in iology. Th molullr
ffiintauss of iologial
is, pt in a trival sns,
trutur of DNA, ut rathr
rom mong an immns va-
l rquirmntsof n volvd
:din th ours of working
som organlsms an get on
:ld and Was at th vanguard of
'utionaryissus' In partitrlar'
t o r s i st o d o u m n t v r i t i o n
l o f m i n o a i d d i f f r n
gnsthat od for thos pro-
of molularpopulation gn.
f i r s t p o p u I a r i o ng n r i i s t o
lvl,using mor rnt DNA
I'luoti 68
Th findings of Lwontirr and his arly ollaorator in this work, Jak
Huy, suggstd-givn th inhrnt iass of tlrir mthods-tht n.ly
vry gn lous is polymorplri, and that a third of all loi ar htrozygous
(uy and Lwontin 1966; Lwontin and Hr"ry|96Q Lw>ntin|974).
-P.. This might sm to b strong vidn in favor of Dozhansky's ..alan''
viw of volution (i..,that thr is onsidralgntivariation within nat-
ural populations' maintaind largly y sltion irr fvor of htrozygots)
and against th ..lassial''position of Dozhansky's arhrivl, . J. ullr
(i.., that natural sltion in favor of optimal gns liminats vri:rtion
within populations). To his tahr'slrilgrin, Lwontin argud instad that
th rsultswr paradoxial' and not t ll sy to plain.
Thr was illst too muh variation to mak sns from th alan pr-
sptiv'Lwontin argud, iting similar onsidrtionsproposd y otoo
Kimura and Jams row' On major prolm is tlrat hlf of t offspring of
a htrozygot ross (x) ar hornzygots (, ) . Heterozygot su-
priority at a patiular gn lous thus ntails th prdution of many inf-
rior homozygots and a orrsponding drasin th rprodutiv apatty
of th ppultion. Th gratr th numr of loi t whih htrozygotsa
srrprior' th gratr th damag don to th rprodutiv apaity of th
population, until th population would b quit unal to prptuatitslf.
orovr, Lwontin argud, no altrativto tl.lalan viw mks any
ttr snsof doumntd pattrns of gnti variation. It is a ..parado of
variation'' that thr is too muh to plaind in trms f natural sl-
tion, lo mattr whthr it is variation-maintainingsort of sltionsuh as
th alan thorists had proposd, or a variation-rduingsot suh as that
proposd y th lassial thorists.Tlrr was also too muh variation to
plaind in trms of th sltiv insignifian or nutrality of th vria-
tion' a position that Lwontin dud ..no-lssia1.''
.Wht
is/was to dorr?or as Lwontin psd th qustion in lris influrr-
tial txt, Th Geti Bsis of uolutilnr bge (7974,267),*ov an
suh a rih thotilstrutur as population gIltisfail so ompltly to
op with the ody of fat? Ar w simply missing som ritial rvolution-
ary insight that in a flash will mak it ll om right' as th Prinipl of ltla-
tivity did for the ontraditory vidn on th prpagatiorr of light? or is
th problm mor prvading' mor dply uilt into th struturof our si-
n? I livit is the lattr.''
On rason for th impass,h argud, is that population gntithory is
not ..mpirially suffiint.''For aml, it inluds paramtrsthat annot
masurd dirtly, or with suffiint aury' to distinguish larly -
twen altrnativ ausal aounts. his rflts Lwontirr's mor gnrl
si n pistmologial onrns-indd, his skptiism. is w|l known-vn
infamous-for prsistntly qustioning whthr gntiists nd volutionary
biologists an possily know what thy want to knW and oftn laim to
know. His ritiqus of adaptationists(inluding his I979 arti|e..TlrSpan-
drls of San aro and th PanglossianParadigm''' oauthord with Stphn
Gould) and gnti dtrmirristsar largly pistmologial.Adaptationists
684 L,llttin

ar-rdgnti dtrminists do not know nough aout th inhritan of th as]in th way that m
traits thy study and th volutionary fors ating on thos trits. Morovr, ally ror a as of sho
no on knows. This is a ntral motif of Lwontin's Geti Bszl and his livrs in th gnti as
stibsquntrviws of tlr stat of population gntis'Th titl lf on suh Lwontin's pisrmrl
rviw wll illustratshis disontnt vn with mtlrods of his oWIl dvising: matrialism, whih play
..ltrophorsisin
th Dvlopmnt of volutionary Gntis:ilston or
Millstn?'' (1991I. D i . r | t i e lm r r i l i
A sond rson for th filur of thory to mk snsof data in popul- mtlrod for solvir-rgpar
tion gntisis that pattr.nsof gnti vriation an du in lrrgpart to sis provids an ovrv
..history''-6or instan, not just whih nvironmnts a population pri- forms of dogmatism a
ns' ut th ordr in whih thos nvirtlnmntsare nountrd(Lwontin tht history ma' lav
|967). Population gnti hory,howvr, is an quilirir.rmthory tlrilt dis- rrrring ar dul as1
oLlntshistoiirlontingnis.Lwontin's disussiorrof th ahistorial har- ard that th onditil
atr of popultion gntis is partly pistmologial: history is ignord dstroyd y th
largly eausth past is so oftn unrovral.But it is also partly politi- ojts in tim nd sp
al. In population gntis,as in othr :rrilsof th rraturl nd soil si- tiorrs.Rmmbr tlrt
ns,quilirium thoris prvaild in tlr twntitl.lntury in part aus lost whn phnom
..prouptionwith tht all th othr avt
of a vry gnral staility'' (Lwontin 1974,269).
For Lwontin, th way forward in sin involvs, in part, posing th appliation to diffr
idologis that hlp to prptu:ltinorrt viws of h world' For ampl' ( L v i n s n d L ' r l l l r i
h rgud tn cnetic B-.istlrt th lssial/alanontrovrsyould not
understood without taking into aount th onfliting politial idals of B|LIoGRAPHY
Mullr and Dobzhansky. Gould, S. J.' and R. -.Lw
A ommon mistak of Mullr nd Dlzhansky' ording t<l Lwontin, Pr-rglssin palrdigm
Was thir gl.ltidtrrrrinistassumptiol-tthat xisting pattrns of gnti di- l|th I|ol SlittI
vrsity hav moal and politial impliations. For his own part' h hs laimd, Huv,J. I-.'and R. (-. [-
..I htrlzygosity in natu
do not liv that th ultimat issus dpnd on how muh divrsity -
in Drosophil psud
ists'' (Lwontin t al. 2001,4).lnstad of trying to did how o organiz I . v i n s 'R . ' n d R . [ , r r ' . l l l
soity on th ll,sisof gnti findings,w shold Llsour st undrstanding R' I-w<lntir-r. ds., h
of gntis' psyhology, soiology, and so forth to dtrminwhat intrvn- Harvrd Urrivsity Pr
tions would rsult in th sort of soity w aspir to (Lwontin t al. 2001, l,wontin, R. . 1964. A m
1 45 : 5 6 6 - 5 6 7 .
4344i Lwontin t al. 1984). Along th sm lins, Lwontin has lftnn
| 9 6 7. T p r i n i p l
onrnd to rr.rtth harg tht his skptiism onrning gnti dt-
M . . K p l a n , d s . ,
minism rndrs him ultural dtrminisr instad.As h quots Karl Mar' l 1|trp rttiI1of u olu
..Th matrialist dotrin that mn ar
th produts of irumstansand up- l972. h pti
ringing, and tht, thrfor, hanged mn ar prodts of othr irum- 8l-398.
sns and hngd upirrging, fogts that it is mn that lrang |974. h Glttt
Ullivrsity Pss.
iumstnsnd that th duator himslf nds duating'' (Lwontin t
1991. ltoph<l
a I .1 9 8 4 , 2 6 7 ) . ilston r millstl:
Th on as whr Lwontin has laimd politial signifianfor his own Lwotirr,R. ., and i. L.
findings in populzrtiongntishas to do with th sorts of studis that h ini_ hrrzygsity in n:lttlr
..Th Apportionrnnt of Huma htrozygosit],, i nrur
tiatd in his lassi 1972 ptlper Divrsity,''
54: 59-5-609.
in whih h argued that th proportion of variation within so-alld ras is
Lwontin, R. ., D. Pul, J.
far gratr (mor than 10 tirns) than th variation twn ras (s also
Llr.tlntin. ln R. S. Sing
Lwontin t al. 2001, 43). Raial diffrns.and all th othr soil nd po- h i kig but L-t'll ut
litial diffrns that lrirv aomp:rnid thm, ar thus not iologially f 2 . 6 l . C e r t l l ' j t l (:. .

about th inhritan of th
ing on tlrostraits. Morovr'
ontin's Geti Basis and his
Th titl of on suh
4r-rtis.
mthodsf his own dvising:
ionary Gntis:Milston or
mk snsof data in popula_
)n an du in lag part to
) n m n r sa p o p u l t i o n x p r i -
(Lwontin
1 t sa r l ] o u n t r d
rn quilibrium thoy that dis_
:ussionf th Lrhistorialhar-
mologial: history is ignord
l. But it is also partly politi-
of th ntul and soial si-
)ntithntury in part aus
'' (L,r,ontin1974, 269)'
nvolvs,in part, xposing th
ws of th wold. For ampi,
[anontrovrsyould not
onfliting politial ideals of
nsky, aording to Lwontin,
istingpilttrnsof gnti di-
lr his own part' h has laimd,
n d r l r th t l w m u h d i v s i t y x -
i n g t t l d i d h o w t o o r g n i z
uld us our st undrstanding
.h to dtrmin what intrvn-
lpir to (Lwontin t al. 2001,
lins, Lwntin has oftn n
;ism onrninggnti dtr-
:ad.As h quots Karl r,
cutsof irumstansand up-
r r p o d t r t so f o t h r i
rhat it is mn that hang
r d s d i r a t i n g '(. L w o n t i n t
litial signifiane for his own
th sorts of studisthat h ini-
onlnnt of Human Divsity,''
i a t i o nw i t h i n s . a l l d
'iation twnras (s also
rnd ll th othr soil and po_
?m' ar thus not iologially
Lotin 685
sdin th way that mny h:rd supposd.Aording to Lwontin, this is -
ally more a as oi showing that gntisis not as politially signifi:rntas -
livrsin th gnti sis of raial diffrnshad imagind.
Lwontin's pistmol>gial and politial intrstsmrg in his dialtial
mtrialism.whih plays a autionirry,huristi rol in his work:
Dialtil matrialisrn is not, nd tlv lras en, 21programmati
mthod for solving partiular physial problms. Rathr, dialtial analy.
sis provids an ovrviw nd a st of wanirrg signs ginsr patiular
forms of dogmatism and nrrownss f thought. It tils us, ..Rmmr
that history my lv n impotarrt tra. Rmm that ing and
oming r dual asptsf natur. Rmrnbr that onditions hang
and that th onditions nrssary to th initition of som pross may
dstroyd y th pross itslf. Rmmr to pay ttnrion to ral
ojtsin tim and strra nd nt los thm in uttrly idalizd astra.
tions. Rmmbr that qualitativ fftsof ntxt and intration may
lost whn phnomna ar isoltd.''And aov all ls' ..Rmmr
that ll th thr avats r onl rnrindrsand warning signs whs
appliation to diffrnt irumstnsof th ral world is ontingnt.''
(Lvirrsand Lu.ontin 198.5,|91-|92)
B|LIoGRAPHY
Gould,S..|..ant1R. . L,ontln, 1979.Th spandrlsof Sn o arrdth
Panglssin paradigm:A ritiquof th adaptationisr programm.Proedigs
lftll Rry'lSlitl|.Lodl,Srists 20.5:'581_598.
Huy,.|.L., arrdR. . Lwontirl.1'966.A mlularapproahto th studyof gni
htrozygosity irrnaturalopulations.l. h nLrm f lllst diffrntloi
in Drlslphilpsudllr'lbscut.. Getics54:577_594.
Lr.ins,R.' and R. Lrvontirr. 198.5.Th prolnrof l,ysnkoism. In R. Lvinsand
R. Lwontir-r, ds.,}c Diltiltsiologist, \6-196. arnridg, A:
HarvardUnivrsityPrss.
Lwotrtin,R. . l964. A moltrlar mssih: Th nw gosplin gntis? Sic
1.45566-567.
|967.Teprir-ripl in volution.In P. S. lorlradilnd
of histliit
. . Kpln, eds.,themtilhllgsto th No-Drilti
.Wistr
Intrprttit,l <l|-u<lllttil,81_88.Phildlhi: InstitutPrss.
|972. Th apportionmntof human divrsity.ultionrBirlllg6:
u m - 3 81 - 3 9 8 .
1974. h Geti l3sisof ulllutilrhg.Nw York: olurr-ri
UnivrsityPrss.
l991. },ltrlphorsis in th dvlpmnt of volrrtionrygntis:
ilstonr millston?Ctis128:657_662..
LwontiIl,R. (].'ntl.].L. u1'.1966.A nlolularapproahto th stud)'of grri
htrozygsity in ntr.lralpopultilns. II. Amuntf varitinand dgrof
htrlzygosity irl rlatualp1rrrlatinsof Drosophil psudolbscur. Gtics
54:595-609.
s i s
l-wot-ttit-t,
R. ., D. aul, |. Batty,ad . B. Krilr.rs.2001.Intrviw of R. .
Lwlntin.In R. S. Singh,. B. Krims,D. B. Paul,ndJ. Batty'ds.,
hiking llout |,Itttitl: Histlril,PbiloslphiId Politi,lIPrsptiL's,
22_61.mridg:()rnridg UnivrsityPrss.
686 Lineus

Lwontin, R. ., S. Rs' nd L. J. Kamin. 1984. No, i r Ges: Biollg, that inludd whals z
Ideolog, d m Ntur. Nw York: Panthon.
inal nomnlatur, th
Singh' R. S., and . B. Krima, ds. 2000. ullutior Gtis
fr-lmrlluls to gnri nam plus a m.
orphollg. amridg: amridg Univrsity Prss.
Singh, R. S., nd . K. Uynoyama, ds. 204. Th ullutio of Popultitl and Homo spies (
Billog. amridg: amridg Univrsity Prss. -I.B. ordr to fit th thous
arviations, omitt
l. H inludd itat
liographi guid or in
Linnaus,arl (or arl von Linn) (|77_|77s)
Linnaus,s systm f
Born and raisd in Swdn, arl Linnaus was passionataout otany from sophial foundations
his youth. By th tim h wnt to Holland tn |75 to finish his mdial d- larnd aout rnt
gr, h possssdan imprssivknowldg of th known kinds of plants and stamn and pistil, as .
animals. Thr h was hird to manag nd atalog th hug privat mu- vntd lasssasdt
sum' Zoo' hrarium, and otanial grdn of Gorg lifford' a walthy asd on dtails of th
mrhant. By th tim LinnaLrsrturnd to Swdn in 1'78,h had pu. asd on only a fw
lishd numrous works, all in Latin, that st out th ruls, framwork' and otanists ould us (
atalogs of ll th plant gnra thn known. H was a foundr and th first forward to th day v
prsidnt of th Swdish Aadmy of Sins.In |741 h was hird y th natural ons. At th l.
Univrsity of Uppsala, whr h taught for th nt 3 yars. rting a of faturs' inluding
dominating influn upon natural history (otany and zoology), h at- sion of th whol pla
tratd studnts from ross urop, som of whom h snt on ollting a onfusing mixtur '
trips ovrsas.H hangd lttrs with distant naturalists who snt him spis.In his lassifi
spimns.Most important, h frquntly produd nw ditions of his ata- vr smd to work
logs of plant and animals: Sstem Nature (12 ditions twn 1735 and insts' and fins for fi
1768)' Gener Pltrm (six editions from |77 to 1764), and Speis naus aimd at th lo1
Pltctrm (thrditions from 175 to 1764). Ths works providd natu- terms gus and s1l
ralists of all nations with on ntral, wll-organizd rgistrof nams. Lin- ill-foundd laim tha
naus'satalogs wr th sarh nginsof thir day. his aims and produ
Linnaus workd in th ontxt of th worldwid olltion of natural his- At th start of his <
tory spimns that aompanid th ploration, trad, nd olonization in latr yars h saw
arrid out y uropan govrnmntsand ntrprnurs.His partiular tal- think that God had
nts, whih inludd unloval traits lik gotism and an osssion with arising latr. ost of
making lists, quipped him to produ an pandal rgistrof th divrsity Baus visil varia
of living things. Although h ptd th nt gnration ro omplt th th ntir rang is n
projt, iologistsin latr nturis,inluding our own' ontinu to nountr sult of nourging
spisnot prviously dsrid.Th projt of making an invntory of lif is spis.Linnaus ha
still unfinishd. his studnts to study
Linnaus's systm ws stritly hirarhial, gouping similar spisinto a at what dt diffrn
gnus' similar gnra into an ordr' ordrs into a lass, and lasss into a alan of natur.
kingdom. Latr taonomists addd familis (low ordr) and phyla (low Th most rmark
kingdom), as wll as suordrs' suprordrs,and th lik; this pandd the Was that th strutur
hirarhy without hanging its prinipls. Linnaus gan th onvnint to ongnial to th
prati of attahing a propr nam to ah individual group, using a tradi- groups. Th old ida
tional on whr possil (suh as Avs and Piss' Latin for ird and fish, did th prdition
rsptivly)ut oining a nW on if ndd (suh as mmali for th lss Baptist Lamark th;
 L'l|1,7us d/

ur Ges: Bioltlg1,, that inludd whals and humans). H also introdud what is alld inom-
). ial nonrnlatur' that is, always using two words to nam spis, th
Gtis from tlluls tl-l gnrinam plus a rnodifir, suh as is lupus (wolf), is fmilirls (dog),
'f.,ii,,,,,, and I7oml spies (human). (urrnt prati is to italiiz suh nams.) In
o|.Popultit
-J.ts. odr to fit th thousands of spis into mangal volums, h invntd
rviations,omittd illustrations,and mad dsriptions as short s possi-
l. H inludd itations of othr authors' so his works funtiond as a bi-
liographi guid or irrdx to th sintifilitratur.
)7-r778) Linns's sysrm fo flowring plants stimultd disussion of th philo-
ssionataout otany from sophial fundations f lassifiation. At th start of his arr h had
35 to finish his mdial d- larnd aout rnt thoris intrprting th ntral parts of a flowr, th
: known kinds of plants and stamn and pistil, as organs of mal and frlsul rprodution. H in-
talog th hug privat mu- vntd lasssasd on th numr and arrangnrntof stalns and ordrs
Gorg lifford, a walthy basd on dtails of th pistil or th fruit. Ths groups Wr artifiial, ing
d ni n 1 7 3 8 , h h a d p u _ basd on only a fw hartristis,ut thy gav ky tht vn inprt
: th uls' framwlrk, and otanistsould us (as long as th spimn was in flwr). Linnaus lookd
ws foundr and th first forwrd to th day whn ths artifiial groups <luld rplad y mor
7741 h was hird y the natural ons. At th lvl of gr-rra
and spis,Linnaus usd a widr rang
n t . ] 3 y a s . t i n g of faturs,inlLrdingth lvs and sds,and llowd his gnral imps-
ltany and zoology), h at- sion of tlr whol plant to affthis judgmnt.His ..sxualsystm'' Was thus
vhom h snt on o|lting a onfusing mixtur of artifiial lasss and ordrs with natrrral gnra and
tt nturlistsrvho snt him spis'In lris lassifiationof animals, he usd variolts harateristis,wat-
: dn w d i t i o n s f h i s a t - vr snrdto Work for ah group' suh as tth for marnmals, wings fo
ditions twn 175 and insts,and fins for fishs.Mny othrwis rlial historians stat that Lin-
77 to |764I' and Spcis naus aimd at th logial dfinitions of nrdival ..Aristotlians'''ut th old
hsworks providd natu- terms gt|us and spcies furrtiond vry diffrntly trom Aristotl's, nd th
rizdrgistrof nams. Lin- ill-foundd laim tht Linnaus Was an ssntialistadly misrprsntsoth
du. his ims and produrs.
id olItinf ntal his- At th start of his rr Linnus insistd that spisannot hang, ut
o n , t r a d , n d o l o n i z a t i o n in latr yars h saw vidn of hyridism and lnutation, whih ld lrim to
prnurs.His partiular tal- think that God had ratd only on spispr gnus' with th othr spis
ism and an osssion with aising lat.ost of his followrs prfrrdto stik with t fixity f spis.
l a l r g i s t or f h d i v r s i r y Baus visil variation within spis is oftn vry slight, speially whn
gnrationto mplt th th ntir rang is not sampld' th assumption of fiity had th positiv r-
.own, otltinuto nountr sult of nouraging th disovry and dsription of prviously unknown
laking an invntory of lif is spis.Linnaus had a livly intrstin th dyrranrisof n:rtu,nlurgirrg
his studrrtsto study whih kinds of plants various animals prfrrd to at'
luping similar spisinto a at wht dat diffrnt migrating irds rturnd, and how insts fit into th
l lss, nd lsss into a balan of natur.
w ordr) and phyla (below Th most rmarkal dvlopmnt toward th nd of Linnaus's arr
l th lik; this prndd th was that th strutur h had imposd in his lassifiationsmdvry oftn
l u s g n t h o n v n i n t to b ongnil to th pattrns of similaritis and gaps of pprntly natural
vidul group' using a tradi- groups. Th old idzrof a hain of irrg,a linar sl,did not fit' ut nithr
s, Latin for ird and fish, did th prdition of Gorgs-Louis Llr, omt d Brrffon and Jan-
:has anrmlia for th lass Baptist l,mark that onntions run irr all dirtions. Instad, pattrn of
Liuig ossl/-.

lumps, sts Within sts! smd natur's own. This unptd and gradually
mrging pitur Was th gat phnomnon lft to Linnaus's hirs. Unlik
Lamark's thory, whih grW Out of th l,rain of eing, harls Darwin's
thory dirtly addrssd th hirrhial pattrn of Linnaan taonomy by
positing a high rt of xtintion and a prinipl lf divrgIr.

BlLIoGRAPHY

B]unt, W. 197|. Th omplet Ntrlist: A Life of Liaes. Nw York: Viking Prss.

F.ringsm1,r,T., d. 1994. Litts: Th n d is Work. anton' A: Sin
istory PuliatiIrs.
L a r s o n ,I . L . 1 9 7 \ ' R s l n d p e r i : T h R 1 l r s e t t i l n N
o f t u r l o r d e r i
th Work oi rl ullt Li. Brkl,v:Univrsity of :rliforni Prss.
Linnus, . |713. Gater pltrum, ror'rmql| hrters tttrIs,sudl
umrum, figurm, situm, i prUprti()nm omium frutifitirlnis pdrtium.
ditio sud, llillibus pltrut Glliis louplett. Pris: ihl
Anronius David.
175. Spies pltrum: hibts plts rit ogits, D gr Th duk-billdplatypL
rlts, um tliffrtiis spis, ltiibus triuilibus, slmis sltis, suh odd org'nisms'th
lois tlibus, sdum sstm sul digsts,2 vols. Inrrdution y and groups of spis'r
\f. T. Starn. London: Ray Soity. alld living fossils'wil
|758. Sstm Lttrp| rg tri turtl, sudum lsss, rlrdis' ( D a r w i n1 8 - 5 9 , 4 8 6 ) .
g,ler' spis, ul hrteribus, diffrtiis, srlmis, /tlls. London:
British Musum (N:1turalHistory).
Nli.illr-Will,S. 2001. Ga dns of rdis.d.tt,or25: 49-54.
.!sit.
ro jt l,innaus. http://www.1 8.rutgs.du/pr/l/proj.l in.html.
.!(inso' Living rlativsof a
lvl. . 2006. Linnaus's iolog was not ssntialist.ls of the issouri
B<ltllilGrd 93, no. |: 2-7. -NI'.\X/. tint, ar osionall
ing fossils' spiall
ths is th olant
Living fossils l i o n y a r s g o ) 'w a s d
Living fossils r dfind infomally; th trm dos not hav a pris thni- fossil spisrprsn
al dfinition' Living anilals nd plarrtstat lrv hrrgdvry littl irr forrn vonin (365 rrrillionyt
and havior' omprd with their rmot anstors,ar said to living fos- l h l t v l n g o t e n t
sils. Also, living rlits of groups that wr muh mor divrs far k in th and a ioint within its
gologi past may alld living fossils. Bausvolutionary hrngtr-rds organs and ys froln
to onntratdin pisods of rapid divrsifiationarly in th xpansion South Afria in 1938
of nw groups of organisms, thsatgoriSlrgly ovrlaP. sin n aught ofi
hals Darwin first applid th ontraditory xprssion ..living fossil'' to nar Madagasar. R
the platypus (s figur) and tlre lungfish' as wll as Pllptrs, th Afrian latd spis has
ihirs and thir rlativ,th rdfislr.Ths nimls all rt;rinprimitiv har- lbs, in Indonsia
atristis that show thm to mor or lss intrmdiat twn typial Ltimri has not t
mmmals and rptils' or mjor groups of living orry fishs, as Darwin span of th spisL.
notd. Th platypus lys ggs' ut sukls its young. It has hair nd is warm modst wys from irs
loodd, ut it hs a lss onstnt ody tmpratuthan typial rmmals. to th gnus rrlpL
arlir naturalists,wlro saw th animal wold as fid st of sparatlyr. tis of Ltitri that
atd ..typs,',rgardd ths ratursas anomalis.To Darwin, and now to gnus-a lustr of l
us' thS living fossils ar th last lodifid dsndantsof rmrt'xtil.lt Thr strrsto r
anstors.Thos anstors'whn thy livd, wr ritial links in th ranh- small groups of los
ing volutionar histor),of highr anirrralsand plants. littl hang irr frm. r

his unxptd and gradually
[t to Linnaus's hirs. Unlik
r of ing, harls Darwin's
:n of Linnaan taxonomy
of divrgn.
es.Nw York: Viking Prss.
s Work. anton, A: Sin
resttirlof NturI rder i
y of alifornia Prss.
Lrtrs turls, s udum
ium frt ctiois prtium.
|pltttl.Paris: ihal
: rit cogits, AD gr
,,iIibus,somis sItts,
ras.2 vols. Introdution y
, sudum lsss,ordins,
:mis' /,lis.
London
r 25 49-54.
1u/pr/l/pro j.lin.html.
. lsof th issori
n t i l i s t
-.P,w.
os not hav a pris thni-
v hangd vry littl in form
jtors' r said to living fs-
h mor divrs far ak in th
ts volutionary hang tnds
a t i o n l y i n t h x p a n s i o n
rgly ovrlap.
y prssion ..living fossil'' to
|| as Polptrus, t|1eAfrian
mals a1lrtain primitiv har-
intrmdiat twn typial
ving ony fishs' as Darwin
ung.It has hair and is warm
aturthan typial mammals.
s a fid st of sparatly r-
alis.To Darwin, and now to
s n d n tosf r m o t , t i n t
:ritillinks in th ranh-
plants.
Liuig Fossl/s 689
y
Th duk-illdplatypus'th mmml tlrat lays ggs.Darwin was fsinatdy
..Spis
suh odd organisms,thirrkingthm highly signifiantfor his purposs.
and groups of spis,whih r lld arant'and whih may fnifully
lldliving fossils,will aid us in flrminga piturof th anirrtflrmsof lif''
( D a r w i n1 8 5 9 , 4 8 6 ) .
Living rlativsof animals long rognizdas flssils,ut thought to -
tint' ar oasionally disovrd.Thy ar most apt tl rfrrd to as liv-
ing fossils, spially in th popularizatitln of sin.Th most familiar of
ths is th olanth.Th first fossil olaanth, of Prmian ag (255 mil-
lion yars ago), was dsridy Louis Agassiz in 1839. Today, 70 or mor
fossil speis rprsnting 27 genera ar known, ranging from th Lat D-
vonian (365 million yars ago) to th l-at rtaous (80 miliion yars ago).
Th living olaanth' Ltimri, rtains haratristilim-lik stalkd fins
and a joint within its rlativly primitiv brin:rsthat sparatsth nasal
organs and ys from th ars nd rain. I'timri was first disovrd off
South Afria in 198. Many additional spimnstlf th sm spishav
sin bn aught off southastAfria and adjant t() th omoro Islands,
nar Madagasar. Rntly, on spimn of a dif{rntut vry losly r-
latd spishas n found on th othr sid of th Indian Oan, off th
lbs,in Indonsia.
Ltimri has not n found in th fossil rord, so th vllutionarytim
span of th spis Ltimri hlm is unknown. It diffrs in only vry
modst ways from its narstrlativ' a Lat rtaousolanth assignd
to th gnus ropom. Stritly spaking,it is th onsrvtivharatris-
tis of Ltieri that ar unquivoally living fossils, and not th spisor
gnus-a lustr of losly rlatd spis-in whih thy our.
Thr smsto nl singl planation for th long-lntinudsurvivl of
small groups of losly rlatd volving lirrags,with limitd spiatin and
littl hang in form. Arthur onan Doyl imagind dirrosaursnd ptrodatyls
690 Liuig Fossils

surviving in his L;sr World' asd in part on his knowldg of th inassil t u r d . A s i n r i l a r i

Roraima Pltau' on th bordr of Vnzula and Guyana. Som living fossils may b th lldst livir
do our in rmot, isolatd rgions, whr thy hav voidd intration yars old ar morphtl
with nwly volvd omptitors. Th platypus and its spiny, hdghog-likrl- spis' Gigko bilb,
tiv, th hidna, liv only in Australia. Sphrrodontids,rlativly unspial- that it rmitrs ullrt
izd anstors of modrn lizards, wr modratly divrs and widly th tr frn_lik a
distriutd around th world during th arl sozoi. Today' two speis Yangtz Rivr asilr i
of tuatara' thir only living los rlativs, ar rstritd to small islands off tolran of nvirrnm
Nw Zaland. strs.
In ontrast, th few living spies of |'lutils and th hossho ras hav l h d p ( ) n l ( )
i n

wid gogrphi distriuttons. Nutilus poryilius, th mst ommon and r-lnifotrrand tlnstn|
varial of thsphalopods with lganthmdshlls,rangs from th Nonthlss, it has m.
Grat Barrir Rf to Nw Guina, th Philippins, and oani arhiplagos lLrsk, Neoplli, wit
as far out in th Paifi as Fiji. Fiv distint spisof Nutilus ar known. othr organs, is a d
Typially, ths animals liv at dpths of 100 to 400 mtrs or mor' on during th arly Pal
stplysloping sa ottors,los in gainst oral rfs.opulations living o{f among ivlvs at al:
isolatd oeani islands display limitd variation. Som onstitut distint largly n displad
spis,whil othrs ar rgardd as loal suspisof Ntilus pompilius' fdillg adaptatilns.A
This has promptd th rnt infrn that Ntillts is urrntly panding thir skltons from g
its divrsity, in a modst burst of spiation.To put this in ontt' not that dpths, low th zor-r
th spisand gnra of anint lozoi nautiloids hav n assigndto sort still our in slral
ovr 125 familis, with an traordinar varity of slill shpesand mods of rrrillirryars :rgo)rrl1
lif. Th prolifi divrsity of nautiloids was rpatdlyut ak in mjor Pa- ar th stalkd hino
lozoi tintion vnts.A furthr shrp rdution ourrd nar th nd f prominnt of all ;rnr
th Triassi (210 million yars ago). Sin tl.ln,th group has dwindld to spis of stalkd rin
th singl gnus that survivs today. n d a t a y s s a ld p t h s
Th horssho ras hav nvr n vry divrs, thoughout thir long and th olaanth,o
history. Fortns not dissimilar to th living animals first appard during th displmnts towrd
S i l u r i a n ( a p p r o i m a t e | 4 2 5 m i l l i n y r s g o ) . H o r s s h o r a s ' x t r n a l faun.r and tlrn tlrs
skltons ar not as asily prsrvd as mollusk shlls, so thir fossil rord i r l n e s h o r s t r i n g 5
may b lss omplt. Owvr, no mor than ight gnra an urrntly T varity of irtl
infrrd to hav livd t any on tim. Thr gnra ar living today. that thr is nl on
On fmiliar spis,Limlus polphmus, is aundant and widly distri- plas irr whilr to n
utd, from }ylain to Yuatan in th wstrn Atlanti. Irr th Irrdo-Paifi, tion. Living fossils do
two spis rprsnting two diffrnt gnra ang from th Bay of Bngal th ft that thy tn
to th Philippins' and third spis rangs from Borno to Japan. Lik major groups to whiL
Limuls polphemus, this last spis has a larg limati rang. It is also has bn a struln (
tolrant of varyirrg salinity and othr nvironmntal varials. In this r- spis,grotrps of v
spt' th horssho rabs ar quit unlik Nutilus, whih is tmpatur- from th stal dpt
snsitiv nd annot surviv in th wrm surfa watrs of th tropis, u i l r l .T r i l o i
rm;.rin
whr it livs. ut thy finally suu
Among living plants, th spor-aing1utnosssand horstails ar r. Thi |ong Jlin t
lits of groups tht dominatd vgtation of th late Palozoi. Both ar from hangs in omt
widly distriutd gogrphially. Th horstils, also known as souring Chn, otrldda
rushs, ar limitd to aout 30 spis of a singl gnus. Typially, thy ar thir ultimat dmis
oppotunisti olonists of wt slops and rivrbanks that ar frqr-tntly dis- ris and futLlrprosp

(nowldgof th inssil
l Guyarr.Sorliving fossils
y hav avoidd intrtion
d its spiny, hdghog-lik rl_
dontids,lativlyul-lspial-
ratl divrs zrnd widly
,lsozoi.oday. two spis
.stritdto small islands off
rnd th horsshoras hav
'lius'
te most ommon and
rrdshlls, rangs from the
ts,and oani arhiplags
isof Nutilus ar known'
to 400 mtrs or mor' on
rl rfs.Ppulations living off
on. Som onstitut distint
isof Nlltilus pompilius'
utilus is urrntly panding
put this in ontt, not that
tiloids hav n assignd to
of shell shirpsnd mods of
:atdlyut ak in major -
tion ourrd nar th nd of
l, th group hs dwindld to
ivrs'thrughout thir long
rals first appard during th
,).Horsslro ras' xtrnal
< shlls,s thir fossil rord
n ight gnra an urrntly
lr gna ar living today.
rundarrtand widly distri-
Atlanti. In th Indo-Paifi,
.angfrom th Bay of Bngl
from Borno to Japan. I-ik
rrglimati rang. It is also
Lmntalvarials. In this r-
tttillts' whic is trprature-
rfa watrs of th opis,
mosssand horstails ar r-
th lat Palozoi. Both ar
i l s .a I s o k n o w n s s o u i n g
gl gnus.Typially, thy ar
n k sr h t r f r q u n l yd i s -
Liuig Fossl/s 6L)1
turbd. A similar hitat was formrly prfrrd y th gingko tr, whih
may th oldst living spis' Fssil lvs nd wood up to 100 rniliion
yars old ar morphologially indistinguishal from th singl surviving
spis, Gingko bilrlb, of this anint group. This plant is so distintiv
tlrt it rrrrainsunrtin whthr it is nror losly rlatd to onifrs o to
th tr frn_lik yads. Orr widsplrd,th gingko survivd only in th
Yangtze Rivr sin in hin until its domstiation. Now, its auty and
tolran tlf nvironmntl insults hav rought th gingko to many ity
stts.
Th dp ;anfloor is nithr as low in divrsity of arrimal lif rror as
uniform and onstnt in its physil onditions as Was on supposd.
Northlss,it lras nlor than its slrar of living fossils. lr rping ml-
lusk, Neoplli, wlt its ap_shapd shll and srial sts of musls and
othr organs, is a dp-sarlit of two familis that livd irr shllow watr
during th early Pirlozoi. Protoranhs with simpl gills prdominat
among ivalvs irt ayssal dpths; orr th ontinntal slrlvs thy hav
largly n displad y mor advand forms with vrid urrowing and
fdingadptations' Arong singl-lldforaminifra, only thos that uild
thir skltons fom grairrs of pristing sdir-rrnt an liv at th grtst
dpths, low th zon t whih aliurrrronat dissolvs.Forams of this
sort still ur in slrallow Watr haitats, ut in ambrian tim (aor"rt510
million ys ago) tlrrl1,this prinritiv cr()up was prsnt.ost striking of all
ar th stalkd lrindrmspopularly known as sa lilis. z\mong tlr most
prominnt of all nimals in shallow-Watr Palozoi faLrnas, about 80
spisof stalkd inoids suviv today, worldwid' on Ontinntal slops
and at yssal dpths. Ths distriutions' togth witlr tlros of Nutilus
and th olaanth.orroorat vidnfrom th fossil rord of sussiv
displalntstoward dp Watr' first of dsndants of th amrian
fnand thn th<lsf a lirtr lozoi fauna, ils nW orrlnruniisvolvd
in narshosttin8s.
Th varity of iumstansin whih living fossils hav survivd indiats
tht thr is no on aus of tlris phnomnon. Tlrr ar lan ways and
plasin whih to marginlizd witlrtlut ing liminatlby naturl sl-
tion. Living fossils do not ollstitut a spial ast of organisms' pt for
th ft that thy tnd to mong tl.rlst spializd mmbrs of th
maior groups tl '"vl.rihthy long. At zrnygivrr tim in arth's histor,v,thr
has brra sptrum of living things: rpidly volving groups of phmral
spis'goups of vragstaility and tnur, anl groups that dpart littl
from th stal .ldaptations lf thir anstors, as long s thir nrods of lif
rmin vil. Triloits Wr th prrninntliving fossils of Prmian im,
tthy finally suumd in th grat tintion irt th nd of th Palozoi.
Thir long dlin fom urant divsity in th mid-Palozoi rsultd
from lrangsin ommunit strutur tht ar not yt urrdrstoodin dtail.
han, outmodd daptation, and omptition all surl ontriutd to
thir ultimat dmis. Tody's living fossils hav similar volutionary lristo-
ris rrd futur prospts.
692 Lorez

|BLloGRAPHY at birds in prtiular, l

not so muh y aquir.
Darwin, ' 1859.o the rigi lfSpeles.Lr-rdon: John urra,v.
ldrdg,N., and S. . Stanly,ds. 1984. Liuitlg Flssils.Nw York: Sringr- havior pattrns, uilt
..fixd
Vrlag. valu. For ths
Fory,P. l998. A lromfrlmhomfor olaanths. Ntur395:319_320. lasd'' y stimuli m
Royr,D. L., L.J. iky,and S. L. !fling.200.ologilonsrvatism in th In th 1930s h onstrr
..livingfossil'' Cllzgl. Plobilllg1'
29: 84_14.
rlsing mhanisrs
Shopf,'J.. 1984.Ratsf volutionand th notior-r of ..livingfssils,''nul
Reuiulof rth nd PltrSis12 245-292. trns. At th sam tim
Thomson'K. S. 1991.Liuig Flssil:hStorof th llazf/:. Nw York: ing.'' LI found tht yo
Nor.ton. -R.D... rogniz mmrs of
formation at a rif, l
printd'. upon th ob
Lornz, Konrad Zachartas (1903-1989) thir instintiv hv
Konrad L<lrnz,th son of a rih and distinguishdVinns orthopdi sur- x m p l ' i s p o s dt .
gon' grw up witlr a passiorrfor aising ninrals. Latr, whn h bam f- th imprinting prid,
mous for his studis of nimal havior, h laimd that his matur sintifi than its own spis.
ptiswr ssntiallyontinuous with pratislr dvlopdin his youth With his hrismat
as an animal lov' He isistd futhrmorthat y raring and osrving an- th naturl haviora
imals undr sminatural onditions h was al to larn things aout animal b o l d t h o r i z i n g ,a d l
havior that wr not radily assilto laoratory sintistsor fild nat- mny rrttits to th n
urlists. his popular ook. o,
Following his fathr's wishs, l-ornz nrolld in 192 as a mdial studnt mad rlir as bi
at th Sond Anatomil lr-rstitutof th Univsity of Vinna, vntully most for attrating th
rning his D thr in 1928.He thrupon nrolld at th univrsity's Zoo- i m p o r t a n o f s t u d v i n
logial Institut, from whih h rivd a PhD in 1933. anwhil' at his pri- f o u n d i n g t h s i n o
vat hom rsah station in th villg f Alnrg, los to Vinn, h had i n l r " r d i n tgh a w a d l
gun making on tam' fr-flying irds th osrvations that would lrlp pro- whih h shard with
pl him to prominn in th 190s, first among Grman ornithologists and BIBLIOGRAPH
tlrn amorrg animl psylrologists.His rsarh provided th osrvtional nd
B u r k h a r d R, . W . ' J r . 2 0
onptual foundations of th nw sin of thology, th iologial study of
d th Ftludig tl
havior,an ntprisin whih Lrnz's work ws bly supportd and om- Lrnz' K. Z. l94 l. Vrl
plmntd y th ontriutions of Niko Tinbrgn, Duth r.raturalist. Heirotb. rg:izun
Lornz's initial ontriution to th volutionry undrstanding of living 1966. ggr
thir-rgswas his promotion of th ida that th rrrthodsrfompartiv natomy !7orld.
ould applid to animal bhvior pattrns just as fftivly as thy ould \)7Q-|91|..\1
vols. Cmridg,
applid to nimal struturs. H maintind that orparing th innat hav-
N i s h t t , . | 9 7 6 . K t t r
ior pttns of diffrnt spis was soffrtimsvn mor virluabl than om-
paring thir physial hartrs whn it ar to ronstruting phylognis.
His most important puliation in this rgd Was a mior monograph in 1941
omparilginnat lrhaviorPattrnsin duks and gs.H ltr put his road, Luy (ustrIopi
omparativ prsptivsto us in thinking mor gnrally about volutionary
pistmology, promoting th viw that th way th human -rindapprhnds In vry sintifi fil<
th world is itslf th produt of a Iong volutionary pross. thoughs that ar ndt
In addition to his fforts to undrstand animal and human havior in vo- all ar suprsddor
lutinary trms, I,<lrenzsoLrghtto nr:rksnsof th physiologial ausation dpr insights ar m
and soial funtiorr of ..innat''' spis-spifihavio pattrns. Looking Thr is no dout tha
 Lu 69

at irds in partiular, he insistd that thy ar adaptd to thir nvironmnts

nt so muh y aquird knowldg as by highly diffrntiatd instintiv -
Jhn urray.
ssl/s.Nw York: Springr- havior pattrns' uilt up ovr tim y volution as a rsult of thir survival
..fixd motor pattrns'' to fftiv, thy ndd to ..r-
valu. For ths
\'ltur95: 19-20. lasd'' y stimuli manating from ppropriat ojts in thir nvironmnt.
logial nsrvatism in th ..rlass,''..innat
In th 1930s h onstrutda thory of instint faturing
rlasing mhanisms,'' ation-spifi nrgis' and innat, fixd motor pat-
-ionof ..livingfossils.''Aul ..imprint-
)o ) trns. At th sam tim, h alld attntion to th phnomnon of
oltb.
Nw York: ing.'' H found that young irds suh as jakdaws of gsdo not instintivly
-R.D.K.. rogniz mmrs of thir own spis ut instad normally aquir this in'
..im-
formation at arief , ritial priod of thir arly dvlopmnt. Thy ar
printd'' upon th ojt that will susquntly srv to rlas rtain of
)) thir instintiv havior pattrns. Howvr, if a young grylag gosling, for
l d V i n n s o t h o p d i s u r - xampl, is posd to a human ing instad of mothr grylag goos in
s. Latr' whn h am fa- th imprinting priod, th gosling will om imprintd upon humans rathr
nd that his matur sintifi than its own spis.
s h dvlopd in his youth With his hrismati, xurant prsonality, his xtnsiv knowldg of
y raringand obsrving an_ th natural havioral rprtoirs of a host of diffrnt animal spis, his
to larn things aout animal old thorizing, and his skills as a popularizr, Lornz was al to attrat
)ratorysintistsor fild nat- many rruits to th nw sin of thology. ontrovrsial in th 1960s for
his popular ook, Oz ggrssio' and rurrntly ritiizd for statmnts h
i |92 as a mdial studnt mad arlir as a iolgist undr th Third Rih, Lornz is rmmrd
',rsity of Vinna' vntually most for attrating th attntion of zoologists and psyhologists alik to th
:olld at th univrsit,s Zoo- importan of studying animal havior from a iologial prsptiv and for
r l 9 . M a n w h i l a. t h i s p r i - founding th sin of thology. His work was rognizd y many honors,
Lrg,ls to Vinna, h had inluding th award of th 1973 Nobl Priz for Physiology or Mdiin,
rvations that would hlp pro- whih h shard with Karl von Frish and Niko Tinbrgn.
g Grman ornithologists and BIBLIOGRAPHY
rovidd th osrvational and
'ology, th iologial study of Burkhardt,R. W., Jr. 2005. Pttrsof Bhuilr:Kord Lorz, Niko Tibrg,
d th Foudingof tbllog.higo:Univrsityof hiago Prss.
was ly supportdand om- an Anatidn.In Festshrift.
Lorenz,K. Z. 1941r.VrglihndBwgungsstudin
)n' a Duth naturalist. Hiroth.rg:inzungsand 3. lorl filr rithlllgi89: 194-29.
rary undrstanding of living |966. ggrssio,. K. lfilson' trans.Nw York: Hrourt, Brand
hods of omparativ anatomy !7orld.
as fftivlyas thy ould 1970-197I. StL|disin iml d Hum Behuiour.R. artin' trns.2
vols.embridg,MA: HarvardUnivrsityPrss.
. omparing th innat hav-
Nistt,^. 1976. Kord Lorz. Nw York: Hrourt BrJovanovih.
vn mor valual than om-
R.vB.
o ronstrutingphylognis.
s a major monograph rn 1941
gs.H latr put his road,
. g n r l l ya o u t v o l u t i o n a r
Luy (AotrIopithecusfrsis)
th human mind apprhnds In vry sintifi fild of ndavor thr ar rtain disovris and rak-
Larfpross. throughs that ar nduring and ahiv somwhat of an ioni natur. Narly
l and humn hvior in vl- all ar suprsdd or signifiantly altrd whn nw finds om to light or
' f t h p h y s i o I o g i al u s t i o n dpr insights ar mad' ut th or innovation or disovry is vrlasting.
: haviorpattrns. Looking Thr is no dout that onstruts lik th Big Bang, th thory of volution,
694 I,

th thory of gravitation' ontinntal dift, th lws of inhritan' and many

othrs hv n nhand or modifid from thir original manifstations,
ut th ntral ida is still thr and th onpt ontinus to known y its
original itration.
Looking ak on my arr as a paloanthropologist-somon who stud-
is human origins-I had oasion ln 1974, in th rmot dsrt rgion of
thiopia known as th Afar Triangl' to mak on of thos nduring disov-
ris. Sh was a 3.2 million-yar-old human anstor who has om pr-
hps th most widly lbratd disovry of a fossil human of th twntith
ntury. Known y th afftionat nam of Luy (namd aftr th Batls
song ..Luy in th Sky with Diamonds''1 s figur), tis 40" omplt skl-
ton (not inluding th ons of th hands and ft) pitomizs our fossil an-
stry. Luy hs om a touhston for omparing and valuating all nw
fossil humn finds. vn for thos who hav diffiulty rmmring hr si-
ntifi name, strlopitheus frsis, ringing up th nam Luy rings a
ll for vryon:..Oh, yah, that anint sklton from Afria!''
On of th ssntialqustions tht all humans ask is' ..!hrdid w om
from?'' Thr ar two ways to answr this qustion. On is to vok a r-
ation story of how W wr onivd in som sort of suprnatural mannr.
Suh planations that ar asd on lif diffr from soity to soity' and
ptanis prmisd on providing som sort of spial vnt or suprnatu-
ral aus, th vidnefor whih is invisil and availal only to thos who
hav faith.
On th othr hand, paloanthropologistsus a sintifiapproah to om-
prhnd th aus, ours' and timing of human volution. ! nd not in-
vok spial xplanations or suprnatrrrlauss.Th study of human origins
is basd on osrval vidn that is availal to vryon' rgardlssof
thir blif systm.
Thr is a fossil rod for humankind urid in th arth's sdimnts.
Luy oms from a priod of tim whn our anstors lookd vry aplik,
with projting fs' small rains (on quartr th siz of a modrn humn
brain), rlativly long arms (lft ovr from tr-livingprusors),and urvd
hand and foot ons (also from our roral past).Howvr' thsanstors,
somtims alld ..ap-mn,''possssda ardinal fatur that plas thm
on th vlutionary ranh f humans rathr than aps:ipdalism.Bipdal-
ism is th ability to stand and walk on two lgs, a havior that stands in
powrful ontrast to all othr mammals, whih gt around on all fours. Ad-
ditionally' ths ..ap-mn,''in ontrast to prsnt-dayhumans, wr prul-
tural, maning thy wr inapal of making and using vn rudimntary
ston tools (thy may hav utilizd prishalraw matrials lik wood as dig-
ging stiks, ut ths itms did not fossiliz). ustrlopitheus fr
. Johanson in th Air
originally doumntd with th 1924 disovry in South Afria of th
..Taung
hild" (th skull of an immatttre ustrlopithus frius| y th
lat Raymond Dart, th di Was ast that Afri-or as harls Darwin r-
frrd to it' th Dak ontinnt-Was th original hmlnd for humanity. In
th dad aftr th puliation of Darwin's ontrovrsial,lasting, and pro-
 L 695

: lws of inhritan'and many

r t h i o r i g i n a Im n i f s t a t i o n s ,
pt ontinus to known by its

opologist-somon who stud.

in th rmot dsrt rgion of
on of thos nduring disov-
anstorwho has om pr-
a fossil human of th twntith
Luy (namd aftr th Batls
gur),this 40% omplt skl-
l ft)pitomizsour fossil an-
nparing and valuting all nw
diffiulty rmmring hr si.
ging up th nam Luy rings a
lton from Afria!''
...W.hrdid w orn
tns ask is,
ustion.on is to vok a r-
r sort of suprnatural mannr.
fr from soity to soity, and
.t of spial vnt or suprnatu-
Lndavailalonly to thos who

s a sintifi approah to om-

nan volution. ./nd not in-
s s . h s t u d yo f h u m n o r i g i n s
al to vryon' rgardlss of

urid in th arth's sdimnts.

anstorslookd vry aplik,
tr th siz of a modrn human
-livingprursors),and urvd
)ast).{owvr'ths anstors'
rdinal fatur that plas thm
than aps:ipdalism. Bipdal-
lgs, a havior that stands in
h gt around on all fours. Ad-
snt-dayhumans, wr prul-
n g n d u s i n g v n r u d i m n t a r y
raw matrialslik wood as dig- ..Luy,''ws disovrdby Donald
ustrlopithcusfrsis.This spimn,
. Johansonin th Afar trianglof l,thiopia in 1974.
sovry in South Afria of th
strlopithusfricus) y th
.ria-o s Charls Darwin -

;inal homland for humanity. In

ontrovrsial, lasting, and pro-
696 Luc

foundly insightful volum' o th rigin of Specis(1859)' it amppar- Humans, apal of

nt to sholars that hr was an ida so powrful that it was apal of lui- yars ago. Somtims l
dating why spis look and at thy way thy do. Furthrmor, it plaind o r i g i n a l l y v o I v di n A l
why thr is a smingly ndlss divrsity of lif on arth. to th dmis of suh
Darwin knw that living raturs look similar to ah othr aus of matly to th popling
\hv om from
ommon anstry. H livd thy ar ah uniquly distint austh
pross of natural sltion has raftd thm to at in mannrs rflting finsus as humn-ip
thir partiular mods of lif. Osrvations y Darwin (1s71) and his los s i z , o d i so f m o d r n
frind Thomas Hnry Huly (1s63) ld thm to mak a tstalprognosti- apparan in Afria. \
ation. Th prmis was that if hmans and th Afrin aps look alik, thy ings with a ommon
mst hav had a ommon anstor' and th diffrns ar rlatd t how largd, nlightnd, an
thy bhav (apslim trs,humans walk on th ground). If th thory was o m f r o m ? . .w i I l u n i t
orrt' nd volution rflts ommon anstry as wll as dsnt with mod- futur.
ifiation (.g.,humans lost thir aility to loomot on all fours whn thy IBLloGRAPHY
am ipdal), and that th Afrian aps and humans sprang from a long.
D a r w i n , . 1 8 5 9 . t h
distant ommon ginning, the oldst tras of humanity rsid in Afria.
| 87 |. h Ds
That proposition is what th Taung hild' Luy, and thousands of othr Murray.
Afrian fossils, hav vindiatd. Our dirt anstors first mrgd in Afria. Huly, T. H. 186,uid
Th ommon anstor to th Afrian aps and humans still luds r- Norgat.
sarhrs and rsts in som unxplord and unxavtd Afrian stratum. J h a n s o nD, . . 2 0 0 4 .L u
ustrlpithsfr
Howvr' paloanthropologists now pitur Luy's spies as having livd at
465-486.
an important infletion point in human evolution. Prior to Luy, th fossil
J o h n s o nD' . . . a n dM . ,
rord is spars. Now, human anstor fossils ar oming to light at 4 million Nw York: Simonan
ears (. nmesis)' at 4.4 million yars (Ardipithus rmidus), and pr- Johanson,D. ., and B.
haps vn as far ak as 6 to 7 million ears (Shlthropus tchdsis). and Shustr.
And tru to th prditins mad y th thory of volution, ths oldr K i m |W , . H . , Y . R a k .a l
a f a r n s i sN. w Y o r k
spis ar looking mor and mor aplik, similar to th prsumd mor
Tattrsall,I., ndJ. Shw
primitiv ommon anstor.
Susqunt to Luy, in mor rnt Afrian sdimnts w se vidn of
thos faturs w think of as mor humnlik. t 2.6 million yars' ston
tools, fashiond y som unknown human hand, ar found assoiatd with Lyll,harls(179
uthrd rmains of savngd animals-mat had ntrd th human's
dit. Thn, at around 2 million yars' a nar douling in rain size (oo I n v a r i a l y m m
hbilis, I:Iomo rudolfesis, Homo rgster) is sn in largr skulls. Th mod. Golog (first dition ir
rn human ody form (long lgs and rlativly short arms) appars shortly was th first hild of an
thraftr, and th first out-of-Afria prin is doumntd in th Rpu- family, and of a Soti
li of Gorgia at 1.8 million ears (Homo erectus). only 15 yars arlir 1
Following a long priod of tim, during whih human spis migratd, h i s f o r t u n w h i l s r v i
volvd, divrsifid' and am tint, vn mor modrn humanlik an- Lyll livd in southrn I
stors suh as , tcsslr and H. heidIbrgesis (500,000 to 1 million his BA from Oxford in
yars ago) walkd th arth. Th first tantlizing glimpss of art, ngrvd took him oftn ro ol
ohr, and th spializd and highly insightful manuftur of on tools l850s, to North mri
appar in Afria som 70,000 to 100,000 yars ago' two to thr tims ar- in th 1820s, and rif
lir than in urop. Now rsarh has onfirmd th ourrn of anatomi- t th nW King's Coll
ally modrn humans (omo spis' supposdly ..wis man'') at los to mans supplmntdb
200'000 yars ago in Afria (thiopia). pially, from lturing

,ies(1859),it am appar-
rl tht it was apal of lui-
lo. Furthmor,it plind
on rth.
ilar to h othr as of
niqulydistint aus th
to at in mannrs rflting
Darwin (1871) and his los
to tk tstlprognosti-
Afriarr aps look alik, thy
iffrnsar rlatd to how
th ground).If th thory was
, as wll as dsntwith nrod-
)moton ll fours whn thy
lhumans sprang from a long-
f humanity rsid in Afria.
uy, and thousands of othr
storsfirst mrgd in Afria.
and humans still luds -
rnavatd Afrian statum.
ry'sspisas having livd at
tion. Prior to Luy, th fossil
r oming to light at 4 nrillion
dipithus rmidus), and p-
(Shlthr opus tch desis) .
ory of volution, ths oldr
imila to th prsumd mor
sdinrntsw se vidn of
e. t )-.6 million yars' ston
nd, ar found assoiatd with
at had ntrd th human's
douling in brain size (Homo
n irr largr skulls. h mod-
y short arms) appars shortly
is doumntd in th Rpu-
tus).
hih hurnan spis migratd,
mor nrodrrrhumnlik n-
rgesis (500'000 to 1 million
ling glimpss of art, ngrvd
:ul manufaturof bon tools
s go) two to thr tims ar-
d t h o u r r n o f a n t o l n i -
..wis man'') at los to
;dly
Lell 697
umans, apal of rading and writing, ntrd urop around 40,000
yars ago. Somtims known as r-agnons' thes human anstors had
originally volvd in Afria' and thir ultural and intlletual apaitis ld
to th dmis of sr.rhspis as Nandrtals (0,000 yars ago) and ulti.
matly to th popling of th ntir glob.
.W
hav om 1rom huml ginnings. ah and vry fatur that d-
fins us as human-ipdalism, tool making, nlargd rains, rdud tooth
siz, odis of modrn proportions' prhaps vn languag-first md thir
apparanin Afri. Ys, Lu hzis lft a lgay that unites us all s humn
ings with a ommon ginning in Afia, and it is my hop that this n-
largd, nlightnd, and profound insight into th qustion of ...!fhr did w
om from?'' will unit all humankind into a fruitful nd lons-livd ommon
futur.
|LIoGRAPY
Drwin' . 1859. th rigi of Sples.London:Jhn urray.
1871. Th Dstof , d Seltili Rltioll /o Sex. London:John
urray.
Huly,T. . 1863.ides trl ris Pli Nture, London:Williarnsand
Norgar.
Johirnson,D. . 2004. Luy, thirty vrslatr:An -ptrndd
viw of
Austrlopitbas frsis.Joull of nthropoklgilRsrh60' no. 4:
465-486.
Jhason,D. .' and . A. dy.1981.Lu: Th Bgiigsof Humkid.
Nlv York: Simlnnd Shustr.
Johanson,D. .' nd B. dgr. 2005. Frlm l-u1,16 Lgug,Nr,r'Yrk:Sirrron
nd Shustr.
Kinrl,!. H., Y. Rak, and D. . Johnso.204.Th Skll of Australoithus
farnsis.Nw York: Oxford UnivrsityPrss.
Tttrsall,I., and J. Shwartz.200,tict ums' Bouldr,O: !stviwPrss.
-D..J.
Lyll,harls(t797_\875)
Invariably rmmrd'vn on his gravston'as th author of Priipls of
Golog (firstditiotl in thr volunrs:1830, 1832, and 1833), harls Lyll
Ws th first hild <lfan nglish mthr' from lndd Yorkshir (Swaldal)
fan"rily,and of a Sottish fthr, lird of a larg Angus ounty statought
only 15 yars arlir y th oy's grandfathr, a farmr,s lad who had md
his fortun whil srving at sa in th Nw World' From infany, howvr,
Lyll livd in southrn ngland, always in London itslf aftr graduatingwith
his BA from oxford in 1819. xtnsiv travls, largly in gology's srvi,
took him oftn to ontinntal urop and four tims' in th 1840s and
1850s, to North Amria. H workd sporadially for a fw yars as a lawyr
in th 1820s, and rifly (181_1833) hld a profssorial hair in his sin
at th nW King's ollg, Lndon. Othrwis, h dpndd on his privat
mns supplrnndy prods from writing and, in th Unitd Stass-
pially, from lturing,sO dvotil-lghimslf almost lusivly t his sin.
698 Lll

vntually knightd, a frirrllf Prin Alrt, .1ndn advisr to thr govrn- with what h had
mnt, h ilm a prominrrt r-riltionalfigu witlr ir h<luson rly Strt, affiliations.
.s7stminstr
fmous for its mdial grnds. His finl rsting pla is in That form, as
Ay. onform gology t
Th yars' frorn mid- l 82 to mid-1826' Wr th most formtiv for th xplaind by al
viws uphld y I,yll in Priipls. arlir, as an undrgraduat,t-yll had fid y Nwtonin
attndd !7illiam Bukland.s gology lturs nd thn don fildwork with thory-a nW vr
puliatiorr in mind, whil rr-rraining unitial of oxford-taught gology. onformal' pot
By mid-1826, howvr, h was pulily gr"ring for a furrdamntaland om- a v o i d i n g d i v i s i v 'i
prhensiv altrntiv to th tahings of th )ford shooi of gology and sin fom os
to th tahingsof ParisiirrrGlrgsuvir, th shool's primary mntor (u- ould only mt i
vir 1813). In mking this shift, Lyll was aov all intgratingth Huttonian roks wr all prol
thory of th arth, as dfndd yJohn Plyfair (1802)' whom Cuvir had nd with th sam i
onsistntly opposd, with th proposls of orrstant Prvost (1823, 1825), futur' th sam so
who wirs lpposinguvir's orvrrintrprtationsof tlr Trtiry formations of insistd, ntitldto
th Pris Basin. Lyll's ngagirrgwith la,vfairin his writings arrdwith Pr6vost by nrost gologists
irr prson wr prol,vokrd y William Fitton, a sretaryt this tim, knw nough of pr
long with Lyll, of th pstigiousGologial Soity of London. Fitton, an fts. By forming
Irish-born' Sottish-taindphysiian' Ws a suppotiv frind of th Frnh- his prim amition
man and an admiring sussorto Playfir as th main writr on gology for urrnt low rput
te dibrgb Reuitu. B1, lat 1823, Fitton lrad rjtd Buklnd's vatd status ompa
Noahian liluvial gology ln thc diburgb RcL,,itL, (s lso Fittorr 1824, takn to th hig
1828), whil it rvas nlltdin th rival Ttlr1,,Anglian Qttrtrl Ruitu Sottish gntry, Lyt
y dward oplston. Coplston, a futur ishop, was Oford's potry pro- th sinthat h ]
fssor, longtim ruttor of Playfair's ritiisrrrsof his univrsity,and Buk. proSPtivprofss
land's oyhood frind (Buklnd Was vn rumord to hav hlpd litist anxity Was
oplstlnwith th writing of his rviw). Th ultimat sta
Th t]iburgh Ruictu oftr-rdfind itslf y its liral, Sottish, r/hig, transndntani
forming opplsition to Tlry, Anglian, nd Oorri:rrrhgmonis in th (a livr in Gd l
n a t i o n ' s p o l i t i s , r l i g i o r l ' l t t r s ,a n d s i n .D t r m i n i n g L y l l ' s v i w o n and govrnmntn
any issu in gology or yond an start fro.r his family's and his own matrial soul om
.shn
alignmnts with th diltburg Rui,s stl1. Lyll wrot in Jan-BaptistLam
1826-1827 on duation and sintifi institutions as wll as on gology rjtd in th arl1
for th Qurtrl Reuita, lr did so s a onsious dissidnt intrudr. It mitmnts bout m
Ws as :r lissidnt intrudr too that h tauglrt :rt King's ollg. With rord s progrs
oplstorr ils n ovrsr' King's ollg ws dsigrrdt Toy, Angli- suh as Lamrk's
n for'rndation, out-ttrr,vight to th young Univrsity ollg, London' am thn to join
whih had n foundd y liral Sottish Whigs and hdd y Lyll's progrssiv nssi
own fathr-in-law, a rothr of an diburgh Reuit,tlfoundr. Lyll's nw tory of lif on arth
r f o r m i n g a m i t i o n i n t h m i d - 1 8 2 0 s w s t o r f u t a n d r p l a w h o l s a l many adaptiv on1
th Oford shool's n<l_(]ttr.irian gllogy, jr-rstwhn that shool had ing suh ontrsts\
ahivldorninan in amridg, in th mtrop;lis,and so in the ntion. orizing as raising,
Argualy, rhrfor' tis anrititll-lis pli;rlirs th mition of a young thology and mor
man wl.rohad om to think that starting ltlt in th Oxford shool was not hildlss and d
a pl whr a liral, Sottish, Whig lawyr lik himslf should vr' g- hugly inf]untial
ologially spaking, hav gon. onvrslv, h now had to at in aord any shol of om1

'nd an advisr to th govrn-
ith a hous on Harly Strt,
i n g p, | e i s i n W s t m i n s t r
r th most formativ for th
an undrgraduat, Lyll had
nd thn don fildwork with
l of oford.taught gology.
for fundamntal and om-
ford shool of gology and
;hool'sprimary mntor (u-
a l l i n t g r t i n gt h H u t t o n i n
i r ( 1 8 0 2 ) ,w h o m u v i r h a d
n s t a n tP r v o s t( 1 8 2 3 ' 1 8 2 5 ) ,
of th Trtiary fomations of
his writings and with Pr6vost
i t n .e s r t a r ya t t h i s t i m ,
ioityof London. Fitton, an
)portivfrind of th Frnh-
Lmain writr on gology for
ln had rjtd Bukland's
ui (seealso Fitton 1824,
, Anglian Qurterl Reuietu
op' Was oxford's potry pro-
Lsof his univrsity'and Buk.
L rumord to hav hlpd
ly its lirl, Sottish, Whig,
oxonian hgmonis in th
Dtrmining Lyll's viw on
'm his family's and his own
n. Whn Lyll wot in
.rtionss wll as on gology
tsious dissidnt intrudr. It
\i7ith
3ht at King's ollg.
lsigndto a Tory, Angli-
Univrsity llg' London,
Whigs and hadd by Lyll's
Ruilu foundr. Lyll's nw
rfut and rpla wholsal
just whn that shool had
r o p t - ' l i sa.n d s o i n t h n t i o n .
as th amition of a young
n th oxford shool was not
lik himslf should vr' g-
L now had to at in aord
I,ll 699
with what h had om to appriat as his authnti familial and prsonal
affiliations.
That rform' as pounded in its fullst xtnt in Pricipls, ws (1) to
onform gology to t'eur us eidential-xplanatory idal (fftsar to
plind y auss known to ist from indpndnt vidn)xmpli-
fid y Nwtonian lstial mhnis; (2) to giv th sin a systmi
thory- nw vrsion of th aosmogonial Huttonian thory (itslf now
onformal, potntially at last, to that idal)-that Was thortial ut
avoiding divisiv, inonlusiv' osmogonil spulation;and (3) to fr th
sin from osai ilial assoiations.Ths nds, as Lyll saw thm'
ould only mt if gology prsumd that th past hangs rordd in th
roks wr all produd y uss still working in th sam irumstans
and with th sam intnsitisand so produing, in th psntnd on into th
futur, th sm sorts and sizs of ffts. It was indd a prsumptin, Lyll
insistd,ntitld to a priori prfrnovr its dnial and so wrongly rjtd
y most gologists beaus thy had prmaturly didd that thy alrady
knw nough of prsnt auss to judg thm inadqut for thos past f-
fts.By rfoming gology y giving it ths prinipls, Lyll ould ahiv
his prim amition for th sin and for himslf: to sav gology from its
urrnt low rput and vn ridiul y giving it sound foundtions and l-
vatd status omparal to th foundations and status of lstialmhanis,
takn to th highst sin of all. As th son and grandson of arrivist
Sottish gntry' Lyll worrid intnsly aout th foundations and status of
th sinthat h had mad his avowd avotion and vn, sin 1827,is
prosptiv profssion in pla of th law. Notorious fr his snory, Lyll's
litist anxity Was at on soial and ognitiv.
Th ultimat status of mankind-our filiation' dignity' and fat-raisd
transndntanxitis that, Lyll insistd, nly rligion an addrss. A dist
(a livrin God ut not in th Bil), h hld to a Divin ration' dsign,
and govrnmnt not only for ntur ut lso for man, whos immortal, im-
matrial soul oms undr God's moral law in this lif and th rrt. H saw
Jan-BaptistLamark's (1809) systm of zoology, tnsivlydisussdnd
rjtd in th arly ditions of Priiples, as dirtly thratning ths om-
mitmnts aout man. Aordingly, h rjtd any rding of th fossil
rord as a progrssion,from low lif to high, so as to disrdit any shm
suh as Lamark's that has humans produd from animal anstors.Lyll
am thn to join uvir' Lamark's othr opponnt' in rplaing any unifid
progrssivnssitatingplan with ontingnt' fitting adaptations in th his-
tory of lif on arth. harls Darwin,s thorizing rtaind this favoring of th
many adaptiv ontingnisovr th on nssitatingplan. Whil rogniz-
ing suh ontrasts with Lmark, Lyll alwys rad this nw Drwinian th-
orizing as raising' if xtndd to man' th sam old intolral thrat to
thology and morality.
hildlss and dstind not to inhrit th family stat' Lyll, although
hugly influntial ovr svral dds f gologial sin,nvr foundd
any shool of omprhnsivlyonsnsual disipls. In his gology' Darwin
I.,h

was vry muh Lyll's most loyal protg6, vn aftr dvloping lris wn dis- Lynh gan his ar
sident iology. Lyll's no-uttonin s,vstmof th arth, itiizd from th of zoopl:rrrktn as a sl
start ovr its laims for a onstant' undelinirrg thrmal onomy for th n s o t a . l n a s r i so f 1
plant, Was mor widly thought implausil from th 1850s on aftr artiulatd a omprh
William Thomson (Lord Klvin) and othrs raisd thir novl thrmodynam. orporating disparat
ial o1tionsto ths lims (Burhfild |975). Th intriat omplxity of gtis, and ulminatin
.Walla's (Lynh 1980).His os
th young harls Drwin and young Alfrd Russl agrmnts
and disagrmnts With Lyll, and with Lyll's agrmnts and disagrmnts rprodutiv sussll(
with Lamark' put th proximat and ultimat dbts of volutionary iology tional wisdom of lif-h
to Lyll yond quik-and-asy summary gnralization. Thos dts ar sking th simplst x1
mor athr thn lss dp ausof that vry omplity. put him J th nt
quartr ntury.
|BLIoGRAPY Th 1970sand 1980
Burhfild,J. D. I97 5. Lord KIuid tb g of th rth. New York: Sin ary iology: th mrg
Histor,vPulitions. uration of th fild of
uvir,G. l813. sso th.heorof the rth.R' Krr, trns.dinurgl-r: rinvntd imslf s a
\fl/.Blakwood.
talished himslf as ]
Fitton' !. H. 1824. Inquirisrsptingth gologilrlationsof th dstwn was instrumntal in .
th hlk and th PurkLimstonin th South-rrst of ngland' Als of
gnetis (.g.,Lynh an
P hillso1lh
8: 365*383.
1828.Addrssdlivrdon th nnivrsary, Fbru:rry1828. Prldigs of Qutititiu rits, l
th GerlltlgiclSoit1,of Lodo 6: 50-62. standard for th fild ol
Lamark,J.-B. 1809.Philoso1lhie zoologiqu.Paris:Dntu. voiutionry quantit.
Lyll, . 1830_183.Priiplesof Golog:Big ttmpt to pli the lizing or positiv dirt
Formr hngesi tb rth's Surfb Rfretl usesNotu i
Kondashov, Lynh was
prtiolt.3vols.London:John urray.
Lyll, K. ., d. 1881 [1970].Life, Lttersnd Jlureils of Sir hrlsLlI, Brt' dltrious mutations t.
2 vls.London:Jolrrrurray. Rpublishdin fasimil,Farnoror-rgh, U.K.: Wilfid Garil, Rinh
Grgg IntrnationlPublishrs. ati study of th ffts
Playfair,J. 1802. lllstrtit.lsrlf th Huttt'lihor oi th rtb. dinurgh: tion xtintion (Lynh t
!illiamrh. ..mutationalmltdo
th
Prvost,. 1823.D l'importand l'tuddsrpsoganiss vivntspur l
th lvanof th r
gIogipositiv.moirs d SoitD'Histoir Nturelld Pris 1z 259-268.
1825.D la formationdstrrainsdsnvironsd ris.Bulltide l mind-st of iologists or
SoitPhilomtiqud Pris:74_77,88_90. In rnt yars, in o
Rudwik, .J. S. 2008.'X/orldsbefordm: Th Rlstrtil of Gllhistori A I l n F o r 'L y n h h s
th ge of Rform. hiag:Univsit,vf hiago Prss. prat ftursof gn
Wilson' L. G. 1972.rlasLll:Th Yrsto 784/. Nw Havn,CT: Yal
transposal lmnts(
UnivrsitPrss.
1998. Lll i mri:TrstltiGolog, 1841*1B5.1. Baltimor: 7999 Ln and Fort
Johns Hopkins UnivrsityPrss. -].H. first prinipls of popul
slightly dltriousmut
nomi faturs that dist
Lynh, ihal (b. 1951) from multillular orga
!7hn onsidring th impat a sintisthas had' on must tak into ount vrsial xplanation in
th radth :rnd dpth of h intllualontriutions.By thos ritri, vry ook The rigis o| ()
fw of his prs hv hd a grt irrrpat On volutionary iology than s y n t h s i so f r h i s w r k .
ihal Lynh. Furthr, his arr hirs n markd y that most valud of volution maks sns
..stady Kimura's lassi Th |
aadmi ahivmnts: progrss.''
 Lh 701

r aftrdvloping his own dis- Lynh gan his arr as an ologist' studying th ommunity strutur
f th th' ritiizd from t of zoplankton as a studnt of Josph Shapiro at th Univrsity of in-
ing thrmal onomy for th nsota. [n a sris of paprs stmming from his dissrtation rsrh, h
ll from th 1850s on aftr artiulatda omprhnsivmodl of lif-history volution in ladora, in-
isdthi novl thrmodynam- orporating dispart lmntsinluding prdatirln, omptition, and nr-
i). Th intriatomplxity of gtis, i:rnd ulminating irr th first f nrany influntial synthti works
l Russl !alla's agrmnts (Lynh 1980). His osrvation that a tad-off twn prsnt and futur
agrmntsnd disagrmnts rprdutiv suss nd not b nssary was a hallng to th onvn.
, dtsof volutionay iology tional wisdom of lif-histor thory. Th undrlying philosophy of lwavs
nraliz:ltion.Thos dbts are skingth simplst plantion has n th hallmark of his arr and has
y omplxity. put him at th ntr of ontrovrsy irlmost unintrruptdly for th past
quartr ntury.
Th l970s and l980s witnssdtwo imprtant dvlopnrntsin volution-
thrth' Nw York: Sin ary iology: th mrgnof volutionry quantitativgntisand th mat-
uration of th fild of molular volution. By tlr mid-1980s' Lynh had
R. Krr,trns.dinurgh: invntdhimslf s qllantittiv gntiist and ovr th nt dad s-
talishd himslf :rs a ladr in th fild. Along with Russll Land, Lynh
.lrlationsof th ds twn
was instrumntal in odifying a nutral thoy of volutionary quantitativ
luth-astrfngland.ls of
gntis(e'g.,Lynh and HilI |986). Th txtook Gnetis d nlsis of
, Fruilry|828. Prldigsof Qutititiu Trits, oauthord witlr Bru Wlsh (1998), is th industy
standard for th fild of quantitativ gntis.
: Dntu. volutionary quantitativ gr-rtis historially fousd on traits undr stai-
|1|1
tt|1\ptto pli th lizing or positiv dirtion:rl sltiorr.Along with Tonroko Ohta nd Al
rarc t uses Nr>tui
Kndrashov, Lynh was among th first to ogniz th importan of slightly
rIsof Sir hrls L,ll,Brt, dltrious mutations to th volutionary pross. Along with ollaorators
tsimil,
Fanorough, U.K.: Wilfrid Garil, Rinlrrd Burgr, nd John ony, Lynh gan sstm-
ati study of th ffts of dltrious mutations on th proility of popula-
or of th rth' dinurgh: tion tintion (Lynh et a|. 199, 7995). Th rsr'rltingsynthsis stalishd
th ..muttionalmeltdown'' in th lxion, and, mor important, stalishd
rs orgarris6s
vivntspour la
irNturIId Pris |:259-268. th rlvn of th rlatiorrship twn fitnss nd ppultion siz in th
ironsd Paris.Bullti de l mind-st of iologists outsid th ralm of thortial population gntis.
In rnt yars' in ollaoration with onry and dvlopmntalgntiist
Rslttitl of Cois!or ilt Alian For, Lynh hs forgd markbl s},nthsisof svrl smingly dis-
go Prss. parat faturs of gnom volution' inluding dr"rpliatgns, intotrs, and
4 / . N w H v n , : Y l
transposallmnts (Lynh and onry 21,2003a, 2003; For t l.
,og, 1999;Lynh and For 2000a,2000; Lynh t l' 2001). oprating trom
1841-1 85. Baltimor:
-1.H. first prirripls of plpulation gntis' Lynh showd that population siz and
slightly dltrious mutations may intrt in suh a Way as to produ th g-
nomi ftllrs that distinguis prokaryots from rrkaryotsand singl.lld
from rrrtrltiliuiirr<lrganisms.ru to form, Lynh providd a simpl, ontro-
ld, on must tak into aount vrsil planation in liu lf a ompliatd' ontrovrsial on. His nt
, u t i o n sB. y t h o s r i t r i a ' v r y ook The origis of Geome Arcbiteture (2007) provids a highly radal
on volutioary iology than synthsisof this work. It is s an-ritilus in sop nd mssg(..Nothing in
narkd by that most valued o{ volution maks sns pt in light of populzrtion gntis'') as otoo
Kimura's lassi Tbe |'leutrl Tbor of oleculr uolutil (198).
702 L1,h

Lynh hs lso mad many important ontriutitlrrs on topis as divrs as

th phylognti mparativ mthod, th voluti<lrro prthnognsis, th
maning of inbrding dprssin' ard statistial popu|ation ]ntis.

BlLIoGRAPHY

Fr, A., M. Lynh' F. B. Piktt, A. Amlrs' Y.-l,. Yn, nd.|. Postltlrr,vit. 1999.
Th prsrvatinof dupliat gns r, omplmntlrrydgnrlltivmutartions.
G tis 15 1 : 1.531_l 545'
Kimlrra, . 1983. Th Netrl Thor of rllulr ullutil, amridg:
mbridg Ut-rivrsity Prss.
Lvnlr, . 1980. Tlr volr.rtinlf 1adrnlif histlis.Q'LtrtrIRuie of
Biolog 55:2342.
althus, Thomas
207, The rigis of Grl Arhittr. Sundland,A: Sinaur Thomas Rort Malthu
Assoiats.
omist. H ws an ordal
Lynlr, ., R. Birrgr,D. Buthr' and W. Gil. 1993. Tlr mutati()1llmltdwn
in sul plpultions..|ttrlof 17ardit84 9-344. yars taught politial
Lynh, N4., andJ. S. onry.2000. h vlutionary fat and Irsqttns of ivil srvants dstind t
dupliat gns.Si290: 1 1 .5l-1 154. frindly and muh likd
2001. Gn dupliation nd vlluti<ln: Rspons tl Long nd Thornton nd His laim to fam is
Zlr:rrrgt aI.Sic 29: 1.5.5 l.
P rinipl of opultil
23a.Th volutionllry dmogirplrof dr"rpliat glls.In A. Myr and
Remrks o te Spul
Y. Van d Pr, eds.,Gom ullutillt,544, Dlrdrht,Th Nthrlands:
Kluwr Aadmi Pulishrs. ers. Malthus was dism.
2003. Tlr origins f gnom omplity. Sci 2: 141'-|404. writrs and thrfor tr
Lynh' Nl.'J. nry, ar-rdR. Burgr. 199.5.utatitlt umrtlationar:rdth mnt ar impossil.Ht
tintion of small popultilrrs.merin Nturlist 146: 489*.518. arlir argLlmnt of B
Lynh, M., and A. Flr.2000e. Gn dupliatior"r nd th origill lf intrspifi
ppulation numrs t
gnomi inrnptiilit,Amri Nturlist 156: 590_60.5.
2000. Th prlailityf duplit lln prsrvltion y supplis an at maximu
..struggl
sufuntior-ra lization. G ctis | 5 4 : 4 59_47. ar invital
Lynh, M., nd W. G. ill. 1986. Phnotypi vlution [y nutral mutation. at th prolm (Mlt
u o l t i o 4 0 : 9 l 5 - 9 3 5 . Malthus's hif intn
Lynlr, M.' . o'ly, B. Y/lslr,nd A. L.lr.2001.Th proaility of
how God had rrangd
prsrr,atinof rrwlv ais gn dupliat. Gatis l59: 1789-1804.
idl tim way aimlssl
Lynh, ., and B' Wlsh. 1998. Gtis rld nlsis of Qutititiu Trits.
Sundrlnd, A: Sinur Assoiats -'B, his work Was takn up
alulations 21sth as
workhouss wr mad(
sty out of thm. on
through wht Malthus
laus that Mlthus ad
spons to ritiism tht
ut loody intrhuman
lt was th sixthditi
nd of Sptmr1838
lizing that thr uld
gud that tl"rrwill b
motiv for hind a n
.Wlla
frd Russl war
s i s w i t h s l t i o t i t s
nd mad it a ruial l
.iutions on tpis s
divrs as
lluti<lrrof prthnogrrsis, th
iill ptlpulatiln gntis.

-. Yan, irndJ. Postlthwait. 1999.

]mlltl]I). dgnrtiv lIl'ltlttlons.

. t r I . u 'i u t i l t . t h i t l g :

tisttlris.Qurtrl Rui lf
althus, Thomas Rort (|7 66_834)
..Sutlrlarrd,
MA: Sirlr'lr
Thomas Rort lthus, known as Bo to his intimats)Was an nglish on-
l993. Th muttrtitln:rlmltdown
ol-nist. was an ordairrd lrgmar irr tlr Anglian Churh and for many
|:9_44. yastaught politil onomy at Hailyury Collg, shool that produd
'ry ft and ons1usof ivil srvants dstind to work in India. As an individual h was warm and
frindly and muh likd; h was an fftiv spakr dspit a lft lip.
spollsto Llng nd Thorntn nd His laim to fam is a ook that first appard in 1798, ss on tbe
Priiple of Populti>s It Affets th Fture Improt,ement of Soiet, ith
luplitgns.In A. iVIyrnd
[4. Dildrht,Th Nthrlands: Rmrks o th Spultions of r. Godi, . odlrt d thr Writ-
ers. Malthus ws dismayd at what h flt Was th fals optimisnr of many
' S i J 0 2 : l 4 0 | - l . 1 0 4 . Writrs and thrfo trid to show that grandios plans for human irnprov.
i o n t l t t m t t I l i onn d t h mnt ar impossil.H did this through a famous inquality, drawing on an
turllist146: 489_.518.
rli argumnt of Bnjamin Franklin and arguing that although human
nd th rigin of intrspifi
' 1 5 6 :5 9 0 - 6 0 5 . p o p u l t i n n t l m b st n d t o i n r s g o m e t r i a l l ( | , 2 , 4 , 8 , . . . ) , f o o d
lrsrvirtiny s u p p l i s a n a t m a x i m u m g o u p a r i t h m t i a l l (y1 , 2 , 3 , 4 , . . . ) . H n t h r
..strr'rggls
ar invitabl for xistn,'' nd plans fo stat aid only ar-
: t i o n v n u t r a Im u t l t i o n . at th prolm (Malthus 1798,14).
althus's hif intnt was natural thologial. H was onrnd to s
)01.h proilit of
how God had arrangd that humans do somthing hr on arth rath than
c tis 1.59:1 789_l 804.
. idl tim away imlssly.Th lthusian inquality is th answr. Howvr,
l sis f Q tititi rits.
-.B. his work Was takn up with nthusiasm y soial formrs' wh usd his
alulations as th asis for drlnian rforms of th Poor Laws' wh
workhouss wr lnad so unplasnt tht th porr would do nything to
staY out of thm' On wy of voiding suh dgrdation would hav n
through what althus trmd prltdtil rstrint (forn rding), e su-
laus that althr"rsaddd to latr, muh-nlargdditions of his ook in r-
spons to ritiisr that h smd to hav implid that God lft n option
but loody intrhuman strif.
It was th sith dition of althus's ook that harls Darwin rad at th
nd of Sptmr188. Gnralizing to th animal and plant worlds and r-
lizing that thr ould no prudntial rstraint in thos aras, D:lwin ar-
gud that thr will a univrsal struggl for istn'and that this is th
motiv for hind a natural form of sltion. Twnty yars latr, whn Al-
.$7alla
frd Russl was indpndntly formulating an volutionary hypoth.
sis with sltionat its ntr' h tlo rmmbrdth Malthusian inquality
ad rd it a ruial lnrntof his thirrkin.Lik althus. th vlutionists

70
704 mmls

strssdthilt struggl rnight silnt and nonovi<lt.ts,

whr organisms at- larg land vrtrats
tl for lif in h:rrd<lndirions.lwv, Darwin n]Walla diffrd in thir million yrs ago). Th
radings of th struggl,with th fornl always inlind to think that it is - aout th ody-siz ral
twn individuals and th lattr prpard to allow that somtims it ours vorS' hrivors' nd
t w ng ( ) u p s . thir aks, whih wr
Plyosaurs wr main]
|LIoGRAPHY
sutropial zons at th'
althus,T. R. l798' lt l.'sslth Prmi1lllf Popttltioz.London:Printdfor By th Lat Prmian,
J..|ohnsn,in St. Paul'shurh-Yar]. Rpint,Nw Ylrk:amillan,1966. dsndants, th ther1
U826| |9|4. l:ss o th Prittiplof Pl1lul,lti.
6th d' l,ondon: nath thir ody' wirh
vryrlrn.
ing, lizardlik postur l
N"ltlll.
Yung' R. l\,1.l9t]5.Dru'ill's|v7t1lh<t: l itt Vitoriultur.
alridg:mi1g Univrsit1.l)rss. -^4.R. th sam tim; this is sr
Latr thrapsids (onv
turs suggstivof inrr
um in th skull for j
ammals indiativ of inrasd
'W small instivorsand
omtonly spak of mammals lrs ing dsnddfrom rptils, ut al-
though modrn rptils long to on of th majlr divisions of amniots,th and hrbivors (somt
sauropsids (whih also irrludirds nd dinosaurs),mammals nd thir fos- might hv bn usdi
sil anstorslongt()th thr mjo 1ir.ision, th syn:rpsids.
Nonrammalian mammals or vn with
synapsi1sar trftnirlldmammal-lik rptils, :rthrmislding nam - swimttlrs, or bipdal r
aus thsanimals r n()tlosly rltdtl thos modrn forms that w all mon in South Afria, w
rptils. graphi distriution aIs
Although th synpsid linag hs volvd nrany uniqu faturs,suh as Aftr th dvastatin
th milk arrd fur tpil tlf rrrammls,sylrapsidshv also rtaind a numr rapsids survivd:th di.
of mor pimitiv ftttrsthat hav n lost o modifid in sauropsids. odonts, initially small
Mammals rtain a glandular skin nd did not volv th thikr sals with hrivorous and arniv
ontriutions fronr t-kratinsn in sauopsids (s wll as th mor gn- Larador dog. ynod
ral alpha-kratin sn ir-r;rll arrrnitlts);thus thy h:lv bn abl to volv anatomial faturssug
mammary glands, srvirtglands, rr1glands that podu various odifrous dgr of mammal-lik
srtions.Unlik suropsids, whih l-ravth divd aility to rt uri inrasd volum of jav
aid and thus produ a onntrtdtyp of smislid Wast' synapsids x- An inrsd rat of lul
rt lativly dilut r.rin,always stord in a urirrary laddr (whih is aphragm (rdution o1
lost in mn sauropsids, irrludingirls).This nlsmmmals to us urin ( s r o l l - l i k ) o n si n t h
for soi;rl intrtins suh as snt-larking.Tlr typially mammalian use air and to rlaim wt
of th pnis for urintion in mais, s wll as for sprm transmission' is also an a signifiant pro
only possil with this typ lf urinirry stup. Diffrns in th intrnal In th latr Triassi t
anatomy lf s1,napsidsanl s;ttlropsidsalso sl-lggsttht vrious fatursvolvd t i o n o f l a r g rs u o p s
onv8ntly within tl-r twl glOtlps; in prtiulr, ndothtny (War- Global hangs in vg
looddnss)larly volvdonvrgtlyin irds and mammals. sd frns y th onit
Synapsids and sauropsids oth ppard during th Lat aronifrous' othr issrrthat rlats
around 300 million ys ]o.At this tinl oth linagslookd rarhr lizard- lvls wr apparntl).l
lik, u svn2rpsidshd hlrtristi lrol in th skull lrindth y sokt, infrd fror th bioltl
whih allowd for jar,v-rnusl patlsitln and attahmnt.(Most sauropsids irds) that suprior lur
latr volvd an anallgousonditiln in th skr-rll'ut with two hols on ah tiv dg. Additinally,
sid, known as th diapsid ondition.) Synapsids wr initially th dominant sistant to dsiation th.

vior'rs,whr organisms at-
anl.Walldiffrd in thir
inlindto think that it is b-
oW thirt somtims it ours
lpLitttk. Llndon: Printd for
{w Ytlk: N4rillan,1966.
optt|titltl. 6tlr d. l-ondon:
|i Vitorion ,,'',',,'-'.n'
snddfrr rptils, ut al-
aior divisions of irmnilts,th
aurs),mammals nd thir fos.
th synirpsids.Nonmammalian
i' a fatf rnisladingnam -
rosmodrn forms tht w all
rany uniqu fatures' suh as
ls hv also rtaind numbr
)st or modifid in sarrropsids.
: volv th thikr sals with
psids (s rvll as th mor gn-
thy hav n abl to volv
-hat prodtrvarious odifrous
. r.lrivdeility to xr ui
smisolidWast' synapsids x-
in a urinry laddr (whih is
; l l sm m m l st o t ' t s u r i n
Tlr typiall mammalian us
ftrr sprm tansnrission'is also
rp. Diffrns in th intrnal
st that vrious fatllrs volvd
artiular, ndothrmy (warm-
irds and mamtnals.
luring th L,atarbonifrous,
.h liegslookd rathr |izard-
th skull hind th y sokt,
L attarnnt.(ost sauropsids
:ull, ut with two hols on ah
;ids wr initilly th dominant
mmls 70.\
th Prmian (z99_z5\
larg lnd vrtrats,spiall prominnt during
plyosaurs, whilr spannd
million yars ago). Th arlist forms wr th
dogs today and inludd arrri-
aout th ody-srzrng sn in dmsti ..sils'' on
had distint
vors, hrbivo..,, nnd fiih .nt..,. Som plysurs
hat xhngrs.
thir aks, whih wr praly usd as nvironmntal
that wr in tropial or
Piyosaurswr mainly io..nd in gographi aras
Amria.
sutropialZonst that tim, suh s North
wr largly xtint' rpld y thir
y tir. Lat Prmian th plyos:rurs
thir lgs mor undr-
d.*..,,d"nt,, th thrapsids. Thrapsids had movd
upright thn th sprawl-
nath thir body, with a stan that was now 1or
tl run and ratlr t
ing, lizardlik postur of plyosaurs, naling thm
of a highr mtaoli rat.
th sam tim; this is suggstivof th voltrtion
linags) volvd vrious fa-
Latr thrapsids (onvrlJntly n*n.,g diffrnt
rat, suh as inrsdvol-
,u'., ,ugg.,.iv of inr]si,rglvls f mtaoli
um in th skull for iaw -i,.l., and mor omplly diffrntitdtth,
Th thrpsids iIrludd
indiativ of inrasej amounts of food prossing.
smallinstivrsndmdiumtolarg-sizdarnivors(sornasigaslions)
andhrbivors(smasigasbison,withhornlikknobsonthirhdsthat
Bt in ontrast with modrn
might hav n usd in iitrspifi omat).
tlrlvr no glidrs,
nrammals or vn with ontmporanoussauropsids,
swimmrs'oripdalrunnrs.LatrinthPrminthrapsidsamom-
this tim. This hnging go-
mon in South Afria, whih had old wintrs at
for thrmorgulrion.
g r a p h i d i s t r i u t i o na l s s u g g s t sa n i n r a s da h i l i t y
two main linags of tlr.
Aftr th dvstating .'1J-'..Ini*'1 xtintiOns
hrivors' and th yn.
apsidssurvivd:th diynodonts' pig- to ow.sizd
latr volvd largr
odonts, initially small and instivorous. ynodonts
non Was muh iggr than a
hrivoousand rnivorous forrs' although
Labradordog.ynodontswranstraltomammlsandhadntlmrous
rat, whih suggsts som
anatomial faturs suggstivf a high rntaboli
degrofmammal.lik-ndothrmy.ororrrpltthandvidnofn
w prossingror food.
inrasdvolum of jaw musls show that thy
vidn of a mr.rsulardi-
An inrasdrt of lung vrrtilationis shown y
y vidn o{ turbinat
aphragm (rdution oft-th postrio ris) and
to Wrl inlming
(sroll-lik)ones in th. .,n,ai avity. Turinats at oth
aus wtr loss fronr th lungs
air anl to rlaim wat from pird air,
vntilation.
n b a signifiantprolm with a high rat of
thrpsids startd to fa omptition from th radia-
Irr th latr rissl
(anstorsof dinsaurs).
tion of largr sauropsids, inludirrgth thodonts
Gloalhangsinvgtation,suhasthrplamntoftlrmlrrhai
volutionary trnds.An-
sdfrns by th onifrs,my hav influndths
that tmosphri oygn
othr iss that latsto this rplamntmy
in rh Prmin, and it an
lvlswr apparntly lowr in t Triassi thn
thodonts (rodilsand
infrrd from th bioiogy of th living rltivsof
providd thm with ompti-
birds) that suplor lu,ig fun.tion may hav
mor highly r-
tiv lg.Additionally,1h. .gg, of irds nd roodils
this {aturrny hav givn
sistant to dsitronthn thos of othr mrriots;
706 mmls

th thodonts a omptitiv dg in th apparntly drir onditions of th ditary itms. Thrians all

Triassi. Lat Triassi ynodonts am progrssivly smallr, prhaps down. I a d t h a t o u l d n r l w s w i r
sizing to sapomptition. At th vry nd of th Triassi (around 200 mil- i n g p o s s i l t h h a r e t r
lion yars ago) only four lings of synapsids rmaind. Two rat-sizd m o d r n m r s u p i l s n d p
linags of ynodonts prsistd into th arly Jurassi, along with th first rathr than laying ggs),a
tru mammals. Only mammals prosprd past this tim, although a rlit it is not lar if viviparit1
ynodont (and also a rlit diynodont) ar now knwn from th aly r- volvd this ondition indt
taous (-140 million yars ago). Th arlistmammals of th Jurassi wr ilar in numrs of spr
shrw sizd, with tth indiativ of a shrwlik instivorousdit. Th ma- n o n i g g rt h n t h s i z
jor larg land vrtratswr now th dinosaurs, a ondition that ontinud monly lld th ag of
throughout th rst of th A4sozoi. malian volutinary hist
Th arlist mmmals hd rlativly largr rains than ynodonts, as wll sizes to fill th typs of lar
s modifiations of th akon to allow for flion up and down. In latr nosaurs. Th volution of
mammals, with a mor moil shouldr, this fatur allowd for ounding tt of th hanging patt
loomotion, ut initially it may hav n important to allow mammals to li hangs in gloal limat
on their sid to sukl thir young. Th tth of mmmals now interlokd rak up during th s
prisly, indiating atual hwing (mastiation) of th food, and formd Gondwana at th start of
only two sts (milk tth and prmannt tth,as in humans). Th volution h d n o t y t r a h dt h i r
of ssntiallya singl, nonrplaing adult dntition ws proaly important lsnof ontinntalpll
for prisly oluding tth ausontinually rplaing tth ould not b of mountain uilding tha
maintaind in pris alignmnt. This typ of tooth rplmntis proaly proimation th world 1
also indiativ of th volution of latation austh ruption of dntition warm' tropial-lik ndi
ould dlayd until th yorrng nedd to at solid food. Th volution of th highr latituds and ir
sukling rquird th volution of lips that ould form a sal and musular Ths hanging globa
hks; ths faturs wr proaly prursors of th ompl sris of fa- typs of mammals that
ial musls, sn only in mammals, that humans now us for sph and filld y animals suh as
faial xprssions.Lik th modrn-day monotrms (platypussnd hid- around 5 million yars a
nas), th most primitiv of th modrn mammals, ths arly mammals must th dsrtifiationof trop
hav n gg laying, would hav lakd nippls and an trnal ar, and sd groups of mamma
proaly had a dns fur oat. m a i n l y o u r r dv i a m i g
During th Jurassi svral diffrnt linags of arly mammals volvd sparating ontinnts)rs
and divrsifid' mainly instivorous or arnivorous in thir dit. Fw wr mammalian typs. Th
biggr than a rat, and thy wr mainly tint y th Crtaous. our tralian marsupials and pl;
knowldg of A4sozoi mammals has gratly inrasd in th past fw yars, marsupial "wolf," or thyl
and w an now add digging, swimming, gliding, and somwhat largr (opos- frnt groups of plant
sum-sizd) arnivorous raturs to th divrsity of known forms. In th Lat losly r1atdto lphan
Jurassi (-180 million yars ago) a distint omnivorous/hrivorouslineag sphr.
appard, th multiturulats (so alld aus of thir ompl hk of th various mamm
tth). h multiturulatsWr a highly sussfullinag of rodntlik adaptd for spifi ol
mammals that survivd into th nozoi (until around 40 million yars at diffrnt tims and pla
ago), whn thy may hav n lipsd y th volution of tru rodnts. is th sar-toothdprdz
During th latr part of th arly rtaetlus, a nw typ of mammal p- ats, within now-xtin
pard, th thrians' inluding th first tru marsupials and plantls.(Intr- South Amrian marsupi
stingly' th first monotrms also dat from this tim.) of havily uilt, musula
Thrians possssdnw' mor ompl typs of hek tth (triosphni fror hrivrous forms
molars) that allowd thir ownrs to prossa roadr sptrum of availal is th larg(ison-sizd
 mmls 707

ditary itms. Thrians also had a mor hinglik ankl joint and a shouldr
ntlydrir onditions of th
ivly smallr' prhaps down_ lad that luld now swing frly, ontriuting to th strid lngth and mak-
ing possil th haratristi ounding gait of many modrn mammals. Both
h Trissi (around 200 mil-
modrn marsupials and p|antalsar also viviparous (giving irth to ais
Js rmaind. Two rat-sizd
rathr than laying ggs),although thy do it in somwhat diffrntways' and
urassi, along with th first
it is not lar if viviparity Was prsnt in thir ommon anstor or if thy
this tim, although a rlit
,known from th arly r. volvd this ondition indpndntly. rtaous mammals wr divrs, srm-
ilar in numrs of spis to th rtaous dinosaurs, ut wr still small,
Lammalsof th Jurassi wr
non biggr than th siz of a small dog. Although th nozoi ra is om-
: instivorousdit. Th ma-
mnly alld th ag of mammals, it in ft rprsnts only a third of mam-
s, a ondition that ontinud
malian volutinary history, whn mammals raditd out into largr ody
'ainsthan ynodonts' as wll sizsto fill th typs of larg land vrtratnihs prviously hld by th di-
nosaurs.Th volution of nozoi mammals is st undrstood in th on-
lxion up nd down. In latr
tt of th hanging pattrns of th arth's ontinnts and th subsqunt
aturallowd for ounding
.tantto allow mammals to li hangs in gloal limat. Although the suprontinnt Pangaa startd to
lf mammals now intrlokd rak up during th sozoi, th southrn ontinnts Wr still unitd in
Gondwana at th start of th Cnozoi, and th northrn ontinntal loks
ln) of th food, and formd
had not yt rahd thir final positions. Movmnt, fragmntation'and oa-
as in humans).Th volution
lsnof ontinntal plats ratd hangs in oan urrnts and pisods
tion was proaly important
of mountain uilding that rsultd in gloal limati hangs.To a first ap-
r rplingtthould not b
proximation th world hangd during th nozoi from initially gloally
loth rplamntis probaly
warm' tropial-lik onditilns to a world that am inrasingly oldr at
l u s t h r u p t i o no f d n t i t i o n
th higher latituds and inrasingly drir throughout.
solid fod. Th volution of
Ths hanging glol limati onditions onsquntly inflund th
rld form a sal and musular
typs of mmmals that volvd. For mpl, th ologial nihs now
s of th omplx sris of fa-
filld y nimals suh as polar ars and amls wr not in istn until
tans now us for sph and
around 5 million yars ago' with th dvlopmnt of an Arti i ap and
.rms(platypussand hid-
th dsrtifiationof tropial rgions. AdditionalIy' th isolation of diffrnt
ls, thsarly mammals must
'ls and an xtrnal ar, and sd groups of mammals on diffrnt ontinntal bloks (lthough this
mainly <rurrdvia migration rthr than th mammals ing arrid on th
sparatingontinnts)rsultd in a grat amount of onvrgnt volution of
:s of aly mammals volvd
mammalian typs. Th st known of ths r th xampls twn Aus-
orous in thir dit. Fw wr
tralian marsupials and plantal mammals lswhr(.g.,plantl wolf and
int by th rtous. our
marsupial ..wolf,'' or thylain),ut mny othr ampls ist twndif-
rrsdin th past fw yars,
frnt groups of plantals. Fo xampl, th Afrian goldn mol ts mor
3, and somwhat largr (opos-
losely rlatd to lphantsthan it is to th tru mols of th Northrn Hmi-
y of known forms. In th Lat
sphr.
lnivrous/hrivorous linag
of th various mammalian omorphologial typs (partiular ody typs
:aus of thir ompl hk
adaptd for spifiologial rols), two typs that hav rpatdlyvolvd
rssfullinag of rodntlik
at diffrnt tims and plas in th past ar missing from today's world. On
ntil around 40 million yars
is th sar-toothdprdator. This typ of mammal volvd twi within tru
h volution of tru rodnts.
ats, within a now-tint family of ..fals sar-tooth'' arnivors, and in
' a nW type of mammal ap-
South Amrian marsupils. (Australian marsupials produd a similar typ
rsupialsand plantals.(lntr.
of havily uilt, musular prdator, th marsupial lion, whih had volvd
ris tim.)
from hrivorous forms and so lakd th distintivanins.)A sond typ
:s of hk tth (triosphni
is th larg (ison-sizd)lawd hrivor that ould ris on its hind lgs to
roadrsptrum of availal
mmls

pull down ranhs. ammals lik this volvd among th hlrs-rlatd rn Hmisphr. Th l
haliothrs in th Northn Hmisphr and Afria, among nativ ungu. onfind to th trtlpis
lats and giant ground sloths in South Amria, nd among womat-rlatd ats) mad an appar
forms in Australia. hrivors)Wrom
A rif history of nozoi mmmalian distriution is as follows. At th rathr than fruit atrs
start of th nozoi, Afria was isolatd from th othr ontinnts and took ovr som of tht
volvd its own spializdfaun (.g.'lphnts);primats wr also known rly ion, around
arly on in Afria, ut othr northrn immigrants (suh as arnivrrsand un- and drying during th J
gulats, or hoofd mmmals) did nt rah thr until th arly ion, th sprd of savann
around 20 million yars ago. Th monotrms Wr th oiginal Australian Long-1ggdungulat
inhabitants; marsupials (immigrating from South Amria via a thn-i-fr in North Amria (hor
Antatia) Wr not known until th rly on, around 55 million yars tiv forms). Howvr, 1
ago. South Amria was isolatd until th middl Plion (2.5 million yars Wr not in vidrru
go), whn th Isthmus of anama formd. Its original fauna onsistd of ing in th lat nozt
dntats (sloths' armadillos, and antats),marsupials (lpossums nd transformation of ths
orhyanids, now-xtint prdatory forms), and nativ ungulats (now all lss prairi' with th l
tint). Around 40 million yars ago it gaind its stoks of primats and million yars ago) oo
aviomorph (rltdto guin pigs) rodnts' most likly from nimals rafting Northrn Hmisphr
ovr from Afria. Alut 2.5 million yars ago many frms migratd from dr tod and wolly
North Amria, nd now ovr half th South Amrian fauna is of rnt nw high-latitud ha
northrn origin (.g.,ats, foxs' tpirs, dr, and mouslik rodnts).North and drying rsultd in
Amria and urasia hav long had fairly road faunal onntions,ut today ial Afria and Asia, rv
their mammal faunas ar mor similar than in many Past tims. For ampl, that was formrly om
North Amrian dr and ison ar reent (Within th pst 3 million yars)
|BLloGRAPHY
ursian immigrants.
During th arly nozoi (th Plon and arly on pohs, from Agusti,J., rrdM. Ant6n
65 to 50 million yars ago) th world was in gnral warm and qual and' Ycrs f mmlit
with th rowsing pssur of th hrivorous dinosaurs movd, was largly Prss.
Bnton,. J.2004, Vr
ovrd in forsts;topil-lik forsts tndd vn within th nfins of
A: BlkwllPu
th Arti irl. Th typs of mammals Prsntwr largly roral inst iflli, R. I-.2001.rly
and fruit atrs'with a fw trrstrilomnivors nd hrivors.Small prd- 12\4-1226.
ators inludd rly mmrs of th modrn ordr arnivora, ut largr J a i s , . . 2 0 0 1 .V i t o
prdators wr not divrs and blongd to groups now tint. Most of d.,Th Book f Li
Krnp,. S. 2005./:e(
ths mammals hav n trmd rhi typs, maning that thy longd
UnivrsityPrss.
to linags that did not prsist until th prsnt day (indd,most Wnt x- Kilan-Jwlrowsk, Z.,
tint for th nd of th Palon).Th start of th on saw th first lf Diosurs:rigi
apparanof primat and modrn ordrs of ungulats(with th first rodnts UnivrsityPrss.
apparing in th latst on) and also th highly spializd ats nd Plugh,F. Ll., . . Jn
whals. Th first known fossil ats ar alrady larly spializdflirs, and SaddlRivr'NJ: Bt
Thwissn,J. G' ., n
w hav littl information on thir anstry. Howvr, in th past dad a
marovolution. Bir
sptaularsris of arly fossil whals hs n olltd that larly shws Turnr,A', and . Antt
th transition from a land animl to a srndrilyaquati n. uolutio of th f
Around 50 million yars gO tmpraturs in th highr latituds startd to UnivrsityPrss.
fall, rsulting in inrasd sasonality zrnd wintr frosts. By th start of th Vughan,T. A.' J' A. Rv
P h i l d l h i S :a u n
oligon (33 million yrs ago) th tropial forsts wr onfind to the
qutorial rgion, and diduouswoodland sprad aross muh of th North.
 mmls 709

lvd among th hors-rlatd rn Hmisphr. Th arhai manrmls Wr now all tint, primats wr
d Afria, among nativ ungu- onfindto th tropis, and mor modrn typs of larg prdators (dogs and
a, nd among womat-rlatd ats)mad an apparan'Rodnts and raits (or quivalnt typs of small
hrivors)Wr omlon, and th ungulatsWr largr and mostly laf atrs
triution is s follows. At th rthr than fruit atrs.In South Amria larg rldnts (lik ttlday's apyara)
.om th othr ontinnts and
took ovr som of th small ungulat nihs. A warming trnd during th
rts);primatswr also known arly Mion, around 20 million yars ago' followd y susquntooling
nts (suhas rnivrresand un- and drying duing th lt ion (12_.5million yars ago), rought with it
:hruntil th arly Mion, th sprad of savann-typ haitats (trd grasslands) at highr ltituds.
|sWrth original Australian Long-lggdungulats'with tthmodifid for ating grass,amprvalnt
rth Amria vi a thn-i-fr in North Amria (horss),urasia (antlop),and South Amria (tintna.
'n'around 55 million yars tiv forms). Howvr, th quivalnt typs of marsupil, th largr kangarols,
dl lion(2.5 million yars Wr not in vidn until th Plion in Australia. Furthr ooling and dry-
[ts original fauna onsistd of ing in th lat nozoi (from around 8 million yars gl) rsultd in th
), marsupials (opossums and transforrationof thspodutiv savanna haitats into lss produtrvtr-
rnd nativ ungulats (now all lssprairi' with th loss of larg mammal divrsity. Plio.Pliston(from 5
rd its stoks of primats and million yars ago) ooling rsultdin th i ags (larglyprindy th
rostlikly from nimals rafting Northrn Hmisphr),and spializdmammals (suh as musk o and rin-
lo mny forms migratd from dr today and woolly rhino nd mammoths in th rntpast) inhaitd th
r Amrin fauna is of rnt nw high-latitud haitats suh as tundra. Howvr, i-ag glol oolirrg
and mouslikrodnts).North and drying rsultdin th dvlopmntof xtnsivsavanna haitats in trp-
l faunalonntions'but today ial Afria and Asia, whih tody prsrvth typ of larg.mammal divrsity
many pst tims. For ampl, that was formrly ommon worldwid.
.ithin th past 3 million yars)
BIBLIOGRAPHY

Lndarly on pohs' from Agusti,J'' and M. Ant6r-r.2002' mmls,S|lrtooths, d Hmiids:65 illio

;nralwarm and qul and,
dinosaursmovd,was largly
d vn within th onfins of
]ntwr largly aroral inst iflli,R. L. 2001. arly mmmalianradition
sand hrivors.Small prd-
n ordr arnivora, ut lrgr
groups now xtint. N4ost of
s,maningthat thy longd
lnt day (indd,lost wnt x-
art of th on saw th first
rngulats(with th first rodnts
L highly spializd ats and
y larly spializdflirs, and
Howvr, in th past dad a
:nolltdthat larly shows Turn,A., and M. Ant6n. 2004' uoluigd: ]llltstrtd
rrily aquati on.
n th highr latituds startd to
ntr frosts. By th start of th
Ll forsts wr onfind to th
rd aross muh of th North-
Yers of mrliultilin urop. Nrv York: lurniaUnil'rsit
Prss.
Bnton,. J.2004. Vrtbrt Plotolog. rd d. haps.5 nd 10' ldn,
MA: BlakwllPulishing.
s..|orlof Plltolog 75:
1214-12.26.
Janis,. ' 2001.Vitlrsy dfault:Th m"rn-ralian sussiln. ln S..|.Gould'
d.,Th Book of Lif' 2nd d., 166_217.Nw York: W. W. Norton r-rd o.
Kmp' T. S. 200.5.Th origi d uolutio of mmls.()frd: Oford
Univrsitrss.
Kilan.Jaworowska, Z.' R. I-.iflli,andZ.-X. l-uo.2004.mmlsfrom th g
of Dirlsurs:rigis, uoltttil,d Strutur.Nw York: llumia
UnivrsityPrss.
Pough,F. H., . . Janis,andJ. 13.isr.2008. Vrtbrt l'if,8th d.Uppr
SaddlRivr,NJ: Bnjaminumrnings.
T h w i s s n , . ] . G . a. 'n d S . B a j p i . 2 0 0 1\.W h l ass p o s t r h i l df o r
marovoluti. BioSi5 |: 107_149.
Guid tlth
urlltiolf th fri l,rg-mmlFu.Nw York: olumia
UrrivrsityPrss.
Vaughn,T. A., J. A. Ryen,nd N. zplwski.)000. mmlg.4t'd.
Philadlhi: Sar-rndrs Pulishing. -.J .
710 Man,s Pla in Natur

NIn,s Ple in Nture (Thomas Hnry Huly)

Thomas Hnry Huxly's 's Pl iNture (1863) (th full titl is ul-
dc s to n's Pl i Nture, but it is alwys known y th shortr titl,
whih appard on th spin) was th first nglish book aftr Darwin's oz
th rigi of Spis to rlt humans nd aps anatomially and in volu.
tionary trms (th rigin had avoidd th sujt).And it did so in punhy
pros for gnral onsumption.
Its inption lay in uxly's disput with th antitransmutationist Rihard
Own. Own, an prt on aps (inluding th gorill' disovrdin th 1840s),
plad mankind in a nw sulass, Arhnphala (ruling rain), in 1858. H
insistd that humns possss uniqu rral hmisphrs,whih divid into a
..hippoampus minor.''
third lo, whos latral vntril ontains a protruding
Huly, a iologist t London's Shool of ins, hatd own's hautur
..thris Gr. ol.
and providntialism and rspondd in 1858 by stating that vry lit-
tl gratr intrval s imls twn te Gorill k the n than ists b- .W.hn
it am to voluti
twn th Gorill & the [aoon|'' (ltur 10, Marh 16, 1858, Royal from harls l)arwin.
Institution Sris,Prinipls of Biology, January 19_Marh 23, 1858' uxly or paradigm; th frmr
Paprs,Imprial ollg, l,ondon, 6.100). This sntimntstands at th hart wondr that Huly at
..Thology Parsondom'' in his talks
of 's Pl i Ntur. ritiizing & rtint' namly human
to artisans in 1859, Huly addd that humans and animals .,must hav pro- dsignd to show yon
dd from on anothr in th way of progrssiv modifiation'' (Huxly grat aps. ost popl

1859). It was, h told a frind, ..as rsptalto modifid mnky as tht ur souls ar insrt

modifid dirt'' (lttrto Frdrik Dystr' January 0, 1859, Huxly Paprs,

Imprial ollg, London, 15.106). Bfor te rigi appard,Huxly had
polarizd th issu lf human volution vrsus ration.
own rassrtdhis laim at th mting f Oxfod's British Assoiation for
haptr (lttr to Hux
th Advnmnt rf Sin in 1860. From th floor am uly's pointd
..on th Zoologial R- t h r l i g i u s p r s sW a
ontradition, and from his pn a sustantiatingssay
tion of mind and sp
lations of Mn with th Lowr Animals'' (1861),whih showed that ap rains
rprint. Th only bar
possssvry ..uniqu'' human fatur.Th disput,whih xplodd publily
shillings, pnsiv fol
in t'etbm tn 1861, alrtd rsptalsoity to human volution.
Mivart was grivd to
Th paupr prssshd long rlishd an aristoray-humlingstial ori- ,,
lb s citis,,, showing
gin, and Hr'rxlywas tapping this groundswll for support. portrayd vo-
rop (unpulishd ltt
lution as slf-ttmnt:it grntd dignity to huml origins. Huly's artisan
Darwin olltion, Un
tlks' running through 1862, fomd th sond haptr of 's Pl in
original).
|Iture(thr wr only thr haptrs).Srndipity providd th third. Sir
I-yll's Antiqit of 1
harls Lyll, who was writing hts tiquit lf , askd uly's viws
tur. Te first mad m;
on Nandrthal lv[an (found in 1856). Huly's nsuing Royal Institutionl-
..On Fossil Rmains sooping th modrn v
tur of Man'' (Huly 1858-1862) would frnis,'s
sand aps for his ans
Plce i Ntur' and h garnrd aounts of living aps for an opning
haptr. |f n's Pl i Ntur was lss assrtivon volution than his l- BIBLIOGRAPHY
turs, it would still prik Vitorian vanitis, not last y suggstingthat hu-
Anonymous.1863,uid
mans might go bak far into gologil tim. Fruary28'287_2
A n i n i t i a l p r i n t i n g o f ] , 0 0 0 o p i s h i t t h o o k s t a l l si n F r u a r y 1 8 6 3 . Darwin, . 1999.h
Darwin had nvr ..rad anything grandr'' than th losing of th middl CambridgUnivrs
 n's Pla in Natur 711

.ry Huly)
r (186) (th full titl is zi-
.aysknown y th shortr titl,
rglish ook aftr Dwin,s on
ls anatomially and in volu-
ljt).And it did so in punhy

t antitransmutationist Rihard
;orill,disovrdin th 1840s),
rala (rulingrain), in 1858.
misphrs'whih divid into a
..hippoampus minor.''
ltruding
ins, hatd own's hautur Shltons ojtha
..thr is vry lit_ Gro. o.
y stating that Clpz. Gorrrl. trL{N.

rill k the than xists -

: 10, arh 16' 1858, Royal
: y 1 9 _ a r h2 3 , 1 8 5 8 , H u l y
rissntimntstands at th hart
rgy & Parsondom" in his talks
,,must hav pro-
rs and animalrs
;rssivmodifiation'' (Huly
l to modifid monky as
uary 30, 1859, uxly Paprs,
origin appard,Huly had
ratlon.
oford's British Assoiation for
l floor am Huly's pointd
..on th Zoologial R-
Lgssay
), whih showd that ap brains
isput,whih xplodd pulily
s o i t yr o h u m a n v o l u i o n .
.istoray-humlingstial ori-
for support. H portrayd vo-
ruml origins. Huly's artisan
lond haptr of n's Ple i
ndipityprovidd th third. Sir
, of , askd Huxly's viws
.'s nsuing Royal Institution l-
858-1862) would frnish 's
o f l i v r n ga p s f o r a n o p n i n g
rtivon volution than his l_
not last y suggsting that hu-
o o k s t a l l si n F r u a r y 1 8 6 3 .
than th losing of th middl
Whn it m to volution'Thomas Hnry Huly hd vry diffrntmotivs
from harls l)arwin. h lattrwntdto providan ovrall sintifithory
or paradigm;th frmr wntd mtrialt fight th hristians.Thr is littl
wondrthat uly at on pikd up on that aout whih Darwin Ws so
rtint'namely human volution.This frontispilf n's Pl in Nture is
dsignedto sow yonddout that w humans ar from th sam stok as th
grataps.ost pplgrd,althoughto this day hristiansusually argu
tht our souls ar insrtdmiraulously.
h a p t r ( l t t rt o H u l y , F r u a r y 2 6 , 1 8 6 , D a r w i n 1 ' 9 9 9 ,| 1 : 1 8 0 ) . B u t
th rligious prss Was stunnd, and many notd th lk of any rogni-
tion of mind and sph. Brisk sals nssitatdan immdiat 1,000-opy
rprint. Th only ar to its falling into gruir hands was th pri: six
shillings, pnsiv for a 159-pag tom. vn so, th atholi St Gorg
ivart was grivd to s translations at Itlian railway stations alongsid
,,obseities,',
showing that th strt-lvlfasination strthd aross u-
rop (unpulishd lttr from ivart to hrls Drwin, April 25' 1870'
Darwin o1ltionU , n i v r s i t y L i r a r y , a m r i d g ,F o l d r 1 7 1 ; m p h a s i si n
original).
Le||'sAtiquit of was pulishd within days of ,s Ple in N-
..a
ture.Te first mad man hundrd thousand yars', old' said on rviwr'
sooping th modrn worldviw, and th sond gav him ..a hundrd thou.
s a n d a p s f o r h i s a n s t o r s ' (' A n o n y m o u s 1 8 6 ) .
BIBLIOGRAPHY
Anlnymous. 186.uid{1sto ||'sPlci Nturc [rviw].Atbum,
Fruary28,287-288,
Drwin, . 1999, h orrs1lod
of hrlsDuill' Vol. 11. amridg:
amridgUnivrsityPrss.
/ tz rgulis

Dsmond, ^. 1997, Hul: From Dui!'s Disipl tl uolutio's High Prist. to work as a unifid v
Rading, A: Addisln-'Wsly.
winin struggl than
Gross, . G.1993. Hippompus mirror and man's pla ill natur:A as stlrdy in
th soial onstrution of nuronatom. ippr',m|>s
matril fom n or1
3: 40_415.
uly, T. . 1858_1862. on fossil rmir-rs of mn. Prtlatligsrlf tb RolI tria. his vi. has
I nstitut i l lf r t B rit i tl: 1Ii,420-422. work on th human
-h
1859. Sinnd rligiorr. Buildr,Januarv 1.5. frrd sidways in thi
1861. on th zoologil rlirtionsof man with th lowr nimals. howvr ltaind, it
Nturl Histor Ruilu: 67_84'
as a omplmnt to D
1863. uiden s to 's Pl i Nture. Llndtln: \/illims nd
Norgt. In rnt yars' ais(
L,vll,. 786. Th tiquit rlf . L.ln:Jhn ur,v. N4argulis (togthrw
Lyons' S. L. |997. ovining mn thy monkvs. Irr A. P. Bar, d., Thlms Sagan) lras n a st
Hr ltl's Pl i Sittd Lttrs: lttttr ssays, 95-l l8. arth as i1norganismi
Athns: Univrsity of Gorgia Pss.
so signifintly that it
Own, R. 1858. on th haratrs,prinipls lf divisin, and primry groups of th
lass mammalian. Jourl of tbc Prodigs tlf th I-i Soit (Zotllog)
ptdly hapd r
'l'_7. gists quat th Gaiz
2:
.W.ilson,
L. G. 1996. Th gorilla and th qustion of huml oigins:Th rlin qually xptdly st
ontrovrsy. Jlurl tlf tll is|or ll|-adiinc d llic! ,\is 57: vind of th ightn
184-2()7. -.D.
lLIoGRAPl])
l\4argulis,
L. |970. rtg
argulis, Lynn (. 1'99) for ,hlrtf th
l,ynn Mrgulis, a longtim profssorof gosinsilt th Univrsity of Mss- o tb Prmbri
200Q.Smbiot
ahustts'Amhst, is proprlv lratdfo hr stongildvoay of th oi.
rgrrlis,L., nd D' Sa
gin of ukurryotlls (lls witlr a nuleus) frotn trtorprimitiv prokary<lt ullti.Nw Yo
lls (lls withlut a nulus). Som vrsion of rhis thrlry gos bk to th
nintnthntury' whn it was pushd y th Russian lihn spialistKon-
stantin Srgivih rzhkovsk (1855_1921), and it was advoatd in th
1920s th Swdish Amrin ]vn !alln. But it lvas argulis wh took lvlarsh,othnil (
up th ida, now known aS te dosmbioti thor, offrd mirlsopi othnil harls al
vidn,rrd pushd it (in hr ook rigi oi ukroti lls, 1970) until nintrrtlr_tury An
it am orthodoy. Paody, and his on
A4argulis laimd that rtain funtining p:rrtsof th ll (organlls)ar rih fossil ds of th \
in fat formr prokaryots that hav n takn up and now funtion for th usd many of his fossi
nfit of th whol ukaryoti ll. Inludd hr ar mitohondria and arsh was orn in
hloroplasts' rsptivly'th powr plants of th ll and (in plants) th or- Shffild Sintifi Sh
ganells th2rtpfom photosyrrthsis.Dfinitir,prof of N{argulisposition trrop rturnd to b
am in th 1980s whn it was disovrdthat tlr DNA f thsorgzrrrlls is unl finnd th P
diffrnt from that of th ll's nulus ut ars signifiant similaritis to pditilns that ars
that of various prokaryots' for instan, in ing irular, hving th sm h was politilly astu
size, and ing l to pfrr th sam funtiotrs as th appropriat or- my of Sins(h s
ganllsor funtions similar to thos of th organlls' ogist of th U.S. G
argr-rlis'sthinking is dply inf]ur-rdy plriltlsophy of lif, wlrih onntioll to Lrosth
ss hrmony and mutualisrrrin natur rath rlrn fighting and arrtagonism. fir fud with dwa
Although som hav argud that th ndosymioti thory an st sn th narrringand dsri
in trms of ptur and slvry-th organllsy th main ll-for hr it is in this volr.rm).H dit
rathr a mattr of diffntlif forms oming togtl"lrin a sussfulatmpt Marsh's tams dis

,lF,uriutil'sigh Priest.
plain ntur:A as sttrdy in
()mpL|s
3: 40-415.
n. Proecligsrl|.th Rowl
u a r l 5 '
''ith th lorr,ranir-rrls.T/.,c
. l-rlnln:\lilliamsand
n Murray.
s. In A. P. Brr, d.,Thoms
(t1r\'ssv-s,95_l l 8.
visin,nd primry grorrps of th
'f th I'itl Slit (Zoolog)
I humrrlrigiIts:h rain
d llid Sis 51:
]nsat th Univrsity of N4ss-
hr strong adv<layof th oi-
oln mo primitiv prokaryot
f this thor1'gos ak to tlr
: Russian lihrrspialistKn-
), nd it Ws advoated in th
But it rvas Margulis who took
' it..thllrl,' offd mirosopi
'f ukrotic lls, |970) unt1|
lrts of th Il (org:rrrlls)
3n up nd now funtion for th
:d hr r mitohondri and
th ll and (in plants) th or-
v p r o o [ f e r g u l i s . sp o s i t i o n
: th DNA of thsorganlls is
a r ss i g n i f i n rs i m i l r i t i st o
ling irular, having th sm
l t t i o n si l s t h a p p r o p r i a t o r -
:ganIls.
, hr philosophy of lrf, whih
t h a n f i . g h t i n gl ] d n t g o n i s m .
iotithory an bst sn
:s y th main ll-fo hr it is
ogthrin a sr-tssftrl
rsb 71
to work as a unifid whol. Sh ssvolution lss in trms of a ruthlss Dar-
winian struggl than s a pross asd on t sidways transfr of gnti
matrial from on organism to anothr, arrid via miroorganisms lik a-
tria' This viw has lr rivd with som skptiism, although rnt
work ln th hurrrirngnom strongly suggststhat gnti matrial is trans-
frrd sidways in this fashion. Sin Drwinism works on gnti variation,
howvotind, it is pobaly rtrto tlrink of this sort of phnonrnon
as a omplemnt to Darwinian volution than s an opposing viw.
In rnt yrs, :.rlsoguidd hr philsophy of hamony and mutualism,
Margulis (togthr with Dorior-r Sagan, hr son from hr mrriag to arl
Saga) has n a Strong dvoat of th Gaia hypothsis, whih ss th
athas an organism and as slf-rgulating'at last Llntil hLlmansdisruptd it
so signifiantlythat it ould no longr funtion proprly. .I.hisadvoay has
ptdly hapd ritiism on argtrlis's had-rany onvntionl iolo-
gists quat th Gaia hypothsis with pr-hristian natur woship-and
qually ptdly sh is mor tilan prpard to wthr th stom' on-
vind of tlr rightnssof hr aus.
-.D,
|LIoGRAPY
Margrrlis,L. 1970. Origi of uhroti alls:uidd Rsrh]mDlititls
for T|trf the rigi t! uo|utilof iro|lil,PIt,d iml lls
o th Prmbrirth. Nw Hvn, T: Yl UnivrsityPrss.
20'SmbiotiPlt: Ntu Llok t t,olutill,Nw York: Bsi Books.
a r g u l i sI, . . , n d D . S a g a r -1r9. 9 7 . S l t d T r u t h s : s s s l G i , S m b i o s i s d
uolutit.l.Nr.York: oprnius Books. -.R.
arsh,othnilharls(1s31-1s99)
othrril als rsh was t most influntial palontologist in lat-
nintnth-ntuyAmri. H xploitd tlr walh of his unl, Gorg
Pody, and his onntions with th U.S. Gologial Survy to plor th
a rih fossil ds of th Wst. H Ws an nthusi:lstisupportr of Darwinisnr and
usd many of his fssil disovristo prvid vidn in support of volution.
arsh was rn in Lokport, Nw York, irr 1831' ws rind at th
Shffild Sintifi Shool of Yal Univrsity nd aftr thr yars of study in
urop rturnd to onrrofssor of plontology at Yl in 1866. His
unl finand th Pody usum at Yal and providd funds for th -rany
xpditions that rsh snt to t !st' Although rsh Was a rsrvdmirn,
h was politially stut and m a powrful figr.rrin th Ntional Ad-
my of Sis(h srvdas prsidntfom 188 to 1895). H was palontol-
ogist of th U.S. Golgial Sr-rrvyfrom 1882 o |892 and xploitd this
onntion to oost his wn privatly finand pditions. H ngagd in a
fir fud with dward Drinkr op ovr ass to W,strnfossils and ovr
th rraming and dsriptitln of spis (salsrth lphatial ntry on op
i n t h i s v o l u m ) .H d i d i n 1 8 9 9 .
attmpt Marsh's tms disovrd mar-rynw spis of fossil marmals and rptils.
714 rsh

'-1i"' Fore-rm. Le'. uPr tlolt. Lov6r olat.

H i s t l l l t i o no f h o r s

(wW
RN.
fl tinuous squnof v
ilfl
l{lr modrn hors, a squ
QuUs.
t1 IJ v i d n f v r l l r t t i l l n
E )(
t, t r n d o f i n s ds p

rfi,
^ww
although th tru storv

I{
4Jl(lt'w\
r s h l s <dl i s o v Jt
PLIoN.
in mon<lgraph.dl
PLloIPPUs.
from Grrrnv.ths\\
l_r
,( i d s h a d v o l v df r . , m
t-l \\ \^J
w s s u s q u n l f1l.l u n
)

ffiffi
n a m d e I a r g t t u m r.

il
o f g i a n t r l ym m m |
l ( l 8 8 6 ) h r r a tl h l r

#h t(
PRooIPPUs
(Ei|!4i'. ansto. Lik Huly,
muh fr.rth llirk ill g
N4arsh was n nthl

.RgF
A
'n
o n r n l r i m s I fr r i r h r l
Io,N.

ir
tf \-cY Lg)
natural sltion woul

,t*
rrsintllignWs
loIPPUs
/]'\ The ltrodutit.l d

(hilhcriuni.

t h t p a | o n t o l o g vp r o v
ing rain siz in th ou
il produtof nint

[
/,111
|l ffiF </
l{l \sss}-/ (\J
ILIoGRAPHY
.l
H r t l v . . H . I 8 3 8 .
oEo
t$
sol{IPPus.
l\ N .Y l r k : plrr
M . t s ho, . . | - 7 .I I

tr
ddrss Dliuardba

At
mwffii?ffi
'\itt t Nlshuill,
oN.
Iili
#h
hous& Tylor.]

\t w, |880. dtrlr
A m,ri . Wr shingttt

oRoIPUs"

P r a l l vl ,o l . 7 .
"fJ" |886.Diort
Y/eshingoll.D(,: Mr,
On a ltur tour t() th Unitd Stts'homas nry Lluly was owld ovr
y othniel Marsh,s olltion of hors fossils.To mak th as for volution in
puli letu,h used this reonstrutionof hosvolution(somthlngw
now know to muh mor ranh-lik),prditingtht soon a fiv-td
anstor would disovrd.T his dlight' soon thrftra spimnwith IyIasstintions
rudimntaryfifth tos was disovd.(arsh al[dit ohipps,..dwnhors.''
Mass tintions ar glr
It was thn disvrdtht spimnshad nfound arlir,althoughnot
rs of spis from di
idntifids horss.Th alirname, 1,rotbrium' taks prdn,
althoughoviouslyarsh's nam has stuk.) frotn rrrlrrginai t
o l o g i a l p r o s s s l.- o
ayrr-rnt or a lk m1
sdimnt. In ()ntrast'l
 ss titios 715

lww
UPr oj!. Low6r ol..
His olltion of hors flssils(sfigur)gnratdWhat appardto on-
tinuous sqLinof volutin frtlm tlr snrll fotr-todrrrohippus to te
rodrn hors, :l squn that Tlrlms Hnry tlxly alld ..dmonstativ
'? vidnof volution'' (uly 1888,90). Th fossils smd to irrdiata
trnd of inrasd spialization y adaptation to running on th pn plains,

frww
although th tru stoy of th hors fmily turnd out to far mor ompl.
M;rrsh rlsodisovrd toothd irds from th rtatlr.rs,whih h dsrid
in a monograp6, ()dltlritbs\1880). Lik th ttr_knownrlltloptr
from Grmany' ths wr haild zrsimportant onfirmation of th viw that
irds had volvd from rptils. Th had of n of arsh's irds, Ihthornis,
was susquntlyfound to long to marin rptil. arsh disovrdand

ffiffi
nald a larg num of dinlsaursnd also studid th volution of group
of gint aly mamtnls h alld Dinort. In his n-rorrogaph on this group
(l886) h trad tlr origin of tlr Ungulata ak to:r hypothtialrtous
anstor.Lik uxly, h livd that th origin of ordrs and lassslay
muh furthr bak in gologial tim than th arlist known fossils.
]VIarshwas an nthusiastisuppotr of Darwinism, although h did not

sffiffi
()nnhimslf with th dtails of how volution rvlrkd.H livd that
natual sltiorrwould gradLrirllvfavr th irrrasingsiz of th rin -
aus intllignwas an importirnt adaptiv advntag.In his 1877 ddrss
T'h ltrodutir> nd Suessio of Vertbrt Lif i mric h argud
tht palontolog providd iar poof of this adaptiv trnd toward inras-
ing rain siz in th Ollrsof vllution.Mash's law of rain growth is a typ.

ffi8
ial produt of rrintr-rth-ntllryprogrssionisr volutiorrism.

B|LloGRAPY
Huly,T. H. l888. AmriAddrsss, tuith Lturo tStudof Biohg'
Nw York: Appltn.
Nlrsh,o. . 1877. llttrodutilnt!Sttssitl lf Vrtbr,ttLif i mri:
ddrss Dliurd bf<lrtll Atltt,rittsstlititlilr th dmtri.

,@@ Sit Nshuill,ll.,ttgltst.0,1877.INrvvn,T: TLrttl,Mr-

Hnry Huly was owld ovr
o m k r h s f o r v o | u t i o ni n
orsevolution(somthingw
ring tat soon a fiv-tod
on thIftra spirrlnwith
alldit obipps,..dawnhors.''
foundarlir'althoughnot
ri um, takesprdn,
hous& Taylo.l
I88. dotoriths: olgrpho th tit TothdBirds rlf Nlrth
mri..Washirlgton, D: Rportof th Gologil}ixplorationof th l.ortith
Parlll.vol. 7.
1886. Dir>crttct: oogrLl|) of tl tiltt rdr tlf Gig|lti h7Ll77I|Llls,
Washingtlrr,L): Mlngrrhsf th U.S' Golrlgill Surv, vol. t0.
-P.J.B.
ass xtintions
ass tintions ar gloal lrniislf rif lurirtionin whilr larg num-
rs of spis from divrs haitts ar destroyd.Thy ar thr:s distint
from mor rgional xtintion Vntsthat oftn our ausof normal g-
ologial prosss.For xrpl, spis that ar rstritd to a main m-
,vn-rnt or lak nly go tint if thir hbitat is gradually infill]with
s d i m n .I t t t r t t t r t,h g l . l l l a l I l J t u t l I n l . t s s t i l 1 t i ( ) l lrs q i t i ss t r r l l e
716 ss titios

gloal-salatastroph' oftn involving th nar shutdown of all photosyn-

thti ativity. Suh disasrsrquir mjor hangsof th arth'slimat and
oan irulation systm. Th most famous of ths g1ol disastrs is th
giant mtorit impat that is thought to hav trminatd th rign of th
dinosaurs 65 million yars ago. This is known as th K-T vnt ausit
marks th oundary twn th rtaous piod' whih is splld with a
K in Grmn, and th fllowing Trtiary. Howvr, this is not th only mass
tintion of th fossil rord. Plots of tintion rats through tim (s
figur) rvl grat vriability, with fiv major paks larly sn and prhaps
twi this numr of smalir xtintion vnts.Th major paks ar known
s th ig 5 mass tintions' with th K-T vnt bing th most rnt. How.
vr' y fr th iggst mass tintion of all tim was round 250 million
yars at th boundary twn th Prmian and Triassi priods. This P.Tr or
nd-Prmian vnt saw th loss of mor tan 90o of all spis.This atas-
troph was as dvastting on land as it was in th sas and oans and,
uniquly, was also a svr risis for trrstrial plants. In othr xtintion
vnts plants gnrally fard muh ettrthan animals.

A U s E so F M A s s x | N | o N s omparisln of tin
Th ida that mass tintions hav puntuatd th history of lif has n larg mtorit rtrl
rognizd xtintion
around flr a long tim. Th rason that most mass-tintionvntsour at
ruption of larg vol
mjor oundaris of gologial tim, suh as etwn th rtaous and
oinids with mass
Trtiary' is that thy mark major disontinuitisin th history of lif and thus
ar usful markrs for th sudivision of gologial tim. Ths tims w
ovious vn whn suh intrvls wr ing dfind in th arly nintnth liquid and last th
ntury. Howvr' th intnsiv study of th auss of mass tintions is thy would stay sus
a muh mor rnt sintht an datd to th puliation of a study y prhaps vn yarsal
.Waltr,
Luis Alvarz, his son and two ollagus in 1980 (Alvaz t al. dvastating for th p
1980). This group masurd th onntrationsof th xdinglyrar tra most osystms.Tl
mtal iridium in sdimntsthat straddld th rtaous-Trtiarytransition. though th food h
For th most part this mtal ours in tiny amounts in th arth's rust, ut whih would di of s
th Alvarzs nd ollgus found a dramati nrihmnt prisly at th snlight. It appars l
lvl of th mass tintion. Thy suggstdtht this iridium am from a nd. This original ki
hondriti nrtorit(on of th most ommon typs of n-rtorit) and got addition of various <
into th sdimnt from fallout in th aftrmath of a giant impat. Th iridium sit is vry unusuai i
anomaly has now n dttdin dozns of loations round th world, and This would hav b
this, togthr with th disovry of th impat ratr in th Yut5n Pnin- ntratd typ of aid
sula of io, has nsurd that thir hypothsis is now widly aptd. this ornr of Mi
Th ratr is loatd nar th villag of hiulu and is urid nath to hit.
youngr sdimnts.It may as muh as 180km in diamtr, making it on Th 1980sand 19
of th largst irnpat ratrs known on arth. impat with etinti<
In th original 1980 artil th link twn th impat and th rsultant plil to all tin
mass tintion was thought to hav om aout from th gloal darknss tintions in whih tl
that would follow suh a larg impat. Th nrgy of suh an vnt would spad vry 26 milli
pulvriz oth th mtorit and targt oks into dust and fin droplts of nation that is unlik
 ss titios 7I7

a shutdown of irll photosyn-

rgsof th arth's limzrt and
o -''
-
: thsgloal disastrs is th .9
.
r trmintdth ign of th
l s t h K . T v n t u s i t
9E
o
iod,whih is splld with a 9
.
vr'this is not th only mass .?
o
Jtion rts through tim (s
lksllysn nd prhaps
l'l
. Th mior paks ar knwn lllanicouga

rt ing th most rnt. FIow-

tim Wxs around 250 million
irli
llrl
l Triassipriods.This P-Tr or
)0" of Il spis'This atas_ xtintion lntensity
rlr;
in th sas and oans and,
9

x

lill
i l p I a n t s .I n o t h r x t i n i o n
o
animls. o
qr

omparis<lnof xtintionratswith th tirningof largvolaniprovinsand

iagmtoritrtrsvr th pst 300 million yars.Not tht all th
d th history of lif has n
rognizdxtintitlnvnts(markdwith n astrisk)oiidwith th
nss-tintionvntsour at
ruptionof larg volanipovins;howvr'onl th lriulub ratr
; twnth rtaous and o i n i d sw i t h m a s s - x t i l l t i onr ' n t .
s in th history of lif and thus
logial tim. Tlrs tims wr
dfindin th rly nintnth liquid and last thm into th outrmost rahs of th atmosphr.Thr
: auss<lfmass xtintions is thy would sta),suspnddin th uppr atmosphr for svral morrths and
th puliationof a study y prhps vn yrs and thus lok most sunlight.The onsqunswould
: g u si n 1 9 8 0 ( A l v a r e z e t a | . dvastatingfor th photosynthsizrsthat form tlr bs of th food hain in
s of th dinglyrar tra nlost osystms.Th rsultant asad of tintions would pss On up
t r l u s - T r rt yi t r n s i t i o n . through th food hain to th hlghr onsumrs' suh s Trlsurus r'
n()untsin th rth's rust, ttt whih would di of stvati<rn, n<ltto rnntion th old usly th lak of
ti nrihmntprisly at th sunlight. It appars that th magnifint dinosur dynasty mt a nrisral
that this iridiunr am from a nl.This original kill snario still holds sway with sil-ttists,
trt witlr th
on typs of mtorit)and got ddition of various othr nasty ffets.For xmpl, th hixulu impat
:rof a giant impat. Th iridium sit is vry unusual in that muh alium sulfatis prsntin th targt strata.
oationsrond th rvold, and Tlris would hav bn vaporizd on impat and tlrn rind ak as a oI1-
t ratr in th Yuat5n Pnin- ntratdtyp of aid rain (sulfurraid). From tlr point of viw of dinosaurs,
lthsisis now widly aptd. this orn f Mio wi1s partiularly trnfortunat pla for a mtorit
.iulu and is burid nath ro hit.
) k m i n d i m rm , akingit on Th 19t]0sand 1990s saw muh dat on th onntion of th mrorit
impat witlr tintion. nd mny sintistsonsidrd th K- snario ap-
n th impat and th rsultant plial t all tintion vnts.Som wlrkrs vn saw a priodiity to x-
tbout fror th gloal darknss tintions in whilr th K-T vnt Was jtlst on of svr:rlvnts that wr
nrgyof suh an vlt would spd ever 26 million 1'rs.A rgular priodiity r1uirsa rgular pla-
s into dust and fin droplts of nation that is unlikly t() om from random arthrr'rnd prosss.Thus
718 ss titios

various astronomial snarios Wr proposd whry th arth was su- of asalt lavas: arb
jtd to dvastating mtorit and/or omt showrs evr 26 million yrs. dioxid is a grnh
owvr, susquntstudy of th fossil rord and improvd ag dating of mosphr will aus1
th roks has shown that svral of th 26-million-yar tintion vntsdo (y ration with w.
not xist (for ampl, thr was no tintion eent 26 million yars aftr th most notiabl
th dath of th dinosaurs).Non of th tintion vnts that rmain show tion of Mount St. H
a rgular spaing. For mpl, thWas a mass xtintion 200 million yars mat. Howvr, su
ago, known as th nd-Triassi vnt, followd y smallr-magnitud- ar raind out of tht
tintion vnt 20 million yars latr in th arly Jurassi.Th nt xtintion that th ruption of
vnts did not our until th Crtaous' ovr 80 million yars latr, and gloal warming' sug
ths wr all vry minor affairs ompard with th K-T vnt. mor importnt li
Attmpts to link othr xtintion vntswith mtoritimpts hav so far gloal tmpraturl
provd rathr unonvining, mainly ausof a lak of vidn.For am- all lif on ath. Thr
pl, th Maniougan ratr of northrn Ontario, anada' is roughly 70% tion of flood asalt
th siz of th hixulu ratr. It is on of th largst ratrs known ut that lads to a runa!
sms to hav had littl or no fft on tintion rats. Th ratr was formd
220 million yars ago at a tim markd y vry low etintion rats. ow-
oNsoUN(
vr, it is worth noting that a giant impat rat datd to th oundary of
th Prmian and th Triassi has rntly n rportd from th northwst If th aus of mass
Australian ontinntal shlf, thus linking in tim a ig ratr with th iggst mass xtintions ar
mass xtintion. Howver, th laims for this ratr hv n tratd with portant turning poir
skptiism y th gnral sintifi ommunity, whih rgards th strutur vnts is that groups
as mor likly to of volani origin, and it rmains to sn whthr nan in th aftrnra
thslaims gain ptan. mammls in th ft
If th link twn impats and xtintions is gnrally wak' th sam for this mass t
annot said for giant volani provins. Th surfa of th ath is markd ially living in th s
by svral suh provins that ar omposd of layr upon layr of asalt nosaus Wr mam
lava flows that typially d 1 million ui kilomtrs in volum. Thes fornrs w s tody
ruptd from long fissurs or raks, and individual flows oftn dd nosaurs may also ow
1,000 ui kilomtrs in volum' a figur that is two or thr ordrs of mag- pard in th Triassi
.Within
nitud largr than thos sn during eruptions in historial tims. th yrs of thir histor}
past 10 yars improvd ag dating of ths volni provins has rvald trrstrial land ani
two rmarkal fats. First, most of th provins appar to hav ruptd in grops. Only aftr tJ
lss than 1 million yars; a gologial link of an y. hus th nvironmn- dinosaurs quikly ris
tal onsquns of suh ruptions would hav n onntratd into a In fft mass t
short intrval. Sond, many of th volani provins hv n found to into th vaatd ni
prisly oinid with tintion vnts. Indd, it now appars that vry of this volutionary
xtintion vnt in th past 00 million yars, inluding th mass tintions, way from ommuni
oinids with th ruption of a giant volani provin. This inluds th n in th rats of;
K-T vnt, whn th Dan Trap provin of India was formd. Howvr, oans ompard
not vry giant volani provin oinids with an tintion vnt.In othr highly sussfulgr
words, ther ar mor volani provins than thr ar tintions. ovi- shlls. Thy suffr
ously, it would ni and nat if vry provin oinidd with a mss - idly within a millior
tintion, ut th history of th arth is dfinitlynot nat. wipd thm out). I
How an volanism aus tintion? uh rsarh has fousd on th of millions of yars
limati fftsof th two major volani gassassoiatdwith th ruption tintion vnt also

l whyth arth ws su-
lowrsvry 26 million yars.
c and improvd ag dating of
lion-1.arintion vnts do
n vnt26 nrillion ys aftr
.tiOn vntsthat rmain show
ss tintion200 million yars
d y a smallr-magr-ritud
Jtrrassi.T h n t t i n t ion
r 80 million ars l a t r , a nd
h th K-T vnt.
t mtoritimpats hav so far
a lk of vidn.For xam-
lrio, nad, is roughly 707"
th largst ratrs known but
)n rats.Th ratr was formd
:r low tintion rats. F{ow-
'atrdtd t<lth oundary of
n rportd fom th northwst
n big rtr with th iggst
rtr hv n traltd with
y, whih rgrds th strutur
it rrrainsto L]sn whthr
rs is gnrallvwak' th sam
surfof th arth is mrkd
i of layr upon lyr of asalt
i kilomtrsirr volum. Ths
rdividual flows oftn dd
t is two or thr ordrs of mag-
; in historialtims.
,olaniprovins has rvald
Ll1Cs appnr to hav uptd in
f an y.Thus th nvironmn-
rav n ()nntrtdinto a
provins hv n found to
[d,it now appars that vry
i n l r r d i n gr h m s s x t i n t i o n s .
rri provin.This irlluds th
lf India was formd. Howvr,
ith an tintion vnt' In othr
.rn thllr tintions. Ovi-
.inoinidd with a mass '
Iynot nat.
h sarhlrs forisd on th
; sa s s o i t dw i r h t h
ss tintios 719
of asalt lvas:rbon dioxid and srrlfudioxid. As is wll known' rln
dioxid is a grrrhousgas' and th injtion of larg volums into th at-
mosphwill aus gloal warming. In ontrast, sulfur dioxid forms arosols
(y ration with wtr vapor) that lok sunlight nd aus ooling. Thus
th most notial fft of mdrn lrg volni ruptions, suh as th rup-
tion of Mount St. Hlns in Washington Stat in 1980, is ooling of th li-
mat. Hwvr, suh fftsar dinglyshot livd austh rosols
- ar rairrdout of th atmosphrin yar or so. Gologial vidn shows
that th ruption of giant volni provins is nrly always assoiatdwith
global wrming, suggstingtlrat th uption of aron dioid may th
mor inrportnt limati fft of volnism. owvr, it is on thing to rais
gloal tmpratur y a fw dgrs and ntirly nothr to wip out narly
ll lif on arth. Tlrrfor,many urrnt xtilltion snarios s th erup-
tion of flod aslt prvins s just triggr in a ompl hirin of vnts
that lads to a runaway grnhoussnario.
oNsoUENEs oF MAss ExINloNs
If th aus of mass tintions is still ativly datd, th onsquns of
mass xtintiorrs r aurrdantly lar. hy lrav long n rognizd as im-
portant turning points in tlr volutiorr of lif. An intriguing aspt of ths
vnts is that groups that wr insignifiant for th tintion ris to domi-
nan in tlr aftrnrtlr.Th quintssntialarpl of this is th suss of th
mammals in th aftrmath of th K-T tintiorr. amnrals had a long history
for tlris mss tintion' ut thy spnt this tim litrally and mtapho-
illy living in th shdlw of th dinosaurs. only with th dmis of th di.
nosaurs Wr mmmals abl to volv into tlr largr sizs and divrs ody
forms w s today. Somwhat pradoially, th initial suss of th di-
nosaursmay also OWsomthigto nlass-xtintionvnt.Dinosaurs first ap-
pard in th Trissi round 230 million yars ago' ut tor th first 30 million
.Within
th yars of thi history thy wr gnrally only a rlativly small omponnt of
trrstrialland animal omnrunitis,whih inludd m:rny othr ptil-lik
groups. only aftr th nd-Triassi mss tilltion 200 million vs ago did
dinosaurs quikly ris to total dominan of trrstrialommunitis.
In fftmass tintions provid lpportunitisfor the survivors to rdiat
into th vtd nihs lft hind y th xtirrtspis.owvr, th pa
of this volutionry radiatilnis highly inonstant and vris in an intrsing
way fron-rommulrity to ommunity' For ampl, thr is a distint diffr-
n itr th rats lf ror,ryof rarursthzrtliv in th wtr olumn of th
oans mpard with thos that liv on th safloor. Ammonoids wr a
highly sussful.roup'distantly rlatd to squid, that had attrativ spiral
shlls. T.hy sr.rffrd nulnrous xtintiolr vnts ut irlwys rovrd rap_
idly within a million yrs or so (xptfor tlr K-T tintion, whih finally
wipd thm out). In ontrast, lams that liv on th saflor oftn took tns
of milli<lns of ,varsto rovr from xtintion vnts. Th nd-rln -
u p t i o l l tintion vnt also shows n intrstingdihtltomy in th rovry rats of
720 rd Smith

plants and trrstril vrtrats. Plant ommunitis only gan to radiat

fatoris, h studid
and rturn to normal around 10 million yars aftr th xtintion vnt'
Haldan was th !7l
whras th terrstrial vrtrats wr divrsifying rapidly within 1 million
studnt of Haldan's i
yars of th tintion. In fat' th rovry intrval aftr th nd.Prmian
at UL without takin
xtintion was ptionally long. Prxtintion divrsity lvls wr not
of th Shool of Biolo
ahivd again until narly 100 million yars aftr th mass xtintion. In.
rmind for th rst
dd, for many safloor ommunitis th rovry did not vn gin until
part of this tim. As
8 to 10 million yars aftr th risis.
shool with onsidr
Th xtme lngth of tim it has takn for arth's osystms to rovr
hology, nuroiolog
should srv as a warning for th urrent man-mad mss-tintion vnt.
at Suss was a m
Gologial history rvals that th arth's osystms ar al to rovr from
world.
vn th most svr nvironmntal atastrophs, ut th timtal of ov-
Haldan rtd a
ry is sprad ovr millions of yars, a tim span that is immasuraly long
rnownd for th lar
from a human prsptiv. If th tim takn for ommnitis to rovr from
of thir knowldg an
modrn dgradation is of a similar duration, thn it is lar tht it is st to
munist for many yar
stop th damag from happning in th first pla.
1956 whil rmainin
B|LIoGRAPHY oth ativ nd su
N4aynard Smith's asi
tlvarez. L. \.. !(/.Alvarz. F. Asaro. and H. V. }ylihl.1980. xtratrrstrialaus
for th taous-Trtiary dan was irasil an
xtintion:primntalrsultsnd thortil
appliation.Sci208: 1095_1108. his ollagus. wat
Bkr,L., R.J. Porda'A. R. Basu,K. O. Pop'T. . Harrison,. Niholson'and tionally good at int
R. Iasky.2004. Bdout:A possilnd-Prminimpat ratroffshorof niority. H was rma
northwstrnAustralia.Sie4: 1469-\476. prtntiousnss.
Bnton,. J. 2003. Wh Lif Nerl Did: Th Grtstss titio of All
H startd his iol
Time. Lond,on:Thams and Hudson.
ourtillot, . \999. uolutiortstrlphs: h Sieeof sstitil. Drosophil subobsu
Cambridg:amridgUnivrsityPrss. volutionary studis.
Hallam, ^.204. tstrophsd Lesserlmitis:b ussof ss mad two notal or
tintios.oxford: Oford UnivrsityPrss. th ffts of inrdi
Hallam' A., and P. B. Wignall. 1997. ss titiosd Thir ftermtb.
D. sbobsur, whih
oxford: Oxford UnivrsityPrss.
Looy, . v.,.!.A. Brugman,D. L. Dilhr,and H. Visshr.1999.h dlayd ltion y fmal hoi<
resurgnof quatorialforstsa{trth Prmian-Triassiologirisis. twntith-ntury vo
Prodigsof the Ntiol dm of SisUSA 96: 1857_\862. til Thor lf N
Wignall, P. B. 1992,h dy th world narlydid. N Sitist,jnuary 2.5' rntly fashi<-lnalgo
51-55. Maynard Smith also r
2001. Larg ignousprovinsand masstintions.rth-SiRuit't,s
-P.B.W. aging. H providd a
53:1-33,
prodution and ota
aging' Drosophil is n
aynad Smith, John (1920_2004) lins of rsarhwr
muh rognition as t]
John N4aynard Smith Was th leading British volutionary iologist of th dvlopmntalgnti
sond half of th twntith ntury. H dvlopd a kn intrst in natural
a prpattrn and Ala
history as a hild, largely without any hlp from adults. As a shooloy at
diffusion prosss'
ton ollg h am aquaintd with th writings of J. B. S. aldan,
Aftr moving to Sr
largly baus Haldn's outspokn Marism ws anthma to his tahrs'
work, spially in v
whos job was to train th nxt gnration of th British ruling lass. Aftr
ausHaldan, by v
studying nginring at amridg Univrsity and doing war work in airraft
ausof th urdn

nitis only gan to radiat
; aftr th xtintion vnt'
ring rapidly within 1 million
r r v a la f t r t h n d - P
ln divrsity lvls wr not
ftr th mass xtintion. In-
ry did not vn gin until
]arth's osystmSto rovr
mad mass-xtintionvnt.
tms ar abl to rovr from
s' ut th timtal of rov.
n that is immasuraly long
ommunitis to rovr from
n it is lar that it is st to
nl.1980.tratrrstrialaus
:ntalrsultsand thortial
v4.Harrison,. Niholson,and
impatratrffshorf
).
'tstsstitio of ll
Sicrf ss titio.
s: h ssof ss
'sd hir Aftrmth.
y'isshr.
|999.T dlayd
n-Triassiologirisis.
: ' sU S 9 : l 3 8 5 7 -l 3 8 6 2 .
N Sintist' January25'
intions.rth -Si Ruius
-P.BW.
volutionary iologist of th
lpd kn interst in natural
om adults. As a shooloy at
writings of J. B. S. Haldan,
was anathmato his thrs,
th British ruling lass. Aftr
Lndding war work in irraft
rd Smith 721
fatois, h studid zoology at Univrsity ollg London (UL), whr
.$ldon
aldn was th Profssor of Biomtry. H am a postgraduat
studnt of Haldane's and took up an appointmnt in th zoology dpartmnt
m i n at UL without taking his PhD. H lft UCL in 1965 to om th first dan
of th Shool of Biologial Sins at th nw Univrsity of Sussx, whr h
rmaind for th rst of his lif' although h was dan during only a small
part f this tim. As an administrator h was sussful in dvloping a
shool with onsidral rsarh strngths, spially in xprimntal psy-
hology, nuroiology, and population iology. For many yars his own group
at suss was a ma for visitors and postdotoral sholars from around th
world.
Haldan xrtd a liflong influn on aynard Smith. Thy wr oth
rnownd for th larity of thir lturs and writings, as wll s th radth
of thir knowldg and intrsts.Lik aldan, Maynard Smith was a om-
munist for many yars, ut h lft th party aftr th Hungarian uprising of
1956 whil rmaining roadly lft-wing in his politial viws. Thy wr
both ativ and sussful ommuniators of sin to th gnral puli, in
Maynard Smith's as through tlvision as wll as writing. Bt whil Hal-
dan was irasil and violnt, Maynard Smith was kindly and lovd y
his ollagus. H was a highly ntrtaining onvrsationalist and rvas xp-
tionally good at intrating with fllow sintists, irrsptiv of thir s-
niority. H was rmarkaly opn mindd ut nvr foold y nonsns or
prtntrousnss.
H startd his bilogial arr in th 1950s y working on th gntisof
Drosophil subobsur, whih Haldan's group had dvlopd as a tool for
volutionary studis. Apart from som asi work on gnti mapping, h
mad two notal ontriutions through this primntal work. H studid
th ffts of inrding on mal mating havior and rprodutiv suss in
D' subobscur' wl,c,ausd him to rogniz th signifian of sxual s-
ltion y fmal hoi of mats. This topi was largly ignord y most arly
twntith-ntury volutionary biologists' apart from R.A. Fishr in Th G-
etil Theor of Ntrl Seletio (190). H vn antiipatd th ur.
rntly fashional good-gnesthory of th volution of fmal mat hoi.
Maynard Smith also usd D. subobsur as a modl systm for th study of
aging. H providd an ingnious dmonstration of th survival ost of r-
prodution and otaind vidn against th somati mutation thory of
agiog. Drosophil ts now a major tool for th iology of aging. Both of ths
lins of rsarhwr yars ahad of thir tim and proably did not gain as
muh rognition as thy dsrvd. H also did som pionring work on th
dvlopmntal gntis of Drosopbila, stimulatd y Curt Strn's onpt of
a prpattrn and Alan Turing's thoris of pattrn formation by ration-
diffusion prosss.
After moving to Suss, Maynard Smith dvotd his tim to thortial
wrk, spially in volution, and gav up doing xprimnts. This was partly
ausHaldan, y whom he always flt ovrshadowd, had did and partly
ausof th urdn of uilding up a nW dpartmnt.His thortialwork
722 ard Smith

was haratrizd y th us of rathr simpl mathmatial mthods that opd' whih appli
onald a good dal of ingnuity in thinking up approahs to iologially Ths mthods ar :
signifiant problms. This sortimsmt with disapproval from applid mth- ioral ology. ayn
.
matiians' who oftn prfr to us ompl mthods to solv insignifiant uolutil nd th
prolms. Maynard Smith ontributd importantly to arly work on molu- luid and informativ
lar variation and volution, using th thn nwly proposd nutrl thory as Aftr formal rtir
th asis for svral important puliations. Th most influntial of ths is Brian Spratt'smiroL
with his SussollguJohn Haigh on gntihithhiking (aynard Smith on molula variati
and aigh 1'974).This showd quantitativly how th spad of an advanta- work was vidnl
gous mutation rdus varition at linkd nutrl loi' y dragging a hro- atrial lls in natr
mosomal sgmnt alng with it. This onpt has lm vry important in ods for xamining th
rlation to th avalanhof data on ntural variaility at th lvl of DNA s- mmrs of atria
qrrns' sin a vally of rdud variaility an intrprtd as th signal DNA-squn lvl (
of a rnt fixation of an advantagousmutation. rmarkal ft of
Anothr fild to whih h md important ontriutions was th volution rputation as an ld
of sx and gnti systms' starting in th lat 1960s. is work hlpd fr N4aynard Smith al
this fild from its domination y th rathr woolly group-sltionist idas of tionary qustions' in
yril Darlington and Gorg Ldyard Stins, whih had had a svrly tion (th lattr trm
ngativ fft on its dvlopmnt. In partiular h mphasizd th prolm quilirium, and th
of th ost of sx, whry an othrwis sltivlynutral asxual variant wrot a sris of <
rising in a sual population xprinsa transmissiondvantagand rap- ogy, ulminating in l
idly sprads to fiation. Maynard Smith was th first prson to apprit 6rs Szathmiry dv
th diffiulty this poss for plaining th prvaln of sxual rprodution tions in volution, frr
among ukaryots. H mad numrous population-gntimodls of pro- th volution of lang
sssthat ould provid an advantag to s or inrasd romination' fts of msothliom
and h summd up th stat of th fild in his 1978 ook h uolutio of rsarh until his d
Se, whih is still th st survy of this sujt as a whol. Th study of th haratristially sti
volution of sx and rding Systms is now a flourishing disiplin within 2003 mting of th
volutionary iology, in no small part aus of aynard Smith's own work
|BL|oGRAPH
and his nouragmntof othrs.
A4aynard Smith's most influntial singl ontribution waS th dvlopmnt F i s h r ,R . A . l 9 3 0 . e
of th onpt of th euolutioril stbl strltg(SS),initilly in ollao. UnivrsityPrss.
N4aynrdSmith,J. 195
ration with th lte Gorge Pri. This flowd out of his lng-standing intr-
longvitof Dros
st in animal havior and his dsir to undrstand why animal onflits 1 9 5 8 b .S u a
usually do not nd in srious fighting. lt uss th prinipl that, for a trait 20_244.l-ondon
valu to rprsntan quilirium undr natural sltin,a nssaryondi- 1 9 7 8 . T h u
tion is that all possibl dviant trait valus ar sltivlydisadvantagous 1982.L|ulltt
UnivrsityPrss.
whn introdud t a low frquny into a population whr most individu-
1989.uolti
als hav th spifid trait valu. Dtrmining an SS provids powrful N4ynardSmith,J.' ..
mans of prditing th outom of sltion in ass whr frquny- volutionary dyn
dpndntfitnsssar gnratdy th iologial ontt, suh as s ratios ynard Srith,.|.,zln
or soial hvior. Although this pproah had n usd for for daling GetilRsr
with sx ratios' notaly y R. A. Fishr and.s7.D. Hamilton, aynard Smith Mynard Smith,J., an
oford: oford U
dvlopd SS thory xpliitly and applid it to many prviously intratal
volutionary prlms. It is n of th most sintifially fruitful appliations
of gams thory. A hug thortial and mpirial litrtur has sin dvl-

mathmatialmrlrlds that
up approahsto iologially
lisapprtlvalfrom applid math-
mthodsto solv irrsignifiant
tntlyt<larly work on molu.
vl proposd nutral thory as
.lr
mtlst influntial of ths is
i hithhiking (Maynard Smith
how th spadof an dvanta-
urral loi, y dagging hro-
h . r s o m V yi m p ( ) r t a n ti n
:iailityat th lvl of DNA s-
:n intprtds th signal
ol1.
ontriutionsWas th volution
l960s. His work hlpd fr
'olly group-sltionistidas of
ins, whih had had a svrly
lar h mphasizdth prolm
tivly nutral asul variant
ansmissionadvantagand rap-
, t h f i r s rp r s o n t o a p p r
vlnof sxual rglrodution
lation-gnti modls of pro-
. o i n r a s dr o m i n t i o n ,
s l978 bok /: uoltio o|
t as a whol. Th study of th
.a flourishingdisiplin within
of MaynzrrdSmith own work
rtriutionWas th dvlopmnt
z/egy(SS),iitially in ollo-
I o u o f h i s l l n g - s t n d i n ign t
drstand why animal onflits
s t h p r i n i I t h a t . f o
:al sltion,a nssaryondi-
ar sltivlydisadvantagous
lpultion whr most idividu-
rg an SS provids powrfl
ion in ass whr fquny_
g i l o n t t .s t r ha s s
ad bnusd bfor fo daling
/. D. Hamilton, Maynard Smith
; to tnanyprviously intatabl
:intifiallyfruitful ppliations
,iriallitrarrhas sin dvl-
rd Smith 72
pd' whih plis SS mhods tO many diffrnt iologil prolms.
Ths ntthods ar a mainstay of muh of th thory that undrlis hav-
ioral ology. Maynard Smith rviwd his ontriutions in his 1982 ook
uollltil nd th horl' of GL1ftls'whih is haratristiall rif ut
luid arrd informiltiv.
Aftr trmal rtirmntin 1985, aynard Smith startdto olloratwith
BiarrSprtt'srniroial gntisgroup' thn at Suss.on th analysis of dt
on molular variation an]volutiorr in tria. An inrportant rsult of this
work was vidn for muh mor hang lf gnti information among
atrillls in ntur than prvior-rsly livd.It lso gnratdnW mrth-
ods for xamining th ffts of infrqunt rominti<lnalhng among
mmbrsof atrilpopulations On pattrnsof variation and volution at th
DNA-squnlvl (Mynd Smith t l. 2000). Mynard Smith ahivdth
rmarkbl fat lf oming a ladr in a nw fild wll aftr stiriishinghis
rputation as an ldr statsmnof volutionry iology.
aynard Smith lso mad many important ontriutiolrst gnral evolu-
tionary qustions' inludirlg suh topis as group sltion vrsus kin sl-
tion (th lattr trm was invntd y him), syrnpatri spiation,puntutd
quilirir"rrn,
and tlr vllutionaryrol of dvlopmntalonstraints' H also
Wrot a sisof xllnt txtooks on various aspts of thrtial iol_
ogy, ulminating in his 1989 ook uoltior Getis. Lat in lif h and
i a t ors Szthm6ry dvlopd a st of spulatividas out th mjor transi-
tions in vllution'fom tlr volution f lif itsIfand th volution of lls to
th volution of languag.Maynard Smith suffrdinrasingly from th f-
fts of nrsothliomain th last two yars of his lif ut ontinud with his
rsarhuntil lris dath irr April 2004. is l:rstpuli ltu wils a rif ut
haratristillystimulating talk on his atrial work, at th Dmr
2003 mting of th UK Population Gntis Group in Susse.
|BLIoGRAPHY
Fishr,R.A. |930. h GtilThor1'of Nturl Seletion. Oford: Oford
UnivrsityPrss'
MaynadSmith,J. 1958a.h fftslftmprtur and of gg-layirrg
on th
- longvityof Drls4lhilsbrlbsur. .lourlof Getis 35: 82-842.
l958. Siralsltin. In S. . Brntt,1',
tltrof Dt'i,
t i t 20_244.Llnlon: Hinllrnn.
|978. h uolutiollllSe.rnridg: (iamridgUnivrsit1, Prss.
1982.ut'lluti<l d th horof Gmes,amridg:amridg
Univrsityrss.
1,989.utll til
r C ctis.ofrd: oxld UIri'"'rsity
Prss.
ynrd Smith,J.' . J. Fil,r-rdN. . Smith.2000.Populationstrutr and
volutilnary
dynarisof pathogniatria.Biosss22: 1115_I|22.
r a i o s MaynarlSrith'J., nd J. High. 1974.Thith-ikingfftof a favourl gn.
GtilRcsrh2: 2]_3.5.
Maynard Smith'.}.'and . Szthm:iry.|995. Th jrt Trsititls i uolutkln.
Oxford:Oford UnivrsitPrss.
724 r

tion. H also ws
Nlayr, rnst Waltr (7904_2005)
Ornithologists' Uni
rnst !altr N4ayr was a Grman-orn Amrin ornithologist who playd N4ayr's ontriut
a ntral rol in volutionary thory and systmatis during th twntith h fousd on th
ntury. H was a strong advoat of two idas: th thory of spiation thory. Sstmtis
y gographi isolation and th Biologial Spis onpt. ayr also tionally and am
mad important ontriutions to iogography, systmatis,and th study was part of a small I
of whol organisms during a period whn biology as a profssin am r- in population gn
dutionist. volutionary pro
N{ayr was born into a profssional Grman family. His fathr did in 7977, ialty, working los
and th family prind onsidral finanial troul during th 1920s. In approh a
Mayr's intrst in natural history strtd with his fathr. gan univr- H passionatiyadl
sity in 192' studying mdiin at th Univrsity lf Grifswald. Dspit that a population
high marks' N4ayr swithd to zool<lgin |925' H alrady had dvlopd ould our. In this
tis to profssional zoologists, of whih th most important wr with th making. on t|
rwin Strsmann at th usum of Natural istory at th Univrsity of onpt. This pla
Blin. Strsmann am Mayr's mntor. Undr his guidan Mayr om- spis.Th biologi
pltd a hD tn t926 on th uropan distriution and iogography of nisms and arrirst
th srin finh, Sris ri srius. Aftrwrd Strsmann hird ayr mhanisms istd
as an assistant. most important m
From 1928 to 1930 ayr plord Nw Guina nd th Solomon Islands, tio (\96) prsn
ollting for svral major musums. Aftr his rturn ayr sought a prma- mpirial and intll
nnt musum position' ut svralpromising offrs in urop faild to mat- ing volutionary stu
rialrize.Rathr than rmain in Brlin' Mayr aptd tmporary work at th nizd th Soityfo
Amrian usum of Natural History (ANH) in Nw York. H gan in uolutio (\947).
Jnuary 1931. In 192Walter Rothshild sold his orrrithologialolltion to vrsational sps l
th musum: 280,000 ird spimns'among othr things. Mayr was givn a ommon purpos ar
prmannt position to urat th nw matrials.This kpt him in Nw York. i\{ayr's impat or
nvr undrtook major fildwork again. N4ayr promotd vo
Whn th National Soialists ros to powr in Grmany, ayr hos not Arthur ain' among
to rturn hom. In 195 h mrrid Margart (Grtl) Simon (7912_1990), nar th spislv
and thy had two daughtrs. Grtl playd an important rol in Mayr's a- ipl of this approa
rr. Sh srvd as srtaryand assistantand also lookd aftr studntsand tionships and to us
visiting ollagus.In 1950 oth wr naturalizd as U.S. itizns. taonomi dision
ayr ontinud at th AMNH until 1953, whn h aptd an Agassiz dmd suffiint to
Profssorship at th Musurn of omparativ Zoology (Z) at arvard thought to adapt
Univrsity. H srvd as dirtor of te Z fm |961 to 1970. For ayr that mad a group
th arvard appointmnt Was a symoli hang in status. H was now a full mriai taonomy a
profssor, lik his mntor. yr am an mritus profssor at Harvard in Profssionally
1'975 and rmaind in amridg thraftr' with th Z as his bas. ology. On th on
As an ornithologist ayr spializd in irds of th southwst Paifi, struggld to onvin
aout whih h wrot marry thnial studis and svral fild guids. From dat in iologial sr
th 1930s ayr was ativ in th Linnaan Soity of Nw York nd on- struggld to draw tl
triutd to suprvising many dotoral studnts.H als mad onsidrabl luding volution.
ontriutions to svral ditions of Jams Ptrs'shek-list of Birds of th iology in th lat
World. Aftr Ptrs did in 1952. ayr suprvisd the beck-lisf's nxt di- bodis and othr gr
 r 725

tion' H also was i]tivin many profssionalsoitis,suh s th Arrrian

onithologists' Union.
'inornithologist who playd ayr's ontriutions to volutionary studis gan in th lat 190s whn
itratisduring th twntith h fousd on th proSs of spiation and mad ontriutions to gnral
d s :t h t h o r y o f s p i t i o n thory. Sstmtisd tbe rigi of Speis (t942) rivd prais intrna-
S p i s o n p t . M a y r a l s o tionally and arlra ornrstonin th Synthtithory of volution. ayr
hy, systmatis'ilnd th study Ws prt of small group of iologistswho thor:ghtthat rnt dvltlpnrnts
Llgvas a prfssiorrbam r- in population gl.ltisand iogography mad plssibla rigorotrs study of
volutionary prosss.During th 1940s h onntratdon this s his sp-
fmily.is fathr died i \917, ialty, working losly with Thodosius Dozhnsky, among othrs.
l i a l t r o u l d u r i n g t h 1 9 2 0 s . In approah Mayr mphasizd gographi isolation and polytypi spis.
h h i s f t h r .H g n t l n i v r - H prssionatlyadvoatdth thoy of allopatri spii1tion,whih rquird
zrsity of Grifswld. Dspit that population physially isoltd for divrgninto zrtrw spis
25. H alrady had dvlopd ould our. In this snsgographiallydistint populations wr spisin
l 1ost important wr with th making. On th sam lins Mayr hampiond th iologial spis
ll llistory at th Univrsity of onpt. This plad a prmium on intrbrdingand gn flow in dfining
J n d h i s g u i d a n a y r o m - spis.Th iologial spisonPt fousd attntion on isolting mha-
ribution and ioglgraphy of nisms rrd arrirs to gl1flow. Altlrough yr thtluglrtthr many isolting
rwd Strsmannhird N4ayr mhanisms xistd as spial ss, h lnsidrdgographi isoltion th
most important lnhanism in n:ltur. is ook iml Spies d uolu-
uina and th Solomon Islands, tio (1963) prsntdhis viws at thir most dvlopd. In addition to his
is rturn Mayr sought a prma- r-npirialand intlltulontrirrtions,ayr playd a ruial rol in shap-
offs irr urop faild to rat- ig volutionary studis into a onrmrrnity.Largly on his own, Mavr orga-
ptdtmporary work at th nizd th Soitv for th Study of volution (|946) and launhd th jlurnal
.lH) in Nw York. L{ gan in uoltio (1947\. h requird ir gat dl of ffort, ut thy prlviddon-
l h i s o r n i r h l o g i al l l t i o nt o vrsational spas for volutilnary studis and hlpd dvlop il sns of
o t rt h i n g s .M a y r w a s g i v n o m m o n p u p o s a n d p t a t i o n '
Lls.This kpt him in Nw York. ayr's impt on systmtis irlvolvd sverl lvls. As a thtlrtiian
ayr prornotd volutionr}.svstIltis rogthrwitlr Gog Simpstln and
'r in Grmany, a,vr hos not Arthur ain, arrrongothrs.This hd spial rlvnfor taxonomi groups
r (Grtl)Simon (1912-1990)' nar th spislvl:suspis,spis,and spisgroups. Th asi prin-
'n important rol in Mayr's a- ipl of this approah was to us lassifiationto xprss volutionary rla-
l also lookd aftr studnts and tionships and to us an undrsttrndingof volutionary prosssto mak
lizd s U.S. itizns. taonomi disions' For instan, tl.lprsn of gographi arirs was
, whn h aptdan Agassiz dmd suffiirrtto draw otrrrdrisin lassifizltions.Likwis' hratrs
, Zoo|og (Z) at Harvard thought to daptations to lol <lnditionsWr highlightdas th faturs
'.
{rom 196I to 1970. For Mayr that mad a group distint. This approah had its ritis. In th 1960s nu-
ng in status.H ws now a full lrial taxonomy and ladistiswr important rivals.
mritusprofssort Harvard in Profssionallyavr workd to inrasth sttt.ts of systmatiswithin i-
with th MZ ils lris s. ology. Orr th on hand, tlris rquird rform within tlre disiplin. Ma1'r
irds of th southwst Paifi, stuggldto onvin ollagusthat thy had rsponsiilityto kp up-to-
; and svrlfild guids. From dat in iologial sujtslik gntis,thology, and physiology. H also
Soity of Nw York and on- struggld to drw thm into ontriuting t gnral iologial sujts,in-
rts' H also mad onsidral luding volution. on th othr arrd,Mayr am politiall ativ within
rcrs,s hk-list of Birls of the iology in t lt 1940s. Whil h was t th Z' h loi1funding
rvisdth bek-list,snxt di- bldisand otlrr grops so thy would not ignor systemati iolgy whn
726 r

gav rdit to thos

prmotd th wori
thm on qual fo
iin and prim
As a philosoph
twn proimatl
intration of stu
proximat auss
work?'' In ontras
words, nswrs to
tion twn pro
,jrtA\ 1 9 5 0 s a n d 1 9 6 0 s

lLloGRAF

l B Bok, W. 1994.rns
vlution.Bil/
2004.rnst]
12]l:67_65|
ayr, . 1942' Sst
UnivrsityPrss
1 9 5 . t h o
Hill.
1 9 6 . i m
1982.Th G,
5 t r s m a n n 't ' ' 1 9 / 5 '
Hrvard Univrs

}yIm
Inhts Ssttltiltis
"td thtl rigi of Spis(1942I, rnst Mayr malmuh of
suh phnomnaas ..ringsof ras,''whr mmrsof touhingsuspisan Th trm meme \^/
intrdut whn th nd populatitlnsirl k trdttlulr,lh. is ..o- ish G t<ldsri
dutivisolation.Hr w sspisir-rth mkir.rg. If th middl populations r units f iologi
(of th warlr PhIklscopustrohiloids)wr Iiminatd,w w.,ulj huu.
popularizd, as vi
two sparatspis'Th suspisin th ring'ar = uirids;I,=ldlotui;
th spawning of a .
T=trohiloids;:obsrtus; andp:plmbitrl,ls.Th rss.hathdra'
in th distrit twnth wstrnS:ryanuntins irnd tlr YnisiRivr, is graphs. ms an
where ui r i dlzsanl p lu it r sus.,u..lop. lifs, hairstyls,
mms to srv as i
any systm witlr di:
tion), sltion' an(
supportirrgth fild. His vision for systmatisis st prsntdin ethods Lik organisms,id
d Prinipls of S1t57g7nti
Zorlllg1,(195), rvhilr aln standrd train. ally inundatd with
ing manual. (othrwis rv wtlul
In th sotld hlf of his Iif, ayr Wrot tnsivIyon th lristory arrd with h thr fr
philosophy of io|ogy, spially on thms rlatd to natttral history and and get rpliatd,
volutiorr. His ook Th Groth of Billogi"ll Thlght (19s2I was a signifi. through imitation, i
ant ontriution to this topi. Drwir-r m l.]nlmportant sr.rjtfo sions t th dinnr l
ayr' as did many othr ky volutionists of th past 100 yars (.g.'Karl lt is important n
Jrdan). ar's ontritionss histlialrntrdon two thnrs.First, h th idas. For m;

12),rnst Mayr mad muh of
l r s t l f t t l u h i n gs, u s p
lak nd touh' thr is rpro-
l i n g .l { t h m i d d l p o p u l t i o n s
liminatd, w would hav
are = uirids; L: ldlotui;
itrus. T ross.hathd ara,
tinsalrd h YnisiRiv, is
s is st prsntdtn thods
whih ama standard train-
xtnsivlyon th history and
rltdto natural history and
tl Thought (1'952)was a signifi-
]am an importnt sujt for
lf th past 100 yrs (.g.' Karl
:ntrdon two thms. F.irst,h
me 727
gav rdit to thos who dvlopd th idas h himslf dfndd. Sond, h
promotd th work of naturalist traditions in th history of iology' plaing
thm on qul footing with othr traditions in th lif sins' suh as md-
iin and primntal iology.
As a philsophr Mayr is hifly rsponsil for promoting th distintion
twn prximat and ultimat auss. By invstigating th opration and
intration of strutural lmnts, h said' xprimntal iology fousd on
..how'' qustions, suh as ..How dos this
prximat auss. Thy askd
work?'' In ontrast, volutionary iology sought ultimat auss' in othr
..why'' qustions. As did his historial writing, th distin-
words, answrs to
tion twn proimat and ultimat auss aros during onflits in th
1950s and 1960s twnvolutionary and molular ilogists.
BlLloGRAPHY
Bok, W. 1994. rnst ayr, naturalist:is ontributionst systmatisand
volution.Billogd Philosopb9: 267-327.
2004.rnstMayr at 100:A lif insidand outsidof ornitholog.Th uh
121:637-651.
ayr, . 1942.Sstmtisd th rigi of Speis.Nw York: olumia
UnivrsityPrss.
1'95.thodsd Priciplsof SstmtiZoolog. Nw York: MGraw-
Hill.
1963.Aiml Spiesd uoltio.amridg,MA: BlknapPrss.
-hoght.
1982. h Groth of Biologil amridg'A: Blknp Prss.
Strsmann, . \97 5. ritholog:From Aristotl to th Preset.amridg,MA:
Harvard Univrsity Prss. -l..
m
Th trm mm was oind y Rihard Dawkins in is 1976 ook The Self-
i s n
ish Ge to dsri units of ultural volution (as opposd to gnes,whih
ar units of iologial volution)' In th intrvning yars th trm has n
popularizd, as vidnd y its inlusi on i he oford glish Ditionr,
th spawning of a ddiatd (and now-dfunt) journal, and svral mono-
graphs. Mms an suh things as songs' tools' styls of drss, rligious
lifs, hairstyls, aphorisms, and myths. Dawkins originally mployd
..Univrsal Darwinism',-th thought that
mms to srv as an instan of
any systm with disrt units that hiit diffrntial fitnss vals (varl'a-
tion)' sltion,and rpliation (hrdity)will volv y Darwinian mans.
'!7
Lik organisms, idas appar to hibit ths haratristis. ar ontinu-
ally inundatd with idas. Thankfully, most of this information is disardd
(othrwisw would hav mmory ovrload). Thus idas ar in omptition
with ah oth for survival within our minds; th fit ons rsid in our minds
and gt rpliatd, whil th unfit ons gt disardd. Rpliation ours
through imitation' it through th writtn word, dmonstration' or disus-
sions at th dinnr tal.
It is important not to onfs mms' whih ar idas, with th rfrnts of
th idas. For ampl, th mm rro|Uhdshrpnig is the id of a mannr
728 dl

of produing an arrowhad, not th rrowhad irslf. his distintion pr- Th stain of studv
srvs th gnotyp/phnotypdihotomy witi iology, or, in Dawkins's ausd outs of sv
trminology, th diffrn twn rpliators and vhils.Rpliators ar This prinlay b
th ntitis that opy thmslvs and od fr struturs that intrat in th of Saint Thomas in I
.,from th bittr strr
world (som thorists prfr th t,erm itrtor to uhile). This :liss sv-
ral points that distinguish mmti volution from ompting onptions 1 9s 4 ) .
of ultural volution. ultual volution is oftn sll s simply improving th At th monastl.
fitnss of th goup or group tnms.For xrlpl,fishhooks ad arrow- oftn-sad tasks of ;
hads volvd to onvy a fitnss advantag to th groups and individuals for.rndhim work as z
who usd suh artifats. ms, howvr, an quit sussfulwithout vr, rquird an
onvying fitnss advantag to thir hosts, ven to th point of birrg dl- oth tims. Aftr tlr
triolls to thir vtors. Dawkins famously lims tlrat rligion is a perriu- Vinna to obtain d
larly sussfulut toxi mm. Mms ar fit wlrn thy hav high fidlity th pratial dn-ron
and high feundity unto thmslvs.Aording to proponents' lms hold lnost rnt wrk in
impliations for our onption of ultur and human minds. N4ndl's talrr
Th srrssof th mme mm' howvr, lis its suss as a trm of art Vinn from athol
in th aadmy. Critis hav idntifid svral problrrrswith mms, nt th ing ndl's tim ir
lst of whih is that it is simply stitlgan old id il-lnw pkaging. Pr- hav krrown of this
aps it is not th mms thmslvsthat ar xplirnatory ut th undrlying trs aquird y plal
psyhologial mhnisms rsporrsilfor th sltion and rpliation of ral loals. Latr
tain mms tht ar usful for xplaining ulturl voiution. Furthrmor, suport for th inh
it is unlar tly what a mm is. It is asy to oll up with xanrplsof l21ntransmutatlons.
mms' ut a lar irnd aptd dfinition rmains lr"rsiv. For xilrrrpl,is possibiiity m11tiO
a mm a part of Bthovn symphony or th rrtirsynrphorry?Until an otanistsJosph KoJ
operational dfinition is found, tl.lfruitfulnssof this onpt rtllins qus- sking to stlish
tional. Finally' som hav argud that th lk of fidlity of mms, as wll hyridization.
as thir smingly nondigital natltr' srrggstsa Lirmarkian rathrr than Having rturnif
Drwinian volutiorrrypross. Th tru impat <lfmms for undrstand. tst varitis of tlr
irrg ultural volutin rmains lagly urrdidd. lryridizatitl11pr
im, h plairrl, u
BIBLIOGRAPHY
hyid progny pp
Blakmor,S. 1999.The m hi.Nw York: Oford UnivrsityPrss. tion with rtainty,
Dawkins, R. \976. h SlfishGe, ()ford: Oford UnivrsityPrss. drtak this progril
Dnntt,D.1995.Dttt'i'sDgrousIt!,I-ondon:Prrguin. -].Z.
hzrratrdiffrn
to hiv stirtistia
..fr.rahing'
was
ndl, Grgor (t822_884) .
portant qlrstion:
..th prist who hld organi forms'' (St
Grgor Johann ndl hs n lld th ky to vo-
lution'' (isly \959'205), ut in lris own day fw iologistshad vr hard th ould forr th
..rvrtd
of him, hrls Drwin inludd. His faril was poor' ut h souglrt sup- turnd or
Port in a monasty whr fortun shon upon him y giving him tlr mans vidntly this rva
to arry out his rmarkal and aritiousxprinrnts. tsting his rsarh
lvIndl, th firstorn of Anton ndl and Rosin ndl (n pulishd. Not unti
Shwirtlih), ws isd in Grmn-spaking Silsizl in th littl vill:rg o{ rigi of Spis.
inzndorf. At 11 lr ttndd slrols far from hom, and at 18 h rrtrd sgs prsntingrr
th Philosophial Institut in Olniitz, ompltil-tgl]is ourss th in 1843. was attaking a tar[

ad itslf.T.his distintion p-
ithin iology, or' in Dawkins's
rrsand vhils.Rpliators ar
|r sttlturstlrat intrat in th
. t o r t o u h i l e ;T. h i s a i s ss v -
,n from ompting onptions
n snas sirnply improving tlr
xampl,fishhooks and arrow-
l to th groups and individuals
:n b quit sussfulwithout
vn to th plint of ing dl-
l a i n l sr h t r l i g i o n i s p r t i u -
fit whn tlry hav high fidlity
ing to prponnts, Inms hold
d hu-ranminds.
lisits sussas a trm of art
lprolms with mms' not th
old ida in nw pakging. Pr-
planatoryut th undrlying
h sltion and rpliation of
;r.rltural volution. Furthrmor,
iy to om up with ampls of
. m a i n s l u s i v .F o r
. rh ntir symphorry?Until an
l s sr ' f t h i s o n p tr m a i n s q u s -
Lakof fidlit1.of m.ls,as wll
its a Lamarkian rzrthr thzrn a
npat of mms for undstand-
dd.
rk:oxiord Univrsit,vPrss.
'fordUnivrsityPrss.
rdon:Pnguin. -.J.Z.
pristwho hld th ky to vo-
ay fw biologistshad vr hard
y Was po()r' ut lr sought sup_
on him y giving him th mans
prlmnts.
dl nd Rosin ndl (n6
rg Silsi in th littl villag of
ror hom, ad at 18 h ntrd
ltinghis ()urssthr in 1843.
dl 729
Th strain of study plus tutoring to provid his own funds had rnanwhil
ausd otsof svrdpssionand tndd tim k honr to rovr.
This prin iay hind his dision to apply to th Augustinian Monastry
of Saint Thomas in Brtinr-r.H knrv, h plaind, that h nddto frd
..from th ittr struggl for istn''
that hd bn his lot hithrto (lltis
t9 s4\.
At th monastry r-rdlprt>vdunal to op motionally with th
oftn-sd tasks of a prist, ut his aot' th undrstnding yril N"P,
found him work as a sustitut tahr inZnaim. Tah rtifiation' how-
vr' rquird an xamination. This lr took twi but though stssfild
oth tims. Aftr th first attmpt th aot snt Mndl to th Univsity of
Vinna to otairr adqut foundations irr th sins.Thr h assistdin
th prtial dmonstrtionsfor th ph,vsisourss arrd was posd to th
most rnt work in plarrt ytology.
ndl's tahr. th otanist Franz Llngr' Was a vitim of attaks in
Vinn from athlli quartrsfo his advoay of spistransllltationdur-
i n g M n d l ' s t i m i n V i n n a ( 1 8 5 1 _ 1 8 5 3 )a n d s u s q u n t l y
. ndl must
hv known of tlris nd of Unge's transplant xpimntsin whih hr-
trs aquird y plants in novl lraittswr lost on rturning to thi natu-
ral loals. Latr ndl rpatd suh primnts and found in thm n
support flr ti-rinhritanof aquird hratrs,thus ruling out Lamark-
ian transmutations.Spismultipliation y hyridization, howvr, was a
m p l , i s pssiility mntiond y Ungr, for h knw th wok of th Grmn
otanistsJosph Klrutr and rl von Girtnr' who had hyridizd plants,
sking to stalishwhthr spisrmain onstant or an hang through
hyridization.
Having rturnd frorrrVinna in 185, Mndl gan th following yar to
tst varitis of th dil pa for onstany of typ in prparation for th
hyridization primnts that oupid him twn 1856 and |862. Ifis
aim, h plaind, was tO ,.dtrminth numr of diffrntforrns in whih
hyrid Progny appar' prmit lassifiationof tlrs forms in ah gnra-
tion rvith rtilinty, and asrtzrinthir numril intrrltilnshi1rs.'' To r.rn_
drtak this program h hos varital diffrnsadvrtisdy nonlnding
harat diffrnsthirt ould l-llrly distinguishd' Awar of th nd
to alrivstatistiallysignifiantrsults,he grw larg populations' Th task
..far-rahing,''
was ut h warnd tht it was th only way to solv th im-
portant qllstion: ..th
dvlopmntl history ftiklugsgeschihte]of
organi forms' (Strn and Shrwood 1966, 2). If som hyrids rd tru,
thy ould form th asis for th dvlopmnt of nw spis'ut if thy r-
turnd o ..rvrtd''to th origirrtingforts, no rrvlywould rsult.
vidntly this was th qustion h st out to solv in 1854 whn h gan
tsting his rsarh mtrial. rls Drwin's thory Ws at that tim rrn-
pulislrd.Not until |862 did N{ndlrd th Gman tanslationof th
origin of Spis. ndl's most frqunt marginali thrin fr to pas-
sagsprsntingnd opposing tlr dotril of spial rirtin.H Drwin
Was attaking targt rht Mndl, it appars,did not pt ithr.
70 dl

Th ontt of Mndl's famous primnts Was thus not Darwin's thory prsnt in th hy
of volution. It was instad th dat among Grman-spaking iologists' sv- lls ar ithr of
ral of whom, lik Ungr, had spousdlmntsf th spulativtahing ominations of p
of Nturphilosophi, prinipally th notion lf th sussivdvlopmnt of ny of th two kin
highr and highr forms of lif ovr an tndd priod of tim. On of th tios that h found
Augustinians in Briinn, Matou Kl5l, also supportd a dvlopmntal viw of gg lls of th sar
spisorigins' ut unlik his frind Mndl, Kl:il was a spulativphilos- th old ida of on
ophr who nvr arrid out a rigorous sris of primnts. Mndl was a nw variant will
drawn hifly to th xprimntal litratur of th Grman hyridists. His hif was also wrong. N
rsour was arl von G;irtnr, who, in his ook F,periments d bseru- dirtly rlvant t<
tios of Hbridiztio i th Plt Kigdom (Vrsuh ud Belbbtugen Darwin had found
br di Bstrdrzgug im Pflzereich' |849), found that frtil hyrids rgation is thus s
yildd progny that tndd soonr or latr to rvrt to on or th othr of th Also from sgr
originating spis. Gzirtnr listd oniy a fw hyrids that appard to rmain in ll sorts of o
onstant in thir offspring, nd it is signifiant that ndl rprodud many pndntly of n :
of ths hyrids and found that thy did rvrt. ndl wnt on to hoos tion with this pro
hawkwds (Hirim) aus of th multitud of forms found in th wild. laim that domst
Although th otanist arl Nigli nouragd him in this dirion, it was not orrsponding wil'
h who prsuadd ndl to hoos this diffiult gnus' as is oftn assrtd. lif.'' Instad, h o;
Mndl rasondthat polymorphi gnramight ow thir multituds of spis Th aptan
to hyridization' ut of a diffrnt kind from that h had found in his lassi adhrn to nonb
xprimnts with th dil pa (Pisum). rativ rol of nat
ndl plaind th rsults of is Pisul primnts y suggsting that lnding stp' thr
thr must a pross of sgrgation (s figur) of th diffring potntials frns littl y li
gnetisinto Darw

|BLIoGRA
dtrN RuD Y.{.Lgt,tl{'flNLD
Batson'!' l 909. ,\
UnivrsityPrs
i s l yL, . 1 9 5 9 ,D
London:Vitor

YR q G:irtnr,. von.184
P|lzerih' S
Iltis,rs. H. 1954.(

fl.#w fliP)
2I*24. Reprt
hiago:Univ
oly, R. 1985.orig
orl, V. 7996' Grg
Strn,.' and . Sh
Blo&.Sanrn

,=.
.siling,
F. 1991.il

P.P|f-l
dil Gti
www.Mndlw.
nglishand Gr
othrs'th follo
Pul' D. B., and
lfilliam Batsonws ndl's mouthpiin Britain in th arly yrs of th prati,1900
twntithntury.his diagram shows his rPsnttionof Mnd['sfirst law,
-l
th law of sgrgation.(From Batsln 909.)

Wasthus not Darwin's thory
:rman-spaking iologists'sv-
nts of th spulativtahing
th sussivdvlopmnt of
d priod of tim. On of th
ortda dvlopmntal viw of
.16l ws a spulativphilos-
; of xprimnts. Mndl was
r Grman hyridists. His hif
,ok primnts nd bsrut-
(Vrsuh ud B lb h tuge
849), found that frtil hyrids
'vrtto onr or th othr of th
y r i d st h t e p p a dt o r m a i n
tht Mndl rprodud many
rt. Mndl Wnt on to hoos
rd of forms found in th wild.
him in this dirtion, it was not
ult gnus, as is oftn assrtd.
t oWthir multituds of spis
hat h had found in his lassi
primnts y suggsting that
1ur)of th diffring potntials
Y(.Lltl {'Rllv|<L}
&w'@
q G:irtnr,. von. 1849. Vrsuheud Blbhtug
rfl&
I'
qr dw
& t
YR Yq
Britainin th rly yars of th
.sntationof Mndl's first law,
edl 7.1
prsnt in th hyrid plant whn th grm lls ar formd suh that gm
lls ar ithr of on kind or th othr, not mid. Thn if in frtilization all
omintions of polln lls and gg lls ar possil' populations of prog-
ny of th two kinds will yildd that will approximat to th statistialra-
.!hr
tios that h found in his primnts. unions r twn polln and
gg lls of th sam kind, oth originating forms ar rovrd pur. Thus
th old ida of on taintd, always taintd Was Wrong. Th assumption that
a nw variant will simply dilutd y rprodution with th normal form
was also Wrong. Nw ominations of haratrsan thus prsist. This was
dirtly rlvant to th prolm of th slmpirxgffect of outrding that
Darwin had found dunting for his thory. Mndl's onpt of grminal sg-
rgation is thus sn in rtrosptas th ky that Darwin lakd.
Also from sgrgtionflowd th possiility of omining hrditry trits
in all sorts of ominations and dmonstrating that th traits hav ind-
pndntly of on anothr. ndl idntifid th sour of hrditry varia-
tion with this pross of rcombitio. Trefor' h ojtd to Darwin's
laim that domsti animals and ultivatd plants ar mor varial than th
..hangdonditions of
orrsponding wild spis us of th fft of
lif.'' Instad,h optd for th gratropportunitis for ross-pollinations.
Th aptanof ndl's work in 1900 am along with an unritial
adhrnto nonlnding haratrs' thus promoting skptiism toward th
rativ rol of natural sltion.If nw hratrsar produd in on non-
lnding StP'thr is no spa for natural sltionto aumulat slight dif-
frnslittl by littl. This matrially dlayd th intgration of ndlian
gntisinto Darwinian thory.
BIBLIOGRAPHY
Batson,!. 1909. dl'sl,riiplsof Hrdit' amridg:mridg
UnivrsityPrss.
isly,L. 1959.Drtai,stur:uolutio d th Who Dislurdlt.
London:Vitor Gollanz.
Pflzreih.Stuttgart:K. F. Hrring.
br di Bstrderzugug
il
Iltis,tr4rs.H. |954. Grgorndl'sautliogrph. Jorlof rdit45:
21-24. Rprintdin R. oly, rigisof edlism,2nd d., |96_198.
hiago:Univrsityof hiagoPrss,1985.
oly, R. 1985.origis of ndlism.2nd d. higl:Univrsityof higoPrss.
orl, V. 1'996.Gregordl:Th First Gtiist,Ofrd: Oford UnivrsityPrss.
Strn,., and . Shrwood'ds.1966.Th rigin rlf Gtis: dlSour
BooA.San Franiso:.W. H. Frman.
Wiling,F.1991. istorialstudy:JohannGrgorndl, 1822_1884.llurl of
dil Gtis40: 1_2.5.
www.ndlw.<lrg. Inludsth nglishtt of Mndl's186.5papron Pisumin
I nglishand Grmanand .,ssaysand ommntary,,' whih ontins,among
othrs,th follwingssays: Oly, R. 1997.ndl,Mndlismnd gntis;
: Paul,D. B.' and B. A. Kimmlman.1988.Mndl in Amria:Thoryand
pratie'190-1919;and Sapp,I.1990. Th ninlivsof GrgorMndl.
I -R.O.
i
72 iber

ologial lationsl
Nlihnr,harls D. (b. 1918) u n d r s t a n da n d i n t
Chrls Mihlrer, ntomologist at th Univrsit1'of Kanss, is th world's ount of th nu.l
lading prt ()n th lassifition,morphology, natural history' and volu- nr as ..irquit' g
tion of s,spially th divrs slitzrryand priritivly soirrlbs whos polmis...[utli
iology olltrsts with tlrt tlf th familiar horrylr(pis tlliierl). tc_ might s;l dadly.''
nr has usd his road kr-rowldgof ths insts tO tst idas aout th Mihnr's ordia
volution of inst soitisand to hlp modrniz th sinof taxonlmy. man of sin,sp
trr-rdhis marr studnts lrav trvll tnsivh. in th Ur-ritdStats, h has ondLltdfil
Afria, Asia' urop, nd Latin Ami to tlllt nd osv s irr thir ntomologists in mu
natural haitats.As a rsult of this work th sh:rvlm modl tatln yrs Mihnr Ws
for omprativ studis of volution. Sins,on of onl1
ilrnr's rsrh is a prmir xrpl lf how volutionr1,iology uss lristory whn h wa.
ullparativ studis of lrvior and mophology ttl undrsttrdth origin of Soity for th Studl
nw trits nd tlr divrsifiation lf spis.Baust>fihrrr'sw<lrkand his pothsis tstingand
nOuragmntof young rsrhrs,thousands of spis lf s n road knowldg of
idntifidwith rtinty.This satisfisth first rquirt-nnt of ltlrpaativ-
sr: aility to idntify tlr organisn-rs. In additiolr, ihnr's work trats th BIBLIOGRAP
phylognti rlationships of diffrnt groups of s' summarizs knowldg H u ] l ' D . L . 1 9 8 8 'S l
of thir iology, nd plains how to rr and osrv thm in th laratory. M i h n r , . D . l 9 6 '
ilrnr'sm<rrthan 400 r.rlitions, inluding two maior bl<lks on s, 12:1.51-172.
|/. lnL'J|)
a a vrital nylopdi t>f inforr-ntirn that has n ploitd by Nlih-
200.h ts
nr and othrs to tra th volution of suh phnomna as parsitism and th '!7st-rhrd,
M. J. :
transition from solitry to soial irr group-livinginsts.His taonomi insight I n M . . [ .R n , d
has nrd him ldr in th tlrorv anl prati of s),stmatis,tlr str'rdyof Smithsniirn Inst
lilssifiationardrlationshipsamong groups of organisls.At th ag of 81,
Mihnr pulishd his magrrum op.he Bs lf th World (2000), a 91-
lvlillr, StanlyI
pag tom that trats 1,200 gnra arrd sugnriland mo than 16,000
spis irr a r"rnifidlassifitiorrd surmrizsmulr of th urrlrt informa- Stanl,vL. Nlillr sp
tio on this implrtant group' insts tllt ar ssrrtiltl ologil and human vrsity of alifrnia
ommunitis as pollintors of flowring plnts, inluding many rops. rakthrough xpr
ihnr's intrstin s gan whn as oy lr gan to Ollt insts sarh of pioti
nar lris hom irr southrn aliforni' l-rtlurag1 y lris moth, an al- gthr lvirh lan1'
tur naturalist, and by P. H. Timrlk, an authority n taonmy, h stps and hmial
pulishd his first ssay on s at th g of 16. FIis ontagious nthusiasm knlrn,ldgrnnt of his
for s has inspird a larg numr of studnts and ollagusirnd has put of-lif fild, ln \97 |
t Univsit of Kansirs On th map as a entr for rsilhand sil-t- to th Natilnal Aat
tifi ladrship. Mihnr also wrrt importent ssays on th taonom of his rol in th r:rti
uttrflis,although ths ar now lss known than his work on s,and h of t,if (ISSOL), th
Was at th ntr of ontrovrsy during datsovr numril tironomy in illr Awrl to
th l950s nr'] l960s. Alog with Rort Sokl, 'hos vilvs wr sotll_ In l9-52-1953,..v
w h a t r o r r a d i a l , M i h l l r a d v o a t d t h u s o f o j t i v ,q u a n t i t t i v and Nol lauratF
dat in lassifitions (meril tolm) as an antidot tO som of th to simulat th ond
ovrly intuitiv rrd lss sirrtifipr<ldursf tlr tim. But h also - primnts h was
l i v d t h a t q t l a n t i t a t i v l n t h o d s . . t l l t t s tn o t l l o w d t o s P a r a t t h s s - on th trlrgnof
. . A l k s a n d rI . o p r :
tmtist from the organisnls thmslvsfor only v.knowing tlrir hvior,
 illr 7

tlltlgialrlatioslrips,th furrt,ionsof thi r,rious stutI'lrs't., an lr

ttr-rdrsttrd :rrrdintrprt tllx()llonli data '' (Nlilrnr l96, 170). In al1 a-
.sity tlf Kansas, is th world's o n t< l ft h n u t l r r i i r l - t a x o n < l m wya r s , H r - r l l( 1 9 8 8 , 1 l 8 ) d s r i d i h -
..a
4v,natural history, nd volu_ rnr as quit' !]ntlman . . . who dlsrrot njoy partiiPting in rauous
pimitivlystlil s whos p l l r n i s . . . [ u t l i n s i r r t i f i o m a t [ h i s I a l s o h i g h l y f f t i v ,s o m
lv /pls mllifr). Mih- might sv dadl.''
llststo tst idas aout th ihnr's ordial, unasslttrringmannr lrs mad hinr n fftiv stats-
.nizth sinof taonomy. mn of sin'spillyin ountris lik Jpan, io, and Brazil, whr
r-rsivly in th Unitd Stats, h hs ondutd fildwrk and promotd ntomologial rsarh;and mong
'lltnd osrv s in thir ntor-ologists in tusums nd laoratoris troughout the world. For marry
s hv <lrrr a modi tzrxon yers Nlihnr Was tlr onl Knss mmr of th Ntiorral Aadmy of
Sins,on of only two Karrsansto hav joind th aalmyin its 184-yar
hw volutionrrryiology uss history whn h ws ltd in 1965. In1967 h srvd tls prsidnt of th
|!]vto undrstrrdth origin of Soityfor th Strrlyof volution. Mihnr's arr shows th valu, for hy-
ruslfMihnr.swork and his pothsiststing and for th undrstandingof iologial divrsitv,of dp arrd
;andsof spisof s zrn b road knowldg of a partiLlli1rgrlup of orgnisms.
q u i r m ntf o m p a r a t i v r -
ion,Milrnr'swork trats t B|LloGRAPY
lf s,summrizsknowldg Hull' D. l-. l988. Sills Prtrss,hiag:Urrivrsity
rlf higPrss.
lsrvtirrnin th lorz1tory. i h n r , . l ) . 1 9 ( l . ] . S o m f r t t L l r d r . l l m n t s i n t i l o n 6 m y . S , , s t m t i Z l r > l t g 1 ,

rdingtwo major ooks on s' 12:151-172.

|974, h SlilBs' mridg' MA: llrvad Univrsity Prss.
rt has n ploitd y Mih_
20. h Bes llf th W;r/d. Bltimor:Johr.rsopkins Univrsity Prss.
lnomnaas parasitismand th Wst-rhrd,. .l' 200.l. Th inl;lortnof taon-ntrdrsrhin iologv.
4 irrsts. His tt>nomiinsiglrt In N{' J. Ryn, r'1.,
Aur lnttltlliti,-7,.sshington,D:
i of systm:ltis,th study of SrnithslninIrlstituionPrss. -..r.w._.
f orgnisms.At th ag of 81,
s of th World (2-0), a 91-
illr, StanlyL. (1930-2007)
3nraand mo than 16,000
s mulr of rh urrrrtinftlrm- StanlyL. illr spnt most of lris rr s profssorof hristryat th Uni-
sntialto ollgialand hurnn vrsity of aliforni, San Digo. [n th arly 1950s h prformd a srisof
inluding many rops. rkthrough primnts that signald th ginning of th systmatir-
r l yh g a nt r l o l l t i n s t s sarh of pioti hmistry nd th mr]n of lif. From thn on, to-
ragd y his nrotlrr, an am- gtlrwith many ollagus arrd studnts, Nlillr invstigatdth possil.ll
luthority ln taxonorny, h stps and Irriallmp<lr.rrrds involvd ir-rth mrgnof lif. As .1a_
t 6 . H i s o n g i t l usn t h u s i i l s m krrlwldgmntof his minnt ontriution t th stalishmntof th origin-
n t s a n d o l l a g u sa n d h a s p u t of-lif fild, in 197 h was th first sintistworking in this ara to ltd
tr for rsahand sin- to t National Aadmy of Sins.In rognition of his ahivmntsand
tnt ssayson tlr tonom of his rol in th rtitlnof th lntrnationISoit.forth Study of th Origins
Lthan his wlrk on bs,arrd h oi Lit (ISsoL)' th soit1,stalishdirftr illr's dath th StarrlyL.
. . s( ) v rn t r r n r i at l x o n o m y i n i[lr Award to prsntd tl young outstanding origin.of-lif sintists.
lkl, whos viws wr som- |n \952'795, working as a graduat studnt in th laoratory of hmist
u s r , f o h j t i v q ' u n t i r a t ir and Nol laltratHrold Urv :rtth Univsit,vof higo, illr attmptd
as n ntidot tO som of th to simulat tlr onditions onlutivt<llif on th prirnordii artlr.Irr ths
s of th tim. But h lso - primntsh rvs tsting for th first tim th oparin.Haldan hypothsis
l allowd to sparat th sys- on th mrgnof lif formtrlatdin th 1930s (sth alphatial ntry
rrlvy knowing thir hvior, ..Alksandr l.
oprirr'' in this volum). illr uilt a glass apparatus tht
74 illr

ontaind in sparatflasks onntd y tus a mitur of gasesrimiking ' i llr.

C l n s q u r t t l yM
th arly hydrogn-rih (rduing) atr-nosphrard watr rpsntingth t o t h s r hf o r t l l o
oan. As an nrgy sour h mployd ltri Jishargsthat stood for thsiz and that oulJ
lightning. Aftr running th primnt for wk, Millr analyzd th aqu- Svral suh andidat
ous solution-1[g ..9g6nn''-fg1 th dissolvd produts of th intraringat- T t i s i m p o r t a n tr o n o l
mosphr'sonstitunts.H found aundant synthsisof rgani lnrpotlnds, o f l i f w s r i g h t l yl i n k .
inluding amino aids, th uilding loks of potins. Signifiantly' th r- plants. Aftr th foun
sults wr not statistiallyrandom: Only a small numbr of iologiall rl- tion (NASA) in 1958,
vant organi ompounds wr produd in high yilds in th pimntout t h s t l l l i s h m not f r I
of the vast nlrmr of potntially rlv:rntorgani sustns'Glyin and Svralof illr's lat
alatrin,th rrrostommon amino aids ir-rprotins, wr th rrrajorproduts. tntil for pritlti h
In work prformd y illr and otlrr rsarhrsin th 1970s, it ws found p r v a l n to n o n o f J L
that th sam amino aids and in th sam rlativ quantitis wr otaind moons.
whn th ontnt of mtoritsthat r:rhdarth wtrs analyzd' Sin mt- ln ordr to disd
orits ar onsidrd lis of th formatiorr of th so[ar plants,thir organi rly on th mpirial d
ontnt ra indiat that hmi| prossssirnilar tl th illr rations i o t i s n a r i o se t l d . l
wr ommon in th solirr systm. fad y rsrhrs.,
Similar primnts y othr hmists dmonstrtd th synthsis undr m o t i n g i g n o n
prioti onditions of svralonstituntsf nuli zrids'for irmpl,th t h p o h | r no f r h m t
nitrogn s dnin. or rntly th istnof a prinrordial hydrogn-
BIBLIOGRAPHY
rih atmosphr that nald organi synthsis lras n qustiond' and
many rsarhrs postulat instad that organis wr importd to arth F r . I . 2 0 0 8 . i l l r ,S t
from spa y omts, mtorits,and dust partils. illr nvrtlrlss- Bitlgrph.vo|.,5,l,
i l l r 'S . I . l q . 5 3A. P r r
livd that onditions for organi synthsis on arth might hv istd.
onditios.Si|
Using more potnt high_nrgysollre' ill suddin dmonstrating,
ginning in rh 1980s, tl-rsynthsisof amino aids nd nuli aid assin
a gas mitur that was lss rduing thn his 1953 ..irtmosphr.''
}vlimiry
illr Was a promotr of th gnti onption <lfth origin lf lif' a-
ording tl l,vhilrlivig sstnlsw first and ftrmostsl{-plitingmo- N4imiry ours rvhn
lular ntitis. Throughout his r th most fr'rndamntalqustion that and, tlrrough that s
proupid him was th natur of th first gnti matrial that ould hav iry is strikingphn
rpliatd, mutatd' and volved to form a primitiv living systrn.illr r o | i n v o l u t i o n a r yi
shard th orrvition f r-t-rnysintistsin th fild tht watrslrdin th t i o n , n d s v r | h l l
mrgnof lif was r-rRNA world, hmial world in whih RNA mol- s y s t m sr m i n t o l . s
uls funtiond oth as gntimatrial and as atlysts.Th id that RNA imir is mpn
moluls ould hav fulfilld this dul rol vry rly in th volution of lif simplst form, mimir'
was suggst{ indpndntlyin th 1960s y Lsli orgl, F.ranisrik, and s m l si V r y d i f f
.W.os.
rl Tlr mo radil lim thr RNA ws th sol plyr irr th vry aggssivnlinliv.wl.
mrgnlf lif anrpopular in th 1980s aft th surprisirlgdisovry that rsmlsth fod
in prsnt.daylls of riozyms. Ths r RNA moluls that in addition o f a n a I l i g e t o s n p p i
to thir gnti rol also t as nzyms, thus arryirrgout th funtion that a t i n gf i s h i n t o t h r u r r
until thn ws known to rmplishdonly y potins. wll_knowrrxampls
On th asis of numrous xprinrnts,htlwvr,Millr, s wll as his ol- fit to its ownr and tl
lgu Lsli Orgl and othr rsarhrs,am onvind that lif ould t h o g n i s m i n gi m
not hav gun with th RNA world. Thy rlizd that th prioti synth- Th mor omplia
sis of an RNA squn that ould rpliat itslf was highly improal. r n ,a s i
I sa s i r r l i l o

s a mitur of gassmimiking
r and Watr rprsnting th
:tri dishargs that stood for
rk,illr analyzd th aqu.
p r o d u t so f t h i n r r a t i n g t -
lnthsisof organi ompounds,
protins.Signifintly,th r-
all numr of iologilly rl-
4h yilds in th xprimnt out
gani sustans.Glyin and
ins,wr th major produts.
:hrsin th 1970s, it was found
lativquntitiswr otaind
arth Was anlyzd.Sin mt-
t h s o | rp | a n t st,h i r o r g n i
similar to th Millr ratlons
ronstratdth synthsis und
nuli aids, for ampl, th
nof a primordial hydrogn-
sis has n qustiond' and
anis wr importd to arth
artils.illr nvrthlss-
on arth might hav istd.
r sudd in dmonstrating,
' aids nd nuli aid ass in
1953..atmosphr.''
ption lf th origin of lif' a-
mo-
l f o r m l l s ts l f - r p l i t i n g
ost fundamntalqustion that
ntimatrial tht ould hav
primitiv living systm. Millr
r fild that a watrshd in th
lworld in whih RNA mol-
rs atalysts.Th ida that RNA
ry arly in th volution of lif
-sliorgl, Franis rik, and
r ws th sol playr in th vry
l s a f t rt h s u p r i s i n gd i s o v r y
i'NA molulsthat in addition
; arrying out th funtion that
y by protins.
vvr'Millr, as wll as his ol-
am onvind that lif ould
alizdthat th prbioti synth-
: itslf was highly improal.
imicr 75
onsquntly, illr's rsarh from th 1990s until his dth was dvotd
to th srh for th onstituntsof a pr-RNA world that wr asirto syn-
thsizand that ould hav volvd into th onstituntsof th RNA world.
Svralsuh andidatswr suggstdy Millr nd his group.
It is important to not that from its inption illr's rsarh on th origin
of lif was tightly linkd to th widr ontxt of th sarh for lif on othr
plants.Aftr th founding of th National Aronautis and Spa Administra-
tion (NASA) in 1958, th young Millr Was among th main ontriutors to
th stalishmnt of th fild of oiology, latr to b alld astoiology.
Svralof Millr's latr primnts lnt rdnto th hypothsis that th po-
tntial fr prioti hmistry xists byond arth, prhaps undr orrditions
prvalnt on on of Jupitr's moons' uropa' and possily on othr Jovian
moons.
In ordr to disredit th study of th mrgn of lif, rtionists oftn
rly on th mpirial datsamong origin-of-lif sintistsovr spifipr-
bioti snarios and on th many thortil and mpirial diffiultis still
fad y rsarhrs.Millr, who strongly dnound rationists for pro-
moting ignoran out sin' nvr doubtd that a hmil solution to
th prlrn f th mrgnof lif will vntually found.
|LIoGRAPHY
Fry' I. 2008.illr' StnlyLloyd. In N. Kortg'd.,Nez Ditiorof Sintifi
Bbgrph,vl. 5, 154-158.Dtroit:harlsSrinr's Sons.
illr, s. L. 1953.A produtionof amino aidsundrpossilprimitivrth
-l.F'
onditions.Sin1'17:528-529.
imiry
Mimiry ours whn th faturs of ln spisrsml ths rf anothr
and, through that rsmlan,onfr som advantag on th mimi. Mim-
iry is a striking phnomnon in natur; its study has plyd an impotant
rol in volutionary iology from th rlist argumnts aout ntural si-
tion, and svralhallngingpuzzls about th volution of ompl rnimiry
systmsrmain to solvd.
i m i r y i s r a m p a n t i n n a t u r ( s W i k l r 1 9 6 8 ; R a n d a l l 2 0 0 5 ) . I n i t s
simplst form' mimiry ours whn a partiular fatur of on organism r-
smls a vry diffrnt organism. Th most familiar xampls ar ass of
aggrssivmimiry' whr a prdator lus its pry with a prt of its wn body
that rsmlsth food of th pry. Th flshy rd proturan on th tongu
of an alligator snapping turtl mimis a lodworm and lurs loodworm.
ating fish into th turtl's mouth. Th lurs of anglrfish and atfish ar othr
wll-known ampls. In ths ass' th mimti fatur onfrs a lar n.
fit to its ownr nd th volutin of th fatur rats no disadvants for
th organism ing imitatd.
Th mor ompliatd flrm of mimiry ours whn on organism rsm-
ls a similar on' as in on uttrfly rsmlinganothr' ut 'ith th twist
76 imir

tht th rsmlanoffrs a nfit to th mimi Llt potntial disadvan- fnding food itm. It
tag to th nrodl (th organism ing imitatd)' This happns whn modl of a paltal spi
and mimi hav vry diffrnt l.laratristisor ologial rols and th su- nss through a rdu
ss of th mimi thratns th modl. For ampl' on oral rfs in th Pa- lowr risk of prdati
ifi, small fish alld ..lanr wrasss'' find food y piking rrrstaan mimi and modl zrr
toparasits from th sals of largr fish, whih visit spifi spots on th Dspit th simpli
rf whr th wrasssour' Thr is anothr spisof small fish, a lnny, rl irpotnt qLlst
that mimis th shap and oloation of th lanr Wrass ut, athr than sian mimis must r
pik parasits from th larg fish, taks small its from thir tail fins. Th mor mimis tha m
blnny's mimiry of th wrss allows it to los in on its targt whn it might rathr than th rvr
<lthrwis atn itslf y th largr rf fish. But on might pt th an- darrs of modl anl
noyn of a rf 6sh at having a pi rmovd frorn its tail will pr<rvoklss as th modl fo s
tolran of th modl' th lanr wrasS' and impril th wrass'sailiry to pipvin swallowtail
gain th asy mal. To undrstarrdth prsistnof this mimiry systm' W Sond, if Batsia
must undrstand th riproal ffts of modl nd mimi on ah othr's gns that ontrol p
fitnss and th onditions undr whih thos ffts ar aland. strutions for th pa
Th most hllnging ompl mimiry systmsar th hundrds of ass th alphabtialntr
in whih dngous or toxi ninrals ar imitatd y harmlss or platal that thos gnsr i:
ons. In ah s th harmlss or palatal spisgains its advantag through the fat that tlrr is
th aility of othr animals to larn th assoiation twna visul or olfa- hritan of mimti
tory u with th dangous nimal and thn assoiat tht u with th gns of larg fft
hrmlss mimi arrd lav it alon. gns wr oviousl
Th most wll_studidass ar thos involving palatal uttrflisthat trastd with th Dar
rsmlunpalat:.rl, toi oIrs.Hnry Bats (1862) rought this phnom- was th umulativ il
non to th ttnti>n of iologists whn h dsridhow rtain Amazonian disput was larifid
buttrflis in th family Pirida or striking rsmlan to unrlatd ut- onrtd inhritan
trflis in th family lioniida. Bats had notid that th hlioniid ut- pattrns and th ff
trflisflw languidly yt wr rarly attakd y irds, whil th pirids flw or-rtrolling gns (s
m<lrfurtivly and lusivly,as if viding xposur to irds. Th hliorriids nw rsarh lrersrv
ar' in fat, toi nd unplatal, and th pirids ar nontoi and palat- t a l . 2 0 0 8 ) ,t h v o l
al. Bats dsrid this systm as a dramati dmonstration of tlr powr linkd ompl tlrat i
of natural seltion' pointing out th advantaggaind y an individual of a Th dsription of
p:rlatalspis whos featurs rsmld thos of a noxious spis that whn diffrnt popr.r
would avoidd y prdators. looking modls in di
Inspird y Bats's osrvations and suggstions' volutior-raryiologists ig pattrn within th
from harls Darwin onward plungd into studying th natur and volution is starting to rvltl
of this typ of minriry. With gratr srutiny am th appriation that mim- Third, th study o
iry is a ompliatd phnomnon; as a rsult, th study of mimiry am to unpalataility and t
oupy a ntral position in volutionary biology as a nrodl systm for undr- that a prdatr wor.
standing omplitd forms of natural sltionthat ould nrold ompl pat- must di so that th
tns from simpl gntiomponl1ts. sam apparan en
omplx mimti systms an lassifid into thr typs. In Batsian sprad via th lass
mimiry a harmlss or palatal spis rsmlsa dangrous or unpalat- gstd a kin-sltion
abl, toi on. It is not diffiult to undrstand how th prdator larns to mzrny toxi uttrfli
avoid a toxi uttrfly;a prdator tht ats on omsill nd vomits, aftr and otlrrs itd thr
whih tast avsion lads th prdator to avoid Drv that rsrnlth of. ity and warning olo

Limiut a potntial disadvan-
d).This happns whn modl
r ologial rols and th su_
rmpl, on ral rfs in th Pa-
d food y piking rustaan
'hih visit spifi spots on th
.spisof small fish, a lnny,
:lanrwrass ut' rathr than
ll its fm thir tail fins. Th
s in on its targt whn it might
l. But on might pt th an-
:d from its tail will provok lss
d impril th wass'saility to
:nof this mimiry systm' w
'dl and nrimi on ah othe's
fftsar aland.
;tmsar th hundrds of ass
tatd y harmlss r paltal
:isgains its advantag through
ation btwna visul or olfa-
n assoiatthat u with th
llving palatal uttrflisthat
s (1862)roughthis phnom-
; r i dh o w r r a i nA m z o n i n
g sr-rrlan to unrlatd ut_
notid that th hlioniid ut-
[ y ids,whil th pirids flw
posurto irds. h hiiorriids
pirids a nontoi and palat-
ti dmonstrationof th powr
rggaind by an individul of a
:hosof a noxious spis that
;stions'vlutionary iologists
udyirrgth natur and volution
Jamth ppriationthat mim-
t, th study of mimiry am to
)gy as modl systm for undr-
ln that ould mold omplx pat-
'd into thr typs. In Batsian
lmlsa dangrous or unpalat_
rnd how th prdator larns to
n omsill and vomits, aftr
void pry that rsml th of-
icr 7.7
fnding food itm. It is also not diffiult t s Bats'Sargumnt that a mutnt
of a pltal spistht rsmlsiln unplatzrlon will hav highr 6t-
nss through a rdud risk of prdatin. If a ttr rsmlan<rnfrsa
lowr risk of prdation, natural sltion will rfin tht rsmlanuntil
mimi and modl ar indistinguishal'
Dspit th sirnpliity of th asi argumnts,Batsian rrrimiryri1isssv-
eral important qustions for whih w hv only partial answrs.First, Bat-
sian mi-lismust rmain rr thn thir modls lst prdators nOuntr
mor nrimis thn modls and assliatth shd pattr wih palatailit1'
rathr than th rvrs.Th pris riationship ptd among tlr aun-
dansof modl and mimi is unlar, spiallywhn a singl spissrvs
as th modl fo svral mimti spis in th sm lotion (.g., th
pipvin swallowtail of th southasrnUnitd Statsand its mimis)'
Sond, if Batsian mimis must rsml thir modls losly, thn th
gnsthat ontrol pattrn mllst inhritd togthr so that th gnti in-
strutions for th pattrn ar not silttrddr-rringsuirl rprodr-rti<ln (s
th alphabtialntry ..volutionof s'' in this volum).Rsarhhas shown
that thos gnsar inhritd as a lok, a pross failitatdin buttrflisy
th ft that tl-rris no rossing-ovr in fmzrls.Th initial studis of in-
hritan of mimti pattrns suggstdthat thy Wr ontrolld y singl
gns of larg fft, a finding usd y som sintists to argu that suh
gns wr oviousl,vth important raw matril for volutin. Tis on.
trastd with th Darwinian viw that th important rw matrial for volution
was th umulativ impat of many gnswith small individual ffts.This
dispur was lrifid only wlrn dtild studis of inhritan rvald th
onrtd inhritan of th svral gns that ontrolld th asi mimi
pattrns and th ffts of othr gns that modifid th xprssion of ths
ontrolling gns (s Nijhout 200 for a tial disussion). Although
nw rsarh has rvidth loation of ths gns in sotn spis (Clrk
t al. 2008)' th volutionary pathway that ror"rght thos gnsinto losly
linkd ompl tlrt is inhritd as a lok rmins to dsrid.
Th dsription of suh Irvolutionary pathway is vn nror ompliatd
whn diffrntpopulations f th sam mimti spisinrittvry diffrnt-
looking modls in diffrnt gographi loations. This is an spiallystrik-
ing pattrn within th hlioniid uttrflis,and nw rsarhinto this Pattrn
is starting to rvi thsp:rthways(Krnforst nd Gilrt 2008).
Third, th study of Batsian mimiry ld quikly to th qustion of hw
unpalataility and toiity ould volv, along with a spifi visual signal
tht pdator would rmrr.]f mr'ttnttlrt is unpalatal nd toi
must di so that th prdator n larn to avoid othr individuals with th
sam apparan and initial tast, it is unlr how th mutant gn an
sprad vi th lssi Darrvinian sltion amon]indivi.luals.Fishr sug-
gstd kin-sltinargllmnt asd on th supposdlygrgariousrraturof
many toi uttrflis,ut rnt rsarh on phylogntis (Bltran et a|.2007
and othrs itd thin) has indiatd that grgariousnssvolvd aftr toi-
ity and wrning ()loration in rany of ths spis, nd so th kin-sltion
78 imir

argumnt annot orrt. Othr hypothss hav n suggstd, ut th gs th qustiotl Ot

prolm rmains unsolvd. i n s p i r d p rq u s
Th sond typ of ompl mimiry is alld iillrian mimiry' whih of sophistiatd rs
dsriesth rsmlanof two or mor unpalatal or dan;rousspis work in gntis, d
to ah othr. This pattrn was lso dsrid y Bats, who osrvd that and omputational
many tropial uttrfly spis formd mimiry rings in whih a group of stoy, undrstandin
spis hiitd a ommon mimti pattrn. Furthr, Bats osrvd that to unfold.
suh pattrns hangd in wholsal fashion from on gographi ara to an-
othr. Svral spis within th gnus l]eliois in th Amrian trpis BIBLIOGRAP
offr on of th st.doumntd ass of a mimiry ring. Miillrian mimiry BtsH , . W . 1 8 6 2 .
and mimiry rings pos a numr of prplxing qustions.Th advantagf rstirlsof t|
a mutual resmlanof unpalatal spisis lar at first; if a rtain num- Bltran,M.' . D. Jigg
pollnfding,pu
r of individuals mrrst di to tah prdators to avoid a pattrn, thn if that
Iiliusttr1
numbr is dividd twn two spis, th risk of prdation is lowr for in- BiologilJour
dividuals of ah spis,and a mutual smlanwill volv. Of ours' lark, R', S. . Brorv
th prolm of how toxiity volvs in th first pla still xists aus this 2008. olour pat
argumnt bgins on two spis ar ah ti. And although many pla- Th transriptio
nations hav n suggstd, it rmains unlar why mimiry rings hang Prodigstlf th
ronforst,M. R., rrd
thir pattrns aruptly vry fw hundrd mils.
divrsityin HIi
Th third typ of ompl mimiry is th most puzz\ing. In ths ass all S r i sB 2 7 5 : 4 9 3
th spisinvolvd ar ti or dangrousut thy vary sustantiallyin that M l l t 'J . 2 0 0 1 . a u s
noiousnss.This maks th rings diffiult to dsri as ithr purely Bat- mimiry.uolutk
sian ompls or purly Miillrian rings. Th most sptaularof thsass N4allt,J., nd M. Jr
Polymorphismss
might th mimiry group that inluds th dadly oral snaks of tropial
nd Sstttis
Amria and a srisof mildly vnomous spisthat shar th rightly andd N i j h o u t ,H . F . 2 0 0 .I
olor pattrn. Th diffiulty in plaining this pattrn is that a it from th x. ig ffts,ar"rd l
trmly dadly oral snak kills most would-b prdators, and dad prdators R a n d a l l , . J. . 2 0 0 5 .A
annot larn to avoid what killd thm. 299-328.
.!7iklr'
Two planations hav n offrd for this typ of mimiry ring. In th !7. 1968.il
first xplanation, alld rtnsian mimiry' th mildly noious spisis in
fat th modl from whih prdators larn and th dadly spis is th
mimi, th rsmlan ing favord y th fat that attak ts on th IyIdrnrptils
dadly animal drasas pdators larn from th lss dadly nountrs. Rptils onstitut
Th sond plantion, alld quasi-Batsianmimiy, rvrssmodl and groups of ttrapod (
mimi and postulats that th mildly noxious spis mimis th highly nox- ing rptils rang in
ious on. This hypothsis was dvlopd for uttrfly rings in whih spis 1 gram to saltwatr
varid in aundan, as wll as toiity' th mildly toi spis ing th vr 1,100 kilogra
rarr. Th rlativ aundans rsml th argumnts about modl and liv-aring and p
mimi aundans that mrg from purly Batsian mimiry, hn th nam (warm-loodd),ma
qusi-Btesi. Te mhanisms positd for quasi-Batsian mimiry involv most of th ologi
vry spifi argumnts aout how animals larn, inluding argumntsaout ttrapods. Som cf
whthr larning is prmannt and how long larning lasts without rinfor- xmpls ound o
mnt (s allt and Joron 1999 for a dpr disussion).Thse opposing aring and th ot}
xplanations rmin to b rsolvd (Mallt 2001). ontrovrsis in rpt
h study of mimiry pitomizs volutionary iology as a disiplin. goup' and in parti
Simpl mimiry is radily osrvd y vn a asual natural historian and ..Birds''in
ial ntry

:s hav n sr.rggstd,
ut th
lld Mijllrian mimiry, whih
p l a t a l o r J a n g r o u s s p i s
d y Bats,wlro osrvd that
ry rings in whih a group of
L. Furthr, Bats osrvd that
.om on gographi ara to an-
.,.liusitt th Amrin tropis
i m i r yr i n g . M i i l l r i n m i m i r y
ng qustions.Th advantag oI
s l a ra t f i r s t ;i f a r t i n n u m -
; to avoid a pattrn' thn if that
isk of prdation is lowr for in-
nlanwill volv. Of ours,
:st pla still ists aus this
,i. Anl although rnany xpla-
lar why mimir1' rings lrng
ls.
rrostpuzzlirrg.In ths ass all
lt thy vry sustantiallyin that
l {sri as ithr purly Bat-
most sptaularof ths ass
: ddlyoral snaks of tropial
s that shrth rightly andd
pattrnis that it from th x_
l prdators,ad dad prdators
his typ of minriry ring. In th
, t h m i l d l yn o i o u ss p i si s i n
r ar-rdtlr dadly spis is th
r fat tht attak rats on th
om th lss dadly nountrs.
rn mimiry, rvrssmodl and
s spismimis th highly nox-
uttrflyrings in whih spis
mildly toxi spis bing th
L nr$untS aout modl and
atsianrnimiry, hn th nam
quasi-Btsianmimiry involv
:arn' inluding argumnts aout
larning lasrs without rirrfor_
l r d i s r r s s i o n )T.h s o p p o s i n g
001).
ionary iology as a disiplin.
a a s u ln a t u r a l h i s t o r i a n a n d
ldr Rptils 7.9
gs th qustion of its origin nd raintnn.rlmpl mimiry systms
inspi d;lr qustions' whih lrav provokd narly a ntury alrd a half
of sopistiiltd sah, from Bats's lriginl osvtitlns to modrn
work in gntis, dvlopmnt, ology, nuroiology, and mathmatial
nd omptrttilnl modling. Wlril tlr disovy of mimiry is n old
story, undstandingits volution is vry muh a nrodrn on that ontlnus
to unfold.
|BLIoGRAPHY
Bats,H. !(/.1862.ontriutionstrl an instfaurlaof th Amazon Vally.
7'rstitls lf th l-i,tStlitl|-Lolll 2: 495_566'
| J l t r a n, . , . D . J i g g i n s^
, . v . Z . B r o w r
, . B r m i n g h ma,n d J . l l t .2 0 7 . D o
pollnfding,pupl-rrrting anllrr,Iggariousnss hr', sirrglliginin
Hlioiusuttrflis? Infrns from mrrltilous DNA squn data.
Blrlllgiitl jourttllftb ]'inSlliat1, 92:121-239,
lark, R.' S. . Brown,S. . ollins,. D. Jiggins,D. G. Hkl,and A. l).Voglr.
2008.olur ttrnspifiatiorr in th Mokr swllowtilP1lilidlrdnus:
Tlr trarrsriptin fatorinvtdis a ndidatfor thmimirylousI{.
Prdigs o|.thRoltlSlit1t of l'odo, SerisB 275: 1181-1188.
ronfrst'N{.R., :lndt,. . Gilrt.2008.Th opulatiln gt-ttis of mimti
divrsityln Heliiusuttrflis.Prodigsof tb Rol Soitof Lodl,
SrisB 275:49_5(|0.
llt,J,2001. ussnd onsquns of a lak lfovlutionin iillrin
,l l , l 9 1 ,I j : , 7 - - . 0 .
l t l i r n t r L' .t l l l t t ! i r ' t t . t r
llt,.}.,nd . Joron. 1999' volution of divrsity in warrring olor and mimiry:
Polymorpisms, shifting alan,rld sititlt-t. ,Aul Rt'ilt',
lf olog1,
td Sstemtis 30: 2,01_2.]3.
Nijhout, H. F. 2003. Polvmorplri minrirv irl Ppilio drt]us:Mosi dorinan'
ig ffts,and origins. ullutio d Duklpmnt 5: 579-592.
Itandall,J. . 2005. A rviw lf mimiry in rrin frses.Zorlll.ilStudics 44:
299_28.
Wiklr,W. 1968.imir.Londot-t:
Widnfldand Niolson. -rT
odrn rptils
Rptils onstitut tln of tlr most mlrphologially arrd ologially divrs
grups of ttapod (fou-lggd)vrtrts.Th roughly l6'000 spisof liv-
ing rptils rang in ldy siz from tiny lizards and snaks that wigh lss than
1 granr to sltwatrroodils that rang up to narly 7 mtrsin lngth and
ovr l,100 kilograms in wight. Th living rptils inlud spisthat r
liv-aring and gg-lying, tothrmi (old-loodd) an.l ndothrrrri
(warm-loldd),marin nd trrstrial,lrrivorous and prdaous-in short,
most tlf th ologial, physilogial' arrd morphllogiI r.riation found in
ttrapds. Sorn of this variation an trmly lail volutionarily-
anrls arrnd of piris lf l<lslyrlatd lizard spis wlr on is liv-
aring and th <lthrgg-lving, ftrr ampI. On of th nrost important
lntrovrsisin rptili-rr.r vtllutiorriryilllgyis :rtlywhat tl inlud irr this
group' and in partiulr, th inlusion of irds rvithin rptils (sth irlpht-
ial rrtry..Birds'' iIr this vllurn).A phylognti dfiitiorrof rptils that most
740 oder R1ltiles

xprts would agr on is ..th most rnt omlon anestor of a turtl, ir Th lpidosallrs ar
snak, a |izard, and roodil, and ll spisdrivd from that anstor.''If Thy inlud two grol
that dfinitiotris usd' the living rpdls inlud lizds, snaks, amplrisanians lizardlik spis rstr
(a group of limlss,lizdlik arrimals),rhynhoplralians(ttutars)'ro- th -7,000 spis of
odilians, bids, and turtls. It also inluds a grat varity of tint linags, tivly alld th squarn
inluding th marin ihthyosaurs and plsiosaurs, th flying ptrosaurs, and tropial rgions, altho
th vr-popr-rlar ornithislrian (ird-hippd)and s.rtlrishian (lizrd-hippd) masss pt polar ;
dinosaurs (stlr alphatial ntry ..)it-tosaus: Th n-rodlsystrnflr volu- uiquitous and prsist
tion'' in this volum). patd volutionry l
Phylogntially th living rptils ar oftn split into thr primary linags. plt limlssnsshas
Th lpidosaursinlud th lizards, snaks,amphisbnians,and ttlatarasand group. Th most fam
.With
r th numrilly dltninirntgroup of nonavin ptils.Arhosaurs inlud narly 3,000 livi
roodiliarrsand irds, s wll as a vast divrsity of xtint spis,inludig of limblss squamats
th traditional dinosaurs. Finally, th tstudins (sorntimsalso known s th th skinks. othr imp
hlonians)inlud th turtls nd tlrtoiss, small group of mlrphologially of omplx vn()l sys
uniqu vrtbrts.Amlng rptile iologists on of tlr ky rnt rsarh f_ omplity and pris
forts has fousd on lrtlw ths majo groups ar rlatd to ah othr, lrd ifid skull and tth all
wht thsrlationshipstll us out th history of divrsifiationof ttrapd dirtly inttl pry with
vrtrats. maxillary on; whr
Turtls ar oftn onsidrdth m<lstprimitiv grop of rptilson th . mourh.
sis of thir shard olr]itionof th lak of an opning (or aps) in th hk Finally, th living a
rgion of th skull (th anapsid lndition).ammals and thir rlativshav hr) and th 22 livin
singl opning (and ar lassifids synapsids)'and all othr rptils hav a
pir of opnings in th hk rgion (diapsids). Orr th sis of thir .
trrnl distintiv morphology and tlr notion tht th anapsid ondition is
th simpIstirnd thrforth most primitiv' turtls hav oftn n onsid-
rd th first linag to split volutionarily from th othr rptils.owvr,
rnt molular nd moplrologial rsrhindiatsthat turtls,whil r-
tainly iz arrd spializd' may wll form of modifid dipsid that
has sond:rrilylost th opnings hratristiof th group. atly whr
turtls fall within th diapsids rmains ontrovrsil;som analyss favor
losr affinity to th lpid<rsaurs, whil othrs favor .rn arhosaurin affinity.
Although th rlationship of turtls t() ()thrttrpods is ntrovrsial,r-
ognizing a tllrtl is not. Both th slrll (a fusd, modifid ri ag) and th
plamnt of the shouldr girdl insid th ri ag (as pposd to outsid
th ris in all othr ttrapods) ar uniqu faturs in th history of vrt_
rt volution. For th pst 220 million vars turtls lrv maintaind ths
uniqu fturs and hav n a onsistnt fatur of lif on arth. Th liv-
ing turtls now ontin aout 325 spisthat ar widly distriutd aross
th tmprat and tropial rgions of th world. Th living turtls ar oftn
dividd into twl major gror.rps. The plurodirs,or sid-nkdttlrtls(aout Th pin snak,itop
irritsth pin fors
75 spis)'hav a horizontlly rtrta|nk joint, whil tli ryptodirs
Nlv Jrsy.Pir-rsnak
(aout 250 spis)hav vrtially ori'ntdnk joints that nl thm to
witlr gophr tortois
rtrat thir had straight bak into th shll. Plurodirsdominat th south- rptils'with narly3'(
rn ontinnts (Afria, Sotlr Amria, nd Australiil), whil ryptodirs r ftursinluding'irr
found throughout ursiir and North Amria. injtingvnom intlpt
 lder Reptils 741

ommol] anesto of a turtl, a Th lpidosaurs r th most divrs and nunrous of th living rptils'
' drivd frorn tht anstor.'' If Thy inltrd two !.roups: th morplrologially primitiv tuatras (a pair of
lizards,sltaks'mphisbanians lizardlik spis rstritdto svral lffshoreislands in Nw Zaland) and
ophalians(tlrtuataras),ro- th -7,000 spis of lizards, snaks, and thi rlativs (whih ar olle-
lratvarity of tint linags, tivly alld th squamatrptils).Squamatsr most divrs in th world's
aus' th flying ptrosaurs, and tropial rgions, although thy our on all ontinntal and most islnd land
nd saurishin(lizard-hippd) masss xpt polr and trm high-lvation rgions. On of th most
rs: Th modl systl for volu- uiquitous and psistntvolutionary thms in squamat rptils is th r-
patd volutionary loss of oth th digits and lims; y on stimat om-
splitinto thr;lrim,vlirrags. plt limlssnsslras volvd minimum of 62 indepndnt tims in th
lphisirnizrns,irnd tutras and group. Th most fmous xampl is th srraks' or srpnts (s figur)'
ian rptils.Arhosurs inlud !7ith narly ,000 living spis'thy ar y far th most sussfullinag
sity of xtint spis'inluding of limlss squamats)ut many othr ampls alound,partiularly anong
:s (somtimsirlso known as th th skinks. Othr important innovations in rh squamats ar th volr.ltion
small group of morphologially of ompl vnom systms,whih rh thir gratstlvl of morphologial
n of th ky nt rsarh f- omplxity and prision in th viprs and rattlsnaks.r th highly mod-
; ar rltd to 'ah othr, and ifid skull and ttlr :rllow injtion of prisl n-rodulatdamounts of vnt>rr-t
tlf ttrapod
lry of divrsifiatior-r dirtly into pr witlr a fang lik a hypodrmi ndlthat is attahdto tlr
maillary on; lvlrn not in us, t fngs an foldd ak into th
tiv gror.rpf rptilson th - mourh.
t opning (or aps) in th hk Finally, th living rhosaurs onsist of th ids (not onsidrd furthr
ammalsnd thir rlativshav hr) and t 22 living spis of alligators' roodils, and thir rltivs.
ds),and all otlrrptils hav a
;ids).on th asis of thir x-
n that th anpsid ondition is
, turtlshv oftn bn onsid-
om th othr rptils' Howvr,
indiatstht trrrtls,whil r-
r forrrrof m<ldifiddipsid tlrt
;ti of th grollp. xatly whr
ovrsial;slnranlyss favor a
s favor an hosaurianaffinity.
)r ttrapodsis ontrovrsial,r-
.sd,modifid ri ag) and th
ri ag (as lpposd to outsid
atursin th histtlry of vrt-
trs turtlshav rnintind ths
fattlrof lif on arth. Th liv-
rat ar 'idlvdistriutd aross
rld. Th livirrgturtls ar oftn
Th pin snak,Pitulphismllurzs, a rtrof th olurid famil,
rs,r sid-nkdturtls (aout
inhaitsth pin flrstsof th southrnUnitd Statsnd th Pin Barrnstlf
:rkjoint' whil th ryptodirs
Nw Jrsy.Pin snaksurrow in sandy soil nd ar o{tnfound in assoiltin
nk joints that nal thm to with gophrtortoisolonis.Snaksar th mst sussfullinagof limlss
Plurodirsdorlinat th south- rptils'with narly 3,000 spis,and hiit a markalrangof daptiv
Australia),whil rvptodirs ar fatursinluding',in rllany spis(ut nt pin srrks),
a omplx systmfo
. injtingvnom ito prv irnd ttkrs.
742 Mods of Spiation

Uniformly prsutdfor thir skins rrd rnat,y |970 aI|roodilian spis portn of hromosorr
on arth wr onsidrd thratnd or ndangrd y th Intrnational -99% of losly rlatd;
Uniln for th onsrvation of Natur (www.iulr'org), an intrntional tions, fusions' or othr r
organiztion that r-tronitorsiodivrsity dlins.Howvr, |995 onsr- a r r a n 8 m n t sa r i s i s h i g
vation fforts around th world had rdudthis y aout half, and as of this nwly arisn rarrangm
writirrg many spiSappar to b rovringto halthy population sizs.R- ish frtility whn thy :rr
flting thir rltivly ls phylgnti rltionship to irds, ro<rdilians ularitis in miosis. Thl
ar oth th most voal and provid th gratstlvls of parntal ar of any arrangmnts,thir F l h'
.!7hit's
group of nonavian rpti1s. Alas' viw of
On of th lssi qustions in rptilian volutionry biology is ..Why did in th nt l5 yasT . h.
th dinosaurs go tint?'' Although it is tru that mny linagsof dinosaurs onditions Wr nss.1
did go xtint roughly 6.5 millin yars ago at th nd of th rtaotlsp- ing spition)whil ls<
iod (and that tintion was tiggrd y a mtorit ollision with th ariss (th individul wh
:rrth),it is important to rmmr that phylogntially,irds atually fall erozygot). mpiril w
within th dinosaus and ar therfora kind of dinosaur. Thus in a vry ral somtims not as strit:
sns th answr to this qustion is that th dinosaurs did not go tint. frnt ways in whih ra
That pigon that yotr saw in th park today was a prftlygood dinosarrr;so rombination) rthr th
was th lrikn that you at for dinnr. His mphsis on th im1
planation for how tlryl
|BLIoGRAPHY
vr' thr r ption
's?ashington'
rnst' . H., and R. \(. Barbour. 1989. Trtlsof th World' D: a dirt fft of a transl(
SmithsnianInstitr-rtion Prss. .Whit
takld svr.rl
Grn.H. w. 1997. Sks:Th uolutirlof str i Ntur.Brkly:
gading spisdiffrn
Ur-rivrsitytlf al iforrriPrss.
L,. S. Y., T. W. Rdr,J. B. Slowiski,nd R. Lawsrrn.2004.Rsolvingrptil rnst Mar's influntiirlvi
rlationships: olularand morphologialmarkrs.In J. raraftand . J. ation, during whih a lar1
Donoghu,eds.,ssmbligth Tr of Life' 45|467. Nw York: Oxford ltiv prssursaftr olt
UnivrsityPrss. of sympatri spiationin
Pinka, . R., and L. J. Vitt. 2003, Lizrds:'V/idolllsto th uolutio of Diursit.
studid today in this ont
Brkly: Univrsit,v of liforniPrss. ..tho
Pough'F. ., R. . Andrws,J. . Cadl,. L. rump' A. . Savitsky,ad K. D. ours), he itiizd
Wells.2004.rpetolog'3rd d.Uppr SaddlRivr,NJ: ParsonPrnti disrimination and othr
all. -t{.8.s. ation. Instad, h suggs
in spiation. Although h
(2004) also not th ovr
Nlods of Specition (ichal J. D. Whit)
natiolr whrl th rltivi
A ommon misonption among studntsof sinis that thoris nrust hyrid prolms i s;.l
.\rhit's
orrt to produ progrss in thi disiplins.For ampl, studntsmisun- txt is rihl
drstand why thy larn th thory of th inhritan of aquird haratrs disussions of mpiria
.!7hit
of Jan-Baptist Lamrk, rmmring him only as ..tht guy who Was Wrong Was an outstndi
about hrdity''' Howvr' that is a shortsightdviw: Lmark prsntda onvind that th phn
omprhnsivthory of volution at a tim whr-rth prossswee poorly spiartion.Althoug so
undrstood. vn if his ontriutitln was impris in dtail, it positivly in- an dout that this t
flund our undrstandingof volution today. studis of spiation.So
With that in mind' it is a plasur to ommnt on ytologist ihal J. D. many ontinu to t
Whit's lassi 1978 book ods of Spitio.This book Was th most om- ..And' at th prs
plis:
prhrrsivtratmnt of spiation until th apparanof oyn and orr's o u t l r h o d so f s p i
2004 tt. h most famous asptof Whit's ook is his mphasison th im- mor justifilto gnr

., 1970 all roodilian spis
rdangrdy th Intrnational
w . i u n ' o r g ) 'a n i n t r n a t i o n a l
ins.Howvr, I995 onsr-
this y bout half, and as of this
to halthypopulation sizs. R-
iationshipto birds, roodilians
'stlvlsof parntal are of any
,olutionary iology is ...Why did
tht many linagsof dinosaurs
at th nd of th rtaous p_
a mtoritollision with th
'logntilly,irds atually fall
l of dirrosaur.Thus in a vry ral
d i n o s u sd i d n o t g o x t i n
asa prftlygood dinosaur; so
D:
f th Wrld..!ashington'
Isterfu Ntr.Brkl:
R. Lawson.2004. Rsolvingrptil
marks. In J. rraftand M. J.
,, 45|467. Nw Yrk: Oford
lotusto thuoltiorlf Diuersit.
runrp'A. H. Savitsky'ar-rdK. D.
JdlRivr,NJ: PlrsonPrnti
-F1.8.S.
' Whit)
lf sinis th:rt thoris must
s .F o r x a m p | s. t u d n t sm i s u n -
rhritanof quird haratrs
..thatguy who was wrong
only s
;htdviw: Lamark prsntd a
,whn th prossswr poorly
rprisin dtil' it positivly in-
ay.
mnt on ytologist i\4ihal J. D.
o.This book was th most om.
apparanof oyn and Orr's
's
ook is his mphasison th im_
ods rfSpiation 74.]
portan of homosoml rarrangmntsin spiation. shit notd tht
-99Y. of losly rlatd nir-nalsdiff by hromosomal invsions.trnsloa.
tions, fusions' or othr rarang-lnts,and that tlr rat at whih ths r-
arrangntsaris is high y volutionary standards (1/500 individuals has a
nwly arisn rarrngmnt)'H also notd that most rarrnglntsdimin-
ish frtility whn thy ar htrozygous zrushtrozygosity indus irg-
ularities in miosis. Thus whn divrging populations om to diffr in
arangn.lnts' thir F1 hyrids ar stril,and spitionhas tlurrd.
Alas' \/hit'sviw of rchil hromoslml spcitioz fll into disfavor
in th nxt 15 yars. Thortial argumnts showd that trmly stringnt
onditiorrsWr nssaryfor th frtility rdr.rtiontO strong (hnaus-
ing spiatiln)whil also not liminating th nw arrangmntwhn it first
ariss (th individual who first ars this mutation will nssarily ht-
rozygot). mpirial wok showd that rarrangtnnthtrozygotesr
somtirnsnot as stril as initially livd.Instad,studisnow suggstdif-
. frntways in whrh rarre1ngmnts ontriut to spiation (.g.,fftson
.s7hit.
romination)rthr tharr tlr dirt frtiliry,dutiorrsdisLrssdy
His mphasis on th importn of rarrangmntsWas orrt' ut his -
plantiorrfor how thy ausd spiation is not widly aptdtoday. ow-
vr' thr r ptions: on lgrrtstudy of Shrom1,syast showd
a dirt fft of transloation on hyrid fitnss(Dlnri t al. 2003).
Whit takld svral otl.lrontntious issus. H rviwd gnti data r-
garding spisdiffrnsarrd onludd tht tlrrWas no vidrrsupporting
rnst ayr's influntil viw of a ..gntirvolution'' that ours during spi-
ation, during whih a larg fration of th gr-romhangs us of nw s-
ltiv prssursaftr oloniztion. tVhit also strongly favord intrprttions
of symp:rtrispiation in many inst groups, suh as Rhagoltis fruit flis' still
studid today in this ontxt. ontrary to mally othr authors of his tim (and
ours), h ritiizd ..thologiai spiatiorrisrs''whl strongiy dvot sual
disrimination and othr haviors as ing of paramount importan to spi-
ation. Instad,h suggstdthat hyrid prolms may mot ommon arly
in spiirtion.Although his viw is not widly aptd toda,v' oyn zrnd Orr
(2004) also not th ovrmphasisplad y som authors on sul disrimi-
nation whn th rlativ importan of ffts from suh disrimination vrsrrs
hybrid proinrs i spirion has rrot rrdfinitiv[ydmrrstatd.
Whit's txt is a rihly dtaild rviw of th litratur on spiation.Th
disussions of nrpirial studis of spition ar tlrorough and loqunt.
!hit was an otstanding ytologist, ut lik mrny iologists, h am
onvind that th phnomna h studid wr of paramount importan in
spiation.Although som of his intrprtationsar unpopular today, nO on
an dout that this tt Was inf]untial irl motivting svral outstnding
studisof spiation.Som of his ontrvrsial viws may yt vindiatd;
many ontinu to ativ aras of study. This quot from his tt still ap-
piis: ..And, at th prsI1ttim' it wlld sm that swpirrggnraliztins
aout mthods of spiation should avoidd as fr as possil (it may
mor justifialto gnra|lzetn 2 -Yarstim)'' (Whit 1978, 24).
744 Mond to an

Th ltrntivtO trri
BIBLloGRAPY Was to find a wr,vttl s
framwork for suh a st
oyn, J. A.' nd . A. Orr. 2004. Spitioz. Sundrlnd,A: Sinaur Assoiats.
Dlnri, D., I. olson, S. Grammnoudi, I. N. Rorts, . J. Louis' and S. G. Olivr. lution in th 1930s and
200. nginring volutilnto study spiationin y:rsts.Ntr 422: 68*72, wr thmslvs prlgr
Ortiz-Brrintos' D.' J. Riland, J. H.y, and M. A. F. No<rr.2002. Romintion t w n p r o g r s s i o n i s nl
nd th lir,rgnof hyridizing spis.Gti 1 l6, ns. 2-3: 167_|78. frnrrvork fo nrrpro
Whit, Iv{.J. D. l978. lds of Spitit;.Sn Franjsl:W. H. Frtlra.
-.'.}J.
partly austh li
Th ook,s valu is r
o m i n s a s w p i n gh i
a o u t i n d i v i d u a ls i n t
Nlond to NI: The onpt of Progress also snsitizradrst,
in uolutionIr Biolog (Mihal Rus)
|BLloGRAPHY
volutinry idas ampopul:rrin N7strniviliz:ltionjust whn onprs
1996.ol
Rus' l\4iha|.
of lruman progrss wr flourishing. The Industrial Rvolution and th gowth Biollg.Cmbidg,,
of sin itslf ld duetd popl to pt limitlss progrss in human
knowldge and in human soity. Many ommntatos hav suggstd that
idologial lifs aout human progrss Wr a motivting fator in lifs .lvlorgan,Thomas I
aout volution nd spially in lifs aout th progrssivnssof volrt-
originlly traind as irn
tion. ilral Rus st out t irrvstigatwhthr prslrl lifs of sintists
th prinipal arhitts
aout progrssivnss in human soityand in volutilnwr orrlatd.
p r i o d 1 9 1 0 - 1 9 3 . 5H. r
ond to n is th rsr"rltof this study. Rus onluds that sintists'
ology or dii (193j
lifs aout th progrssivnssof volution wr indd orrlatd with
dmonstrating tht M
thir lifsaout soial progrss.This provids a hallngto th o1tivity
rod-shapd bodis founr
of th sintifilif in volutionary progrss.Judgmnts of progrssivnss
normous influn nlt'
annot dmonstratdon str:rightforwardmpirial grounds; thy ar nor.
in mryology and sp
mativ ssssmntsof what oul1ts as ..forward'' hirng. ould th mt.
his group rsolvd th qt
phorial irrfrnfrtlm soil progrss to v<tlutionryprogrss b takn
work of Ggor d
sriously by sintists?o nsrvr this qrrstirnRtls asks thr qustions
hypothtil onstuts1
aor'rt lrg numr of lristorial and ontmporaly volutionists.Fist, do
hrdity or rprsntd
a sintist'sidas aout iologial progrss orrspond to what on would
transmittd as disrtb
pt if th sintistwr inflr'rndy ids aout soial progrss?Sond,
p o r t n t d t s i n r i
ws that sintistonsious of th rlation twn soial progrss and io-
itd valu austh l
logial progrss?Third, to whilt xtnt did th iologial viws outstrip th
T. H. organ ws [-
vidn?h largr th gap twnvidenand thor, th mor likly it
r h n d o f t h U . S .( . i v
is that th sintistwas arrid i1rossthat gap y his or lrr soial iss.
h i s s s yo n h r i d i z . r t
Rus's answrs ar intriguing. First, almost ll nintnth-nturyvolu-
f a m i l y t h t i n l u d dh i s
tionists and most twntitl.l-ntuyvolutionists Wr progrssionistswith
gnral and ladr of }
rspt both to soity and to volution. But sond, thy sm to hav r-
Sott Ky (1779_184)
ognizd that this ratd a ias. Th ognitionis rvaldin diffrns-
th Stat Univrsity of
twn thir tratmnt of voltrtionary topis and rhir nonvolutionary
rrd Johns Hopkins U
rsarh.volution Was a t()pi ior puli ltursnd popular writings, br'rt
Willim Kith Brtl<lks(
it was usully not a topi fo lrrd<lrsintifi rs:rrh.Thomas nry
sis on th volutiony
Huly, Darwin's strongst sllppotr, was prft xampl of this tn-
C r u s t a n d A r h n i
dny. H wrot dozns of popular rtils on volution ut largly onfind
onludd that th gou
his sirrtifipuliations to PaPrs on omparativ morphology.

:ndrlnd, MA: SinaurAssorats.
lrts'. J. Louis,and S. G' olivr.
tionin ysts.Ntur422: 68-72.
A. F. Nor. 2002.Romination
tid116,nos.2_3: |67_\78.
.W.
Frniso: H. Frman.
^4..F.N.
rogress
Rus)
:rn ivilization just whn onpts
ustriai Rvolution and th growth
pt limitlss progrss in human
)mmntators hav suggstd that
/r a motivating {ator in lifs
)out th progrssivnssof volu-
rthrprsonal lifs of sintists
in volution wr orlatd.
y. Rus onluds that sintists'
ion wr indd orrlatd with
vidsa hllngto th ojtivity
.ss.
J u d g m n t so I p r o g r s s i v n s s
J mpirialgrunds;thy ar nor-
rward'' hng. ould th mta-
l volutionary progrss takn
| s t i O nR u s a s k s t h r
ttmporaryvolutionists.First, do
;s orrspond to what on Would
asaout soial progrss?Sond,
twnsoial progrss and bio-
1 th iologial viws outstrip th
nand thory, th mor likly it
gap y his or hr soial iass.
nost all nintnth-nturyvolu-
tionists Wr progrssionists with
}ut sond,thy sm to hav r-
nitionis rvaldin diffrns-
opis and thir nonvolutionary
ltursand popular writings, ut
iintifirsarh.Thomas Hnry
as a prft ampl of this tn-
s on volutionut largly onfind
nparativmorphology.
org 745
Th altrnativto trating volution as a popular (ut not a rsarh)topi
Was to find a way to study volution that did not assum progrssion. A
famwork for suh a study finally appared with th synthti thory of vo-
lution in th 1930s and 1940s, ut many of th arhitts of that synthsis
Wr thmslvsprogrssionists.only in rnt yars has th assoiation -
twn progrssionism and volutionism fadd. This is partly aus th
framwork for nonprogrssionist volutionary studis has advand, and
partly bausth iif in soial progrssionismhas fadd.
Th book's valu is not solly in its onlusions about progrssionism.It
omins a swping history of th sinof evolution with intriat dtails
aout individual sintists'rsarhs,prjudis,nd prsonal livs. It an
also snsitizradrsto sutl progrssionisttndnisin thir own thought.
BlLIoGRAPHY
Rus,iha|. |996. od to : h optlf Progrssi uoltior
Biolog.amridg,A: HrvardUnivrsityPrss. -R..
organ, Thomas Hunt (|866_1945)
originally traind as an mryologist, Thoms Hunt organ am on of
th prinipal arhitts of th Mndlian hromosom thory of hrdity in th
priod 1910_1935. H was th first gntiist to Win th Nol Priz in Physi-
ology or Mdiin (1933) for th work h and his laoratory group arrid out
dmonstrating that Mndlian gns wr physial parts of hromosoms' th
rod-shapd odis found in th nulus of all ukaryoti lls. This work had
normous influn not only in th rapidly dvloping fild of gntisut also
in mryology and spially volutionary iology. Th work of Morgan and
his group rsolvd th qustion whthr th hrditary fators postulatd in th
work of Grgor ndl (1822_1884) in th nintnth ntury wr mrly
q u s i o n s
hypothtial onstruts that wr onvnint for plaining rtain pattrns of
hrdity or rprsntdral, matril omponnts of th organism that wr
transmittd as disrt odis from parnt to offspring. This was not n unim-
portant dat sin ritis f th nw gntis maintaind that it was of lim-
itd valu austh si units, gns,had no physial rlity.
T. H. organ Was on in Lingtn, Kntuky, tn 1866, on yar after
th nd of th U.S. ivil \/arand th sam yar in whih ndl pulishd
his ssy on hyidization in pas (Pisum). organ am from a distinguishd
family that inludd his unl.}ohnHunt orgn (182-5-1864)'onfdrat
.War,
gnral and ladr of Morgan's Ridrs during th ivil and Franis
..Th
Sott Ky (1779_1s4), author of Star-SpangldBnnr.'' duatd at
th Stat Univrsity of Kntuky (now th Univrsity of Kntuky), h n-
trd Johns Hpkins Univrsity in 1886 as a studnt of morphology undr
.s7illiam
Kith Brooks (1848_1908). rivdhis PhD in 1891 with a th-
sis on th volutionary rlationship of th sa spidrs (Pynogonida) to th
rustaa and Arahnid. From a dtaild anlysis of arly mryology h
onludd that th group was most llslyrlatd to th Arahnida, ut that
746 rlrg

th two groups must hav divrgd from a ommon anstor in th distnt o.o YELLow' sPo.
o.' !iL I.
p a s t . A f t r a s m m r( 1 8 9 1 ) r r dt h r r y a ( 1 8 9 4 _ 1 8 9 5 )a t t l r N a p l s m a - ro wl]le oslN' IlRRY.

I.O BNOB}tIL
rin laoratory (Stazione Zoologi) working losly with th mryologist
Hans Dris'(1'867-1'941), Morgan gav up th largly dsriptivand sp-
i! ]FID.

ulativ sil-lof morphology and am ir omrittd primntalist.x-

primntation, h am tl bliv,povidd a mor rigorous and analytial
way to approah biologil rsarh than th splrlativ and untestal
phylognti hypthss of th morphologists. Morovr, th qustions -
primntlists suh as Drish wr asking onrnd important ilogial
prosssourring in th prsnt (for mpl, th mhanisrrrof mryoni
diffrntiation)rathr than histori1(phylognti)rlationships. olla-
oratd with Drish on a sris of primnts On th dvlopmnt of .' LAL lll.
tnophors (popularly alld om illis)' arin invrtratsthat hav a'. Ali'

suprfiial rsmlnto jllyfish. h itmnt of ing al to analyz

ompl physiologial or dvlopmntal prosssditly y manipulating
th organism or its rlvironmntonvind Morgarr this ws th dirtion in
whih modrn iollgy ndd to prod.
N4organhld th tahing positions during his arr:at Bryn Mawr ol-
lg (1891_1904), olumia Univrsity \\904-1928), and th lifornia ' LAL v.
..o sAL
Institutof Thnolog 092.8-1945), whr h foundd th Division of Biol-
ogv. Throtrglrout all ths yars h spnt su1rsat th arin Biologil
Laoratory in \/oldsol, assahustts, whih h and othr iologists 4o.t LIIAL lv.

onsidrd th Amian Npls. During his Bryn Mawr priod organ

workd mostly on prolrs of primntal myology, using organisms
RLrDlNARY
suh as frogs, arthworms, nd sa urhins. lt was only ftr arriving at o- c . . rRD,
sz.o BAR.
lumbia Univrsity' starting round |907, that h am farilir with th
fruit fly, Drosophil mlogstr. Th work for whih h and his laoratory
group at olumi susquntly (aftr 1910) am most famous, sta-
lishing th ndlian hromosom thory of hrdity,was inititd and ar-
rid out during th olumia yars (sfigtr).Although h transportdhis
..fly la''
to Psadna in 1'928,his administrativdutis took him away from
dy-t-dy fly rvork, nd Whn h did rtlln t th loratory' prolms of
mryoni dvlopmnt rsufds his min intrst.
Two mjor aspts of organ's Work rlt to th dvlpmnt of volu-
tiorrary thlry in tlr twntith nttrry.Th first r,vrs his arly opposition to
Darwin's thory of natural sltion (1903_1915) as a mhanism for how
volution, spially daptation, ould our. Th sond ws his skptiism
aout oth th Mndlian and hromosom thoris of hrdity.
Morgan's initial ljtionsto Darwin's mhanism f natural sltion wr
void in his ook utlltil d dpttil (|903\' irr whih h argud that
sltion ould nvr produ wholl1' nw spisy ating on slight individual
diffrns (rfrrd to at th tim as ontinuous or flutuating variations), as homs Hunt organ and
Darwin had laimd. Baus suh variati-lnsWr s-rall,vn if thy on- nd lvin B. Bridgs)rnp
..swampd soms of Drosophil. Tis i
frrd som advant;.rg,thy would tnd to gt out'' in sussiv
dli Herdit (1915)
gnations. In additiln, Mrgan argud tht flutuating vari:rtions ourrd
only rarly and 1' han, so that slti<lt.l, ting on suh vritions, ould
 orgn 747

:ommon allstor in th distant L o.o YLLow' so. . o sAK

(1894-1895)at th Napls ma-

t3i,T,l'tl.*'n,
.n.
f N.
g loslywith th mryologist
th lagldsriptivand sp- "" '-"*" rYlsg.
ommittdxprimntalist.x- [
a mor rigirousnd analytial
th spr"rlativ and untstbl
s . o r o v ' h q r r s t i o n s -
1,.,..u* - lc.. D.{II$

onrndimportant iologial
ll' th mhanisnrof mryoni
1nti) rltionships.H ollab-
imnts on th dvlopmnt of [-'""*- .' PlN' l.{'.

marin invrtratsthat hv a
tmnt of ing al to ana|yze
)sssdirtly y manipulating
o g a nt h i s w a s r h d i t i o n i n
.{?
[""":":- LA.

ng his arr:at Bryn Mawr ol- .4o'. PU&PL, {q lDN.

904-1928\' and th aliforni

1"":-',-:",
h founddth Division of Biol_
Lmmrsat th Marin Biologial [';;;;:
, whih h and othr iologists
his Bryn awr piod N4organ .l.,o vsIoIAL

l m r y o l o g y .u s i n g o r g a n i s m s
.8. oNY.soot.
It was only aftr ariving at o. Lo,! oBltlD.
B^R.
lrt lr m familiar with th '.o
.u*',. ..r UYD.
<for whih h and his laoratory
l0) am most famous, sta-
".
lf rdity,r,irsinitiatd and ar- LALe

r).Although h transportd his l.3!'l

.ativdutis took him away fr<rm
'n to th laoratory, polms of
in intrst.
at to th dvlopmntof volu-
: f i r s tw a s h i s e r l y o p p o s i t i o n t o
- 1 9 1 5 ) a s a m h a n i s mf o r h o w
L.Th sondwas his skptiism ..i.! aRC.

: thoisof hrdity.
:hnism<lInturel sltionrv .eo.o sP.
(90|,in whih h argud tht "r.. oRULA'

isy ting on slight individual

uous or flutuatingvaiations),as
)ns Wr small, vn if thy n-
gt ''swampdout.' in sussiv
t flutuating varitions ourd
' ating ln suh vrritions,ould
homas Hunt organ and his young assoiats(A. F{. Sturtvant'H. .I.ullr,
and alvin B. Bridgs)mppd th rd of gnson th fou (pairsof) hrom-
somsof Drosophil. This is th frontispiof thir ook, Th ehism of
edelin tlerdit (19 15).
748 org

lrvr p()duth fin-trrnladaptations sn thror'rghoutth living world. B. homologous hromosom

twn 1903 nd 1910 Morgn advoatdHugo d Vris's (1848-1935) muta- th signifianof th hrt
tion thory, whih limd tht nw spisros in on stp y lrg-sal ogist Morgan had a strong
mutations that luld mak an offspring a diffrnt spis from its prnts. oris bus thy sm
This thory, for whih thr appard to sound xprimntal vidn in d prformation, whih hd
Vris's nlodl organism, th vning primros (othr lmrchi), wzts w a s m r | y h u n f i l l d i l r g.
spilly :rppling to Morgn nd nrany of his ontrnporzrrisnot only - Rfring ompl nl ]l
tls it irumvntd th ojtions to Darlvirrin thory outlind arlir, lrut ls in th grm lls, tl
also aus it ws laimd to an eprimtl alproah to volutir.D planatory ut in fat ultir
Vris wirs al to hyridiz diffrnt varitis and spis of oother zlnd Ironially organ g
produ th supposdly nw, mutatd spis at will. Although d Vris's mu- th spring of 1910 and tl
ttion thory was latr abandlndausit was found to th rsult of :r from his own studislf D;
varity of hromosomal anomillis puliar to elthr (and thus did not in his olumia lorzrro
produ nw spis),it svdfor nany at th tim, lik organ, as vial i o n s r fd V r i s i e nt y p i l l
ltntivt Drwinian nirtualsltion. lsrvd among his ultu
Dr'ling th sam priod (190-1 910) organ was also skptial f th ys. It ws a mal, and h
ndlian thory as an planation of th rsults of hyridization and th red-yd. But whn r
hromosom thory as an xplanation of suh phnomna as s dtrmina- ratio of roughly on whit
tion. To organ th Mndlian thory, lik numrous othr partiulat asially th ptd M
thoris of hrdity in th lat nintnth ntury (rlrosof Darwin, rnst rozygous fo ssir't
akl, August !7ismann' or ilrl Nigli' to nam trv),was ouhd in rd, whih Was d()n1ll12
trms of hypothtial ..fatrs'' or prtils nd w:rs thrfor ompltl n < l r dr h r l l t w h i t -
sptrlativ.Asid from tlr fat that th istnof ths supposd f:rtors y _ o l o ri n h r i t a n ( 1 9 1
ould not onfirmd, Morgan's ojtionsto Mdlian thory wr thr- must somhow sgrgiltv
fold. First, if, s ndlians smdto laim, sx Wi1sa hrditary trait, tl.r mts.Bur this pl.rt
ndlian thory ould not aount for th 1:1 s ratio found in most ani- apparnt iologial asisf.
mals (and plants). Sond, Morgan thought that rrdlians trtd phn<l- soon am tO s' th firt
typi trits (an organism's pparan) as disrt, sharply diffrntitd s o r y h r o m o s o m s( i n p r
atgoriswhn in rality most (for ampl, nimal o21tolor) rngd ovr solns' vn if on of tlrrr
a wid sptrum. orgtrr .rlso notd that dli:.rnssmd to invnt as .raskdy th rd ftor t
many pairs of fators as tlry r-rdd to plain any givn hrditary ondi- vr, in mals, with lnly ol
tion; fr xampl, if on pair of fators would not srrffito plin a parti- .raskdnd thus woull sh
ular pttrn of hrdity,two wuld invokd (s in assof pistasis,whr gan Was onvind tht N
on pair of fators affetsth xprssion of anothr). H also qustiondth sis if it wr assuml1
Mndliarr ssumption of sggation'th sparation of the two fators fr l 9 0 9 l a l d8 . , 7 s( ). ( . t
any trait 1uringgrm-ll formatiorr so that h ndd r-rpin a diffrnt mosom.
gl-nt. S u s q u n t l yt h i d a r
.Wit
gard to th hrornosom thory, until l9l0 organ limd that som was givn futhr sr
thr wtrs nl vidn that lrrrlmosomshd anything to do with hrdity, ( 1 8 6 1 - | 9 2 6 ) a n d o t h sh
i n l u d i n g t h l a i m y h i s o l l a g u . B . W i l s o n ( 1 8 5 6 - 1 9 8 ) a n d f o r m r mor trits togthr. Alth
studnt Ntti . Stvns(186|-19|z) that th so-alld assoryhromo- nomnon in trms lf th
soms (wht w today rfr to as and Y) ar rsponsilfor dtrmination his studnts,did. Indl,I
of s in a wid varity rfnimals. Although hrlmosomsWr larly vis- atr-rzrllv mapping gn's
il in mirosops at the tim' nd nruh dsriptiv work had n don gan notd, linkags appa
dtailing thir movmnts itr mitosis (th pross of distriutirrg th th two homologous ro
hrlmosoms during ll division) and miosis (th proess of sparating lld tually hanglp

throughoutth living world. B-
;od Vris's (1848-1935) muta-
aosin ()n stp by larg-sal
frntspis from its parnts.
und primntalvidn in d
: (othr lmrki), was
his ontmporaisnot only -
nian thory outlind arlir, ut
tl approah to volution. D
; and spisof eotbr and
at will. Although d Vris's mu_
was found to b th rsult of a
o othr (and thus did not
lr tim' lik Morgan' as a vial
)rgan Was also skptial of th
rsultsof hyridization nd th
h p h n o m n a s s d t m i n -
k numrous othr partiulat
ntury (thos of Darwin, rnst
to nam a fw), was ouhd in
; and was thrfor ompltly
stnof ths supposd fators
to Mndlian thry Wr thr-
1' s Was a hrditry trait, th
l : l s r t i of o u n d i n m o s t a n i -
that Mndlians tratd phno-
disrt,sharply diffrntiatd
:,animal oat olor) rangd ovr
M n d l i a n ss m dt o i n v n t a s
,lain any givn hrditary ondi-
l d n o t s u f f i r x p l a i n a p r r i -
d (asin assof pistasis'whr
anothr). also qustiond th
lparation of th two fators for
at ah ndd up in a diffrnt
until 1910 Morgan laimd that
ad anythingto do with hrdity'
W i l s o n ( 1 s 5 6 _ 1 9 8 )a n d f o m r
: th so-alldassoryhromo-
lr rsponsilfor dtrmination
;hhromosomsWr larly vis-
dsriptiv work had n don
h pross of distriuting th
: i o s i s ( t h p o s s f s p a r a t i n g
org 749
homologous hromosoms duing rdution division in gamt formation),
th signifianof th hromosoms was still unlar. Finally, as an mryol-
ogist orgn hd a strong vrsion to both Mndlian and hromosom th-
oris aus thy smd lik rvrsions to th long-rjtd thory f
prformation,whih had laimd that th pross of mryoni dvlopmnt
was mrly th unfolding of miniatur mryoni parts into adult haratrs.
Rfrring ornplx and dynami iologial prosssak to invisil parti-
ls in th grm lls, organ laimd, might sm at first glan to x-
planatory ut in fat ultimatly plaind nothing.
Irlnially Morgan gan to hang his rind on all thr thoris twn
th spring of \9| and th wintr of 1'911. Th major imptus am first
from his own studisof Drosophila. organ had gun dingth fruit fly
in his olumia laortory around 1908 to s if h ould produ muta-
tions of d Vrisian typ in animals. H had gottn nowhr whn on day h
osrvd among his ulturs of normally rd-yd flis on that had whit
ys.It was a mal, and h rd it to a rd-ydfmal;th offspring wr all
rd.yd.But whn h rd ths offspring with ah othr, h otaind a
ratio of roughly on whit-yd fly for vry thr rd-yd flis. This was
basially th ptd ndlian ratio for a foss twn two parnts ht-
rozygous for a rssivtrait (in this as whit ws said to ressivto
rd, whih was dominant to Whit). Mor important, howvr, rgan
notd that ll th whit-ydf]iswr mals. In his first pulishd papr on
y-olor inhritn (1910), rgn assumd that th fator for whit y
must somhow sgrgatwith th fator for malnssin th formation of ga-
mts. But this planation Was not wholly satisfatory sin thr Was no
apparntiologial asis for assuming suh sltivsgrgationunlss,as h
soon am to s' th fators for dtrmining y olor rsidd on th as-
sory hromosoms (in partiular th ). Sin fmals had two hromo-
soms' vn if on of thm had th nw mutant fator flr whit, it would
maskd y th rd fator on th homologous' or prtnr hromosom. F{ow-
vr,in mals, with only on hromosom' th mutant fator would not
m a s k da n d t h u s w o u l d s h o w u p a s t h w h i t - y dp h n o t y p . B y 1 9 1 1 M o r .
gan Ws onvind that Mndl's thory ould givn a onrt matrial
asis if it wr assumd that fators, or what !ilhlrn Johannsn had in
1909 lald8s' oclpid a spifiphysial rgion, or lous, of th hro.
mosom.
Subsquntly th ida of gns as arrngd linarly along th hromo-
som ws givn furthr support y osrvations On what \/illiam Batson
(\86|-1926) and thrs had idntifid as linkag, th inhritn of two or
mor traits togthr. Although Batson himslf did not intrprt this ph-
nomnon in trms f th hromosom thory, othrs, inluding organ and
his studnts,did. Indd,Morgan usd th ida of linkag to dvis a way of
atually mapping a gn'sptlsitilnon th hromosom. on oasion, Mor-
gan notd, linkags appard tl rokn and th gns romind, as if
th two homologous hromsoms on whih a group of gns wr llatd
had atually hangdparts:
7.'0 org

A B al parntal hrroslm N4ainshin,J. 1991. Thol

i Amrin Bfu'l,'g1,
Fml parrrtlhomosom N4organ' T. H. |90. ull
1910. S liritdi
Y
I 1916. A ritiqe
Univrsity Prss.
A A4orgn, T. H., . H. Sturt
offspring hromosoms hnism of dl,
B Shin, I.' and S. \/rol.19
Lington: Univrsity .
Morgan supptlrtdthis ida from work y a Blgian yt()l()gist'F. A. Janssns
(186-1924), who had dsrid in a 1909 ppr what h a||edchis-
mtpie, or th intrtwining of th two homologous mmrs of hromo- lV1ullr,Hrmann Jt
som pair during miosis, and what appardt thir susquntraking .W.hn
H. J. ullr was
and rjoining. organ nd his studnt A. H. Sturtvant (1897_\97\| rea-
hiit on evolution at
sond that th frquny with whih rominatiln ould osrvd in
mmrd ing imprs
offspring was a funtion of th distan apart lf any two gns on th hro-
siz and anatomy hang
mosom' and this providd a way to stimtat last th rltivpositions of
known as gntiist ar
gns with rspt to ah othr on th sam hromosom. This mapping
tht -rys indu gn
produr soon am for Morgan and his laoratory group-inluding, in
his profssional and prs
a d d i t i o n t o S t u r t v a n t , a l v i n B . B r i d g s ( 1 8 8 9 _ 1 9 8 ) a n d H . J . M u l l r
(1890*1967)-th asis of an laoratrsarhpro!]ramthat was tndd a t t n d d o l u m i a U n i v
two grat Amrian iol
by othr workrs to hundrds of organisms (orn, th mous, and th rat) .!7ilson
organ. was a
ovr th nt 0 yars. It was for stalishingth validity of th hrlmosom
gnration of iologists
thory of hrdity that organ was awadd th Nol Priz in 1933.
volvd in hrdity, dv
organ's own turnaout with rgard to gntisand th hromosom th-
vntually provd lil/ilso
ory ld him to s that Mndlin variations-th diffrntwnrd and
mlnogstr. In his snr
whit ys or twn wild-typ and add wings-might atly th
foundd a iology lu
sorts of varitions on whih natural sltionould at. I|916, for a sris
Bridgs njoyd sharing'
of ltursat Printon (pulishdas A ritiqu of th Thor of ulutio),
that was atingsuh s
organ notd that th many variations h had osrvd t Droslphil wer
.Whras t h k y f i g u r si n p u t t i n
disrt,stal,and inhritd. rlir h hd argud that small indi-
fruit flis univrsally kno
vidul or flutuating variations would always tnd to swampd, th dis-
Mullr rivd his Pl
rtndlian varitionswould not. Ths ould aumulatdindfinitly
whn N4organ and Sturt
nd thus ould lad vntually to th formation of a nw spis.In his ltr
him to a study of what h
writings on volution (1925 and 192) he arplifid ths viws, larly hv-
10 yars to pulish ths
ing om to apt th asi outlins of Darwin's thlry. It was for oths,
xtndd ovr many gn
howvr' with mor mathmatial lanings to us th nw glltisto formu-
of how Darwinian natur
iat th matur form of Darwinian thlrvtht m to known as th vo-
g n st h t o u I d i t h ri n
lutionary synthsrs.
organ systm or trait. N
B|LIoGRAPY v a r i a l , n d h i s o l a t d
Alln, G. 1968'ThomasHunt orgn nd th prolrnof nturlsltiln. Jourl ations. ullr, lik Darr
of th Histor of Bkllog 1: 11_19. nd that nw organ syst
1'978.hlmsut lrg:Th tl His Si,l)rinton, NJ: N4ullr workd first at
PrintonUnivrsityl)rss.
of Txas in Austin. !7hili
1980.h volutionarysynthsis: rgirrrad nirtualsltionrvisitd.
In
. Mayr and !. Povit.t,ds',Th uolutiorS),thsis:Prspctit,s
o th mutation and th gn. l
Uifititlof Biolog,356_.]82. anrri1g,
MA: HarvrdUnivrsityPrss. individul gn and argt

omos()m
:hOmosm
Blgianyt()logistl. A. Jarrssns
)9 papr whirt h a||cd cbis-
lologlusmmrs of a hromo-
l to tlrirstrsqr"rnt raking
H. Sturtvant ( 1 8 9 1 _ 7 9 7 1 )r a -
rlintilnould osvd irr
rt of an,vtW()]nsln th hro-
t lastth rlativpositiorrsof
m h o l n o s r l m T. h i s m a p p i n g .
laoratorygroup-inluding, in
( 1 8 8 9 _ 1 9 8 )a n d H . J . u l l r
:arhprogram tht was tndd
s (orn, th lnous, arrd th rat)
g th validit of th hromos()m
d t h N o l P r i z i n 1 9 3 3 .
;ntisnd tlr hromosom tlr.
i-th dif{rntwn rd and
Ld wings-lnight atl,v th
ln ould t. In 19|6, for a sris
iqe o| th Thor of ullution\,
had osr,lin Droslphil were
ir h hd argud thirt smll indi-
rys tn{to swilmpd, tlr dis-
ouid arrmulatdindfinitl
Ltionof a nr'vspis.In his latr
rmplifidtsvirvs,larly lrv-
arwin's thlrv.It was for othrs,
to us th nw glltisto flru-
hat arnto knwn as th vo_
prolmof ntural st.ltitl.
.|ottll
nd is Siltt,.Prill:tlr-r,
NJ:
ln and nlltural sltin visitd.In
r 51,thesis: Prsptit'so th
l' N,IA:Hrval Univrsit1.Prss.
ullr 751
airrshin, J ' 199 1. Thomas Hunt rgn. haptr 8 of Trsformig Trditios
i mric Biologl, 1890_1L)1.'.Bltimor:Johns Hopkins Univrsity Prss.
lvlorgn, T. H. l90. ulltitl d dpttil. Nw Ylrk; N1milln.
191().S linritd inhritani Droslphil.Si 2: \2-122.
1916. ritiqu lf th Thulr o|-ullutiltt.Printot-t,NJ: intn
Univrsity Prss.
rgan, . H.' A. H. sturtvnt,. .|.N,lullr,nd . B. Bridgs. t9 | 5. T
hism of dli Hrdit. Nw York: Holt.
Shin' I., ar-rdS. !(lI. 1977. hlms Hut lrg, ilr lf Gtis,
[,ingt()n:Univrsity f Kntuk Pss. -C.,..
ullr, HrmannJosph(|s90-1967)
!hn H. J. ullr was aout l0 yars o[d, h wnt with his fath tO s n
hiit on volution at th Amrian usum of Natural History. H r-
rnmbrd ing imprssd y th small siz of fossil horss nd how thir
siz and natomy hangd ov rnillions of yars. Although Nlull is st
known as a gntiistand won his Nol Priz (in |946) for work proving
tht _rays indu gn mutations, lr onsidd himslf an volurionisin
his profssilnaland prsoIrallif. ullr was third-gnrtionAmrian and
attndd olumia Univrsity ln sholarship. Thr h tlok ourss with
two ]atAmrin iologists' dmurrd Behr wilson and Thoms Hunt
.silson
Morgan. was a ytologist whos writings hlpd onvin a youngr
gnrtion of iologists who t<lok his lsss that th ll was dply in-
volved in hrdity, devlopmnt, an.lvolutio. <lrgan' skptial at fist,
.Wilson's
vntually provd thris y using th tiny fruit fly' Drosophil
mlnogstr.In his snilyar Mullr mt two soplromoswith whom h
foundd a iollgy lu. Alfrd Hnry Sturtvant and alvin Blakman
Bridgs rrjoydsharirrgwith ullr thir idas On th nw 6ld lf gerrtis
that was rating suh a sti among thir tahrs.Ths fiv indlviduals wr
th ky figurs in puting togthr th fild of lassial gntisand making
f u i r f l i su n i v r s l l l l yk n o w n m o n g s i n t i s t s .
ullr rivd his PhD for working out som omplitis that aros
whn orgn nd sturtvnt disovrd rossing ov. N{ullr lvok Id
him to a study of what h alld gn-haatrrlations. It took him aout
l0 yirrs to pulish ths rsults austh pimnts Wr tdious :lnd
tndd ovr mny .nrtions'For ullr it was ruial dmonstration
of how Darwinian natural sltionworkd through th sltionof modifir
gnesthat ould ith intnsif1'or dinrinish hif gn'snrajor ffton n
ogan systm or trait' Mullr usd gns for wing shap that wr highly
vrial,and h isolatd and mappd th modifir gnsthat usd th vri-
ations. ullr, lik D:rrwin, rognizd that volution is a gradual proess
and tht nW organ systmsdo rrot ris sr.rddnlyut a uilt up gradually.
ullr workd firs at Ri Univsity in Tas nd thn t th Urrivsity
of Ts in Austin. Whil at ths institutions h wokd on th prolms of
mutation irnd tlr grr. stitd th trr muttio to hang in th
individual gn nd agud that th gn Was th asis f lif aus all
752 ullr

omponnts of th ll and th organisnr wr produts of gns, as shown y mans to a radiatin dls

th anormalitis produd in thm whn th normal gns Wr altrd y tually soity would hav
mutation. Mullr also masurd th first spontanous mutarion rats at a to rdu this mutationa
tim whn thy wr so rr that h said that finding a nw mutation was lik amount of tim nd ost
finding a dollar ill on th strt.B 1926 Mullr had workd out th tools
|BLIoGRAPHY
for masuring mutation rats y dsigning fruit.fly stoks in whih h ould
dtt a lass alld rssivlthals. If n of ths wr indud on a nor- arlson,. . 1'98.G
mal hromosom in a sprm' th resulting daughtr luld rd, and ullr.Ithaa,NY:
Mullr' H. J. 1968.Studrc
among hr sons on atgory of hr two hromosoms in hr immatur r-
Blomington:Indiana]
produtiv lls would prdu frtiliztions that killd half hr sons. Th his paprs,and this l
othr half of hr sons would b killd y th hromosom ullr dsignd.
This hromosom' alld lB (ontaining a supprssor of rOssing ovr, 1
a lthal, l; and a visil dominant y shap alld Bar ys, B) allowd
Mullr to min thlusandsof vials of ontrol and -rayd dsndantsfor
vials ontining only daughtrs and no sons. found ths in aundan
with th doses h usd (aout 4,000 rontgns).Thos doss ar 10 tims
gratrthan th dos that would kill half of all humans xposd to it. It is not
that fruit flis ar hardir than humans. any yars latr Mullr showd that
-rays rak hromosoms, as wll as indu gn muttions. Brokn hro-
mosoms in dividing lls lad to ll dath or' if ths lls ar sprm' to
aortd mryos. IndpndntlyBaraa lintok showd th sam yl
of raking lading to ll anormlitis in orn (maiz) nd alld this th
rakage-fusion-ridgyl. ullr immdiatly rognizd that this yl
in fuit flis and maize must th asis of radiation siknss, spially
whn tht ourrd among tns lf thousands of atlmi-omvitims in Hi-
roshima and Nagasaki. ullr rivd his Nol Priz at a tim whn th
tomi ag Was ing ushrd in, nd h quikly am an advoat fr
radiation prottion in industry, mdiin, and th military appliations of
tomi Wapons.
N4ullr mad many ontriutions to th study of voiutin. was th first
to propos an intrprtation of why po|yploidy (having thr or mor stsof
hromosoms instad f two) is rar in nimals and ommon in plants. In
animals s is usually dtrmind y s hromoslms,whih ar rar in
plants. This maks it impossil for triploid nimals to produ frtil mals.
H dmonstratd that th Y hrornosom (not normally found in fmals)
gradually loss its gns and shrinks in siz (ovr millions of yars of volu-
tion). Mullr also anlyzd hyrids of th rltd spis D. melogster
and D. simuls and sudd in gtting on of th tiny fourth hromo.
soms of simuls into an othrwis a|\-mlogsterfl..l/hat impaird
ths flis Was not som vagu physiologial inomptiility ut spifign
diffrns twn th two that h was al to idntify.
In th 1950s ullr turnd his attntion to human volution and workd
out quations to follow th fat of gn mutations in a populatiln. H sti-
matd that in th asnof ntural sltionth load of mttions prsnt
in a population would doul in aout four nturis.H also stimatdthat
this douling of th load lf mutations Was th quivalnt of sujtinghu-
 uIIr 7.'.

]produtsof gns,as shown y mans to a radition dos of aout 150 rontgns.Mullr sugg,std that vn.
h norml gns wr alterd y tually soity would hav to prti som fom of voluntry sltivrding
pol-ttan()usmutation rts at a to rdu this rrrr'rtational
load' or humanity would suffr disproportion.rt
finding nW mutation was lik amount of tim and ost to path up its infirmitis.
ullr lrd workd out th tools
BIBLIOGRAPHY
:ruit-flystoks in whih h ould
of thswr indud ln a nor- arlson,. A. 1983' Cs' Rditil,d Slit:h Lif d Work of H. .[.
uller.Ithaa,NY: ornllUnivrsitPss.
ng daughtould rd, and
r.rllr,H. J' 1968. Studisi Getis: SltdPprsof H'.|, ullr.
hromlsonrsirr hr immtur r- Blomingtorr: IrrdianUnivrsit1,
Prss.[Mullrsltdr.ts fronrmost of
:ls that killd hlf hr sons. Th his pprs,nd this lokontainsa lmplt iliogrph1'
lf his works.]
t hrmosomullr dsignd. -',.
l supprssorof rossing ovr, ;
ap lld Bar ys, B) allowd
'trol and -rayd dsndantsfor
rs. H flund ths in aundan
:gns).lros doss r 10 tirns
all hrnsposdto it. It is not
1y yars latr Mullr showd that
t gnmutations. Brokn hro.
.h or, if ths lls ar sprm' to
[lintokshowd th sam yl
L orn (mt-riz)nd alld this th
diatl'rlgnizdthat this yl
of radition siknss, spially
ds of atlmi-omvitims irr Hi_
; Nol Prize at a tim whn th
quikly am an advot for
and th military appliations of

:ud,vof voltion. wlsth first

idy (havingthr or mor sts of
rimals :rnd ommon in plants. In
, hih ar r in
h o m o s o m sw
l nimalsto produ frtil mirls.
(not normally found in fmals)
l (ovrnrillions of yars of volu-
rlatd spisD' melogstr
on of th tiny fourth hromo-
ellgsterfly. What impaird
I inompatiilityut spifi gn
l to idntify.
to human vlution and wlrkd
t:rtionsin populatiln. sti-
ion th lod of mutations prsnt
.ntttris. H lso stimatdthat
; th quivalntof sujting hu-
 vn as lt appars supr
and plants with digitl l
th distriution of plnt
athr than survyors'ir
ral faturs hav hang
In partiular, as linlat
r a l i s t sa p l a y i n g r i t i
organisrrrs in rspons tt
ar providing th raw m
Natural histry tati()n tl rpidly han

Ntural history is th arftrlosrvation of niltur, from study of how soil BlLIoGRAPHY

harrgsfrom on rgion to anothr to how fmal uttrflissarh for a Brtram,W., and F. Hp
suitl plat upon whih to ly thir ggs.Th prtitionrs of natural his- d i ti , A
. t h n sU
: niv
tory ar trmd ttturlists'and wll for sintistsaIld thnrslvsgol. Hilldolr'B.,:rnd. O. \
ogists rr otanists, indivlduals who studid th ntural world wr ;rlll p lortil.mrid
W i l s o n ,F . .o ' 1 9 9 4 .N t u r
naturIists.A tru naturlist r"rltivats prtis in aras from tlr ours ot
rivrs to th haitats of plants.
Ntural histor' was th for.lrrdatirrr
for tlr dislvry of volr.rtiln.Natu.
ralists lik Jan-BaptistLamark and rasmus Darwin osrvd th haits Nturl Selction
of anirrrls rrd n<ltdtlr mtlr rwn haits ad partiLllervalrrs of Philip Slrppard's Ntur
traits, lik th oft-itd ampl of giraffs ploiting thir long nks to for- just flrth ntnary
ag in trs.Nturalists notd t rlarionshipstwnslil tys nd th o- nrjor onrriution to t
urrn of individul spis,nd naturalisswho travld widly rordd not last thir lllow io
th distributions of plnts and arrimalsin spa,notillg arfully whr simi. sltion was norv a fulI
lar spisrplad on anothr gographially.lrarls Darwirr's apPrla- languag of Thomas Ku
tion of natural history ld him to osrv lrow mtrh th tortoiss of th fat, and lnddh rv
Galipagos diffrd in th shap of thir shlls from on islnd to nothr and saying that although th .
how th ak sizs and shaps lf th finhs of th arhiplago varid with th ook is disppointin
thir food haits. Whn Darwin oupld thsosrvatilnswith his appri- btrt ..in survying mldr
ation of th young ag of th ahiplgo' th ida of volution was plantd it is muh lss satisftor
firmly in his rind. vryon gd,:lus
Although th inrptus for th disovry of volution grW out of ntural Th work mplifis
histol.},'th mpiril study lf natural sltioll was drivn y it. Tlr British w s l l d h s y r r h t it
.Ufldon's
naturalist '.F. R. osrvations of mrirr animals' <lupldwith Darwinism. First, th o
his ppriation of th anging natur of thir hitt, ld hirn to gin tion undrpinnd y lVI
what might wll alld th first attmpt to dourrrntth atitln of natural 50 yars aftr ndl's
sltion (sth alphatial ntry ..\' F. R' Vldon'' in this volum). Nat- Wr nor widl kno,tt
uralists wh hd rnad rful studis of individul varition in a haratr sntials. Thos intrstd
lik srrailolor nd anding, mindful of Darwin's hypothsisof ntural sl- tion, pulishd fiv yar
tion, gan to study th fats of individuls with liffrntfatursand thus and Frarris Crik, rrk
gav ris to th quantitativ study of sltion.Th thousands of studis of howvr, not only is DN
naturl sltiontlrat lrav rrpulllishd ovr th dadsar th linzrld- r.rtthr is also a whol
sndantsof th work of ths lutiful naturalists. work of popl lik Ri|
Although gln through th prlfssionljournls of volutioirary ioi. lulr diffrensin <lg
ogy may suggstthat natural histoy is a lost pursuit, it ontinus to thriv and thniqus.
7-t1
 N a t u r a IS l t i l ann d H r d i t y 755

vn s it appars suprfiiallymuh hngd.Naturalists doumnt animals

nd plants with digital imags rathr than paintings anl drawings and map
th distriution of plnts and animals with gogrphi information systms
raththn survyors' instrltmnts.Altlrough th tools tlsd tO dsri natrr-
r| faturslrzrvhangd,th rrdto do sl and th vlin doing so rmaitr.
In partiular' as limat hangs zrnd humn ativity ltr landsaps' natu-
ralists ar playing ritial rols in doumntinghangs in th distriutions of
organisms in rsponsto ths altationsand through thir doumntation
irr providing th ri]w matrial for sttrdyingwhat may :r nw ra of adap-
ttiOnto a rapidl,vhanging arth'

BlBLIoGRAPY
lf natur'fronr a study of how soil
ow fmal buttrflis sarh for a Bartral,W', and L-.Harpr. |998. Th Trulslf WiIIim.lrtrI:Nturlist's
ditio.Athr.rs:Univrsityof GorgiaPrss.
s. Th prtitionrsof natural his-
. sintistslld thmslvsgol- olldolr,B., nd . O. \ilsn. |994. .|orto th Ants: Storof Sitifi
pllrtitl.
arridg'MA: BlkllpPrssf HrvdLJllivrsity Prss.
id th naturl lvorld wr alld Wilson,, o, 1994.Nturlist.Washingttlll' D: IslandPss. -].,
)rtisin irrasfom th <lr.rrs
of

th disovryof volution. Natu-

Nturl Sletion nd Herd,b/ (Philip . Shppard)
tsmusDarwin osrvd th haits
n haits nd partiular valus of Philip Shppard's Nturl Sltiotld Herdity, whih ppard in 1958'
l plitingthir long nks to for- just bfor th ntnary of hrls Dirrwin's n th ()rigi of Spis,was
hips trvrrsoil typs anl th o- a rlajor lntriutiollto th ffort y Britislr volutioniststo prsuadpopl'
lists wlr<ltrvldwidly rordd nt last tlrir flllw iologisrs' rlrat th thory o[ voluti<lnthrugh ntllral
..paradigm,''in th
ipa' notitlg arfully whr simi- sltionWas now a fully funtiorrirrgra of rsarh'
ially. harls Darwin's appria. languag of Thoms Kuhn (1962). This was still not a gnrally aprd
,, how muh th tortoiss of th fat, and indd th rviw of th ook in Hrdit was amazingly hostil'
lls from on island to another irnd saying that although th writing was lar, ..onsidrdils a whol, howvr,
hs lf th arhiplagovarid witlr th ook is disappoirrting.''It might dtl for ..advandUnivrsity studnts,''
..in
hs<rsrvations with lris appri_ t sr-rrvying modrn viws ot-tth thory of ntr-rrlsltiot,howvr,
th idirof vllutionwas plarrtd i t i s m u h l s ss : r t i s f a t o r y(' 'R s 1 9 5 8 , 5 2 3 ) .O v i t l u s l y , v n a k t h n ,n o t
vryonagrd,austh book was to go through flur ditions.
. of volution grW out of natural Th work mplifis autifllyth asi strutur of what in Amria
ltionwas drivn y it. Th British ws lld th synthti thory of vrltionand in ngland was alld no-
; of marin :rnitlrals,oupld witlr Darwinism. First, th or of th thory is prsntd.This is natural sl-
of thir lrirbitat,id him to girr tiorr undpinnd y Mndlian gntis.Rognizing thirt vn mor th
to dottrntth ation of r-ratul 50 yars ftr ndl's laws wr rdisovrd' tlr si idas of gntis
. R. !ldon'' in this volum). Nat- Wr not widly known, Shppard thrfor givs a short primr on th s_
individual variation in a hratr sntials.Thos intrstdin th diffusion of idas will not that th first di-
lrwin's hypothsisof natural sl- tion, pulishd fiv yars aftr th grourrdrakingwork of Jams \i7atson
Llswith diffr-rtfaturs nd thus and Franis rik, maks no mntion of DNA. By th fourth ditin \1975),
;tion. Th thousarrdsof studis of howvr, not only is DNA introdrrd omfortaly and without ommnt'
ovr th ddsar th lirrl d- but thr is zrlsoa whol nlv lrapt on protin volution, drawing on th
lralists. wtlrk of popl lik Rihard Lrvrntinwho had dislvrdsignifiant mo-
nal journls of volutionry biol- lular diffrnsin rganisms, 1iffrns unrmarkd y lassial studis
lost pursuit' it ontinus to thriv and thniqus.
756 Nturl hlklg

\/ithhis foundatiln now givn, Shppard gos ()n to disuss a numr ()f faitlr nd authority,
importnt issus in no-ontologial volutionary studis.Partiulr attntion on th ross filr our sil
is paid to polymorphism-th wys in whih natural sltion an ilt th t h r a t h t t r l t t t l i .
sam tim hold diffrnt forms in a population. Muh of this naturally draws xists nssrilyfom
on Shppard's own work, partiularly th studis lf srrails thet h had don ural thellogiallaim.
arlir in th dad' with fllow Darwinian A. J. ain, in th Oford oun- In th hristin rlrg
trysid. Shppard had spnt a yar in th loratory of ThodlsiusDlzhan- llogy. Howvr, sp
sky in Amria, and it is prhps aus of this that h is not ntirly always had n imptlrt
dismissiv of gnti drift as a ausal fator in volution, :rlthoughth strong thology, and famouslr
imprssion givn is that it is natural sltionthat ally ounts in dtrmin- istn of God, s wt
ing th gnti omposition of populations. By th 1975 dition Shppard is stnd his natur. Prott
vn lss kn on drift, although h dos admit tht it might nlattr in prltin susiiln ut hv gn
voltion. Anglinhurh prti
Othr topis that gt major disussion a th voluti<lnof dlminan, thority of th tholis
prottiv olortion, and mimiry. Hr w suly s somthing of a traditionally givn natu
nationalisti ias, for ths wr all topis of major intrstto Ronald Fishr volutionary thory'
inis Geticl Theor r>fNturl Sltitl(190) and wr mrrtiondlit. in sevrl rspts.on
tl in th lading Works of Amrian volutionists. Dozhansk,s Getis mirals. ny natur
d the rigi of Spis(third dition, 1951 ) has vry littl on n-rimiry,and mirals-r ki ngs lf
vn that is gorr y th tim of th d fatl fourth dition, Getis of th of th deity. \/atouJ
ullutior Pross (1970). l,wontin, a arxist onvrt' has somwhat so. L{rwvr,ny sin
ontmptuously rmarkd that th nglish intrst in uttrflis,th hif supposs tht th worl<
sour of information aout mimiry, rfltsth dilttantish,uppr-lassna. thory supp()rtsth ui
tur of nglish volutionary illogy (sLwontin |974). With good rson' In th fa of this o
A. J. ain (of vry huml origins) rtd vhmntly against this. But for h a v n t a k n .o n t h
thos intrstdin th ultural fators that influnsin,if th mphasis twn th ordr of na
is put only on partilrlar prolms, Shppard'svry attrativlywrittn littl prod aording ttl I
ook is highly suggstiv.It is vry muh writtn y irn nglishman who volutionry thory do
lovs his ountrysid and th dnizns thrlf. t h y w i l | r .l l p o i n
BIBLIOGRAPHY argud that mir:rls
a matt of maning. J
Dzhansky'T. l951. Gctisd thc rigi of Spels.3rd d.Nw York:
physial ris frlm th d
olumiaUnivrsityPrss.
Fishr' R. A. 1930. h Gtilblrf Nturl Sltil.Oxford: ()ford still livd. vllutionry
UnivrsityPrss. Anothr pla whr
Kuhn, T. |962. Tb Struturof SitifiRt,olutitls.
higo:Univrsitof om into ontat is th
hiagoPrss. tn of God is th isr
Lwontin,R. . 1974,Th }titssisrlf ullutillrhg'Nw York:
and Aushwitz. No Il-
olumiUnivrsityPrss.
Rs,' 1958.Rviwof NturlSltiod Hcredit.rcdit'12:522_52. happn. Darwinism s
Shpprd,P. . 1958.NtttrlSeltiotllld rcdit.l-ondot.t: uthinson. posits a r-riquitousstru
-i4.R. wys throughout th lrv
o n r s p o n s m i g h t
t l f v i l , n a m I y ' h tC r l
Natural thlogy that God ould hav r
Gld didd rrot tl lt h
Thology dals with our knowldg of th natur and istnof God. R- that popl would nv
vald thology is th ara that trats of <lurundrstandinglf God thrtlugh thn fir without pain.
 Ntrl Thcolog 7.57
d glsn to disuss numr of faith an{authoritv, this authority th hurh lr th Bil..That Jsus did
) n ys t u ( , | i l\).a r t i r r l ar t t n t i o n on th ross for otlr sins is a rvldthologial laim. Naturl thllogv is
lih ntural sltin atl at th th azlthat trats lf lur trndrstandingof Go{ througlr rason. hat God
ion.uh of this naturally draws xistsnssarilyfrlm his vry dfinititlrr(th lnologialargumnt) is a nat-
tudisof snails that h had ]on u r r I t h o l < l g i ll i m .
r A. J. in, in tlr oflrd oun- In tlr hristiarr rligion rraturalthology taks a ak sirtto rv:lldth-
)oratoryof ThodlsiusDozhan- ology. Howvr, spially givn th Grk influn on hristianit1,,it hs
of this tht h is not ntirly always lrad an importnt rol. atholis partiularly m:rk muh of r.ratural
in volrttion,although th strong theologv,nd fnlously Sint Tlrlms Aquins ] llv a sris of proofs of th
ln tht llyourrtsin dtrmin- xistnof God, as wll as disussiorlsof how w might rasonaly undr-
By th l975 ditiorr Shppard is stand his natur. Prltstantshav rgardd rason in ligion with mor
i t t h t i r m i g h t m i l t t ri n p r o t i n suspiion ut hav gnrlly allowd som pla for nturl tholog,v.Th
Anglirrnhurlr partiulrlv, sking middl r,vytwnth xtrm iru-
ar th volution f dominn, thority lf th atllolis and th trm ilialism of man1,Protstatrts'hs
w suly s somthing of a traditionrllygivn natural thlllgya signifiantpla.
lf major intrstto Ronld Fislrr voltrtionar1,thly,spi[IyDrrvitrisnl, irrrpingstln tru.tltlrolog1,
( | 9 )a n d w r m n t i o n d l i t . in svrlrspts.on ovious pint of tnsion ()urs()vrth xistnof
utionists.Dlzhansky,sGetics mirals. Many natural thologians hv rgtld that th isttl of
i 1 ) h a s v r , vl i t t l o r r m i m i r . n d mirls-rakings of th lrvs of natu-points t th ti.,.intvI-ltIon
to fouth ditiln,Gtis lf tbe of th lity.\Wtrould not hav rrurnd into win lrd God not don
arist onvrt' has somwhat so. Howvr, any sintifith<lry,in partiular n volutionary thtlry, p-
r intrstin uttflis,th hif suppossthat th rvlrld wllks aordirrgto lw, nd th sllss of sr'rha
ltsth dilttantislr,uppr.l:lssna- thor suports th uiquit,voI Iw irrrdhn tlr rrtlnxisrnof rrrirIs.
lwontin |974).With good rson, In th fa tlf tlris rjtion'th livr hs two optiolls' oth of whih
1 vhmntlyagainst this. But for hav -ntkn. on tlr l hand, it lrn argud that thr is a diftrn
influnsin,if th mplrasis twntlr lrdr of natur rrd th >rl of g. Nomlly th rvorld dos
rd's vry attrativlywrittrr littl prod aordirlg to law, lrut somtims God fls th nd to rtrvn.
t wittn y an nglishman who vlutionary thory dos not prov th:rt mirls i.rrimpssil, only that
of. thy will rar, point aptdy tlr livr' on th otlrr hnd, it an
ergud that rilsdo not nssrrily imply a akirrgof law. It is mor
mtt of maning. Jsus's rsurrtion' for instn, did not involv a
d.Nw Yok:
tlf Spis,3rd
physial ris fronl th dad tlt th flir.rg in th hrts of his disiplstl-rath
trl Slctl.oxitld: ()tord still livd. voir-rionary tl-r<lry
has norhing ro s:ryln this.
Anotlrr pla whr volutionary thory and natul thology sm to
'lutios. <lf
hiago:LJnivrsit1, om into ()ntatis th prolm of vil. A lssi r]umntagainst th is-
tntlf Gtld is th istnof physial and mol vils, srrhs arthquks
,lltillt.tr hgt'. Nw Ylrk:
and Atrshwitz. N ll-prvrful,all-good god luld hv llowd ths to
happn. Dawirrism smingly onfirs th istn of physial vil, for it
I rdit.Hrdit 12: 522_52.
r dit1,.[,tlndon
: posits triquitorrsstruggl fl xistl-l,invlvir-rgpain nd suffring al.
"..'.,'''.'.n:n,. o. ways tlrrlughor'rttlr livirrg r,vlrld.
on splt-ts right om fror an qually lassi ountr to th prolm
of vil, nrly, that God ann()tdo th impossil. Prhaps th only world
tlrat Gld ould hv rrd Ws on'tlrt inluds pain rrd sr-rffing'If
God didd ntlt tl lt urning aus pain, thn h would hv hd o llow
naturand xistnof Gol. R- that popl would nvr lan to far fir. It is ttr to hav pain with fir
ur undrstandingof God though than fir ,ithrlut pin. Prhps th t>r-rl1, way in whi God ould hav
7.'8 Ntrl hrlklg

ratd th living world was though a mhanism lik sltion,and hn It should mntio
h nnot lmd for its ad onsquns.(Kithr 2007 raks this t l Ii g n t D s i g l lT h t l r i s r
ritiism, that Darwinism xaratsth prolm of vil, and Rus 2001 lik natur of th living
rgus tht it may not quit suh a prllm for th hristian.) tht hn w must mai
Th majtlr pla whr volutionary thory, Drwinism in partiular, Although mmrs of th
oms into touh with natural thology is th qustilnof dsign (Rus 2003). idntifid with th h
From Plto on, it was rognizd that th living world is not random ut of thm. As might 1
shows omplity, an intgratd nd funtioning omplity, for xampl' sion of ..rationismlit.
th hand nd th y and th nos. How ould this ' givn that normally on should rrvr invlk
lind laws lad to randomnss? only thrlughth intrvntion of a dsignr, ar just not ndd in v
Orlsth rply. This rgumnt was pikd up y hristins, Saint Thomas is rquird. (S Dms
patiularly, and mad th asis of th most onvining of th argumntsfor th lading ontrovrsia
t h i s t n o f G o d : t h l i v i n g w o r l d i s a s i f d s i g n d ;t h i s i s a u s i t i s In onlusion, it must
dsignd,and th dsignris God. rethr tump taditin
Philosophrs' most partiularly David Hum, showd that in major r- just historil intrst.
spts' vn if thr is a dsignr,it anntlt muh lik th hristian god. th dsignlik natur of
Apart from anything ls, why stop at on dsignr?Howvr, Hum had to plaining. orovr, lik
admit that it dos sm s though thr is somtingat work. Using a vrsion hoil languag of d
of what is known as th argumnt to th st planati<ln'Hum agrdthat Ad1l ttil d Nturl
lind law dos not ld to dsignlik phnomna, and givn that th organi is produd as th funt
world is dsignlik,thr nrust slmethingor somon hind it all. With to srv that funtion, l
this pssiility, susquntnaturl thollgins,notaly Arhdaon William onsilus dsign. Th dt
Paley t th ginning of th nintnthntury' wnt on arguing for th - flr a rtin glll or fu
istnof God on th sis of th dsignlik natur of th living world. v o l v d w s f a s h i o n d y
Darwin's mhanism of natural sltion offrs an altrnativ to th god bautiful and historill
hypothsis.It is agrdthat th living world is dsignlik,ut it is argud that h y was aov all t}
this is ausof natural sltinrathr than Gld. No longr is th st - so1thingthat had ttl h
planation th hristian god. Drwin hs shown that th world is indd th y o u l d n o t h a v o m
produt of lind law, ut lind law of a spial kind, rramly,that whih pro- its, thos long-gon
dus daptiv funtioning by a pross of omptition and atrition. m i l l i o n y r sa g o a n d l a s
Not tlrat Darwin dos not hr disprtlv th istn of Gld. In fat, h flis (s figurs on pag
himslf was a livr in a dity whn h wrot th rigi of Species' tht it is ompl, with l
vn though latr in lif (mainly ausof th prolm of vil) h baman on ag 761).Why is th
agnosti. Rathr, Darwin shows that th dsignlik ntur of th world dos on th prolm rlizd
not nssitata god. In th words of today's popular sinwritr Rihard hristiaan Huygns had
Dawkins, ..Drwin mad it possil to an intlltually fulfilld athist'' if you Want to avoid h
(1986.6\. aus th diffrnt om
Dos this now man that th livrshould hav no thologial intrstin i n d i s ) .N a t u r a t t h t
th living world? any of today's hristins dny this strongly. vn though s o n a n d L v i - S t t i1 9 7 5
on an no longr hav a ntural thology-a proof of God-on an hav a ritis of univrsal D
thology of natur. On livs in God n faith (rvaldthology), ut and StphnJay Gould (
thn on's appriation of God's powr and majsty is {lshed<ruty tln's wronghadd sllution;
nountr with th living worid. Th uty of ros' th harm of a mr. Natural thology l
haffinh, and th urnof a fun shlw tht th ator was grat and prisly what should
loving dspit all th hatrd irnd vil in th world. a now-outmodd thlog

rnism lik sltion,and hen
s . ( K i t h r2 0 0 7 m a k s t h i s
' o l r ror f v i l , a n d R u s 2 0 0 1
for thhristian.)
cry, Dawinism in partiula'
qustionof dsign (Rus 2003).
ving wold is not random ut
'ning onrplity' for ampl,
rld this ' gil,r.rthat normlly
r th intvntionof a dsignr'
rp y hristians, Saint Thonrs
:onvinir-rg of th argumnts for
f d s i g n dt:h i s i s e u s i t i s
t m , s h o r v dt h a t i n m j < l - j u s t h i s t o r i a li n
l muh lik th hristin g<ld.
sigrrr?Howvr, Hum hd to
r t h i n gl t ' o r k . U s i n g v r s i o n
planation,um agrd tlrat
t n . n t Ig i v n t h t t h o r g n i
or somOnhind it all. \/ith
ls, nrltalyAhdaon Willinr
lry' Wllt on arguing for th x-
ratur <lfth living world.
ffrs altrnativ to th god
; dsignlik,ut it is argud that
n Gld. No longr is th st x-
wn rhat th wlrld is indd th
l k i n d , n a r l y t, h a t w h i h p r o -
)nlptirionnd attritiln.
:h-ist of God. In ft, h
,rot th rigi lf Speies,
rprolm of vil) h am an
ignlikenalturof th world dos
; populr sinwitr Rihd
n intlltullyftrlfilIdathist''
d h a r ' n o t h o I o g i iln t r s i n
dny this strongly.vn though
r poof tlf God-on an hav a
:r faith (valdthology), ut
majstvis f]shdout y ()n's
lty tlf :r r()s' th harnr <lf
l tlrtth rator WaS grar nd
orid.
Nturl Tholog 759
It should mntiond tht thr is t<lda,v an ative group' th so-alldIn-
tlligntDsign Thrlists,rvho argu thirt sltionannot plain th dsign-
lik natur of tlr living world, tlr:rtit shtlws ..irrdtri[omplxity''' nd
that hn W 1Llstmak planator rfrnto dsigning intllign.
Although mmrs of this group dny that this intllignmust nssarily
idntifidwith th hristian god, in 1atthis is th dirtion takn y most
of thm. As might lr ptd,th volrrtionaryommunity gardsthis vr-
sion of ..rationismlit'' With som dis]ailr,arguing on gnrillgrounds that
on slrould tlvrirrvokmirals in sinnd on spifigounds that thy
ar just not ndd in volutionar}.stlldies.Ntural sltionn do all tht
is rquird. (S Dmski and Rus 2-4for a olltion, ditd y two of
th lading ontrvrsialists,prsntingoth sids to th problm.)
In onlusion, it must strssdthat, vn though natural sltion dos
rathr trump traditional natual thology, th onntiln is of mor than
s t D
. a r w i n i a n s a g r f u l l y w i t h n a t u r a l t h o l o g i a n st h t
th dsignlik nttlof th lir,ing wlrld is tb cruial thing tht nds _
plaining. lrlvr, lik natural th<lllgians,Darrvinins us th mt:l-
phorial Ianguagf dsign unrsrvdly. Gorg Willirrrs, in his sminal
dpttio d Nturl Sletio,writs: ..!hnvrI livthat an fft
is produd as th funtion lf an adptation prftd y rratural sltion
to srv that funti<ln,I will us trms appropriat to humn atifi and
ltlsiousdsign. T dsignatiorrlf son-rtl-ring s th mns or mhanism
for a rtilin glal o furltiorl or pllros will implv thzrtrh mahirrr in-
vlvd ws fashiond y sltitlnflr h goal ttriutlt<lir'' (\966, 9).A
autiful and historially rdolnt mpl is furnishd r' th triloit y.
Th y was aov all th organ on whih th natural thllogianssizd as
somthing that h]to hav a dsignr-Paly (1802) mad muh of it. Th
y ould not hv om lut y pur han' Darwinians agr.Th trilo_
ites, thos long-gon n-lrineinvrtrats(thy ppard mor than 500
million yars ]oand lastd ovr 2'i0 rnillion yars) lr:rdompound ys lik
flis (sfigurs on pag 760).|f on uts through a lns vrtilly, on finds
that it is ompl' with two parts' spaatdy a urvd line (sth figur
on pg 761),Wh is this? \What dsign prinipl dos it sv?Rsarhrs
on th problm ralizd that irr th svntnthntury, Rn6 Dsartsand
Christian Huygns lrd solvd th prolm. It is th kind of lns you uild
if you want to voil hromati ar:rtion(n indistint fous oming -
aus th diffrrrtOmponnt olors lf whit light lrrvdiffnt rfrativ
indis).Natur t th sintistsy hundrds of millions of yars (lark-
s o n a n d L v i - S t t i\ 9 7 5 \ |
.Vfntworth
ritis of univrsl Darwinism, lik D,Ary Thompson (1917)
and StphnJay Gould (1997) gnrallv do not Want to laim that this is a
r,vronghadd solutitln;ut th Warn that on swall<rwdos tlot mak a sum-
tnr. Natrtral tholog1,lds ou to pt r'riqrritous adaptation, :rnd this is
prislywht slrould provn not assumdas part oi tlr unsnaggagof
a now-outmodlthologialmtaphor.
760 Ntrl Tholog

l-_-:=:_\

i------j

-/:\-

Triloit (rdoidsgrdilis,iddl (]ar.nbrian

spilrlnfronr Bohmi).
Top: Rn6 Dsrts(/ef
hromti arration.B
and Lvi-SttiI975.)

lLIoGRAPHY
lrkson' . N. K.' and R.
Dsartsand Huvgn
Dwkins,R. 1986.h Blt
1995. Riur()u
Dn.rski,\/.A., and M. R
amridg:ambrid
Gould, s' I. 1997. Th Dar
n o . 1 0 : 4 - 7 .
Kithr, P, 2007. Liuillg tu
Nw York: oxford Ur
.!7.
P a l y , [ 1 8 0 2 ]1 8 1 9 .N a
Rivington.
Rus, . 2001. Dr
d R I i g 'i, ' a m h r
23.Drid
Triloit e (Rdops strbrgi, Dvlttitrtriltlit frlrl l}tlhmia). MA: Harvrd Univ
 Nturl Theolog 76

Upr lns
unrt

Intralnsa
owl

i l i l l s l . t i r r l n i t l t l - tB o h m i ) . *l
lp:Rn Dsrts(left) and hristian Hr.rygns(right) shr.trrg
how to avoii
hromati arrtin. Blttotll:ross-stions.o1
triloItys. 1.3,,-,lrkson
n d L v i . S e t t |i 9 7 5 . \

lBLloGRAPl{Y

larkson' . N. K.' and R. Lvi-Stti. 197.5.Triloit ys

and th optis of
Dsartsirnd u1'gs.Ntur 254: 66-667.
Dwkins' R. 1986. b Btid Wthmkr. Nw Yrk: Norton.
1995. Riur out of de. Nw York: Bsi Books.
Dmski, W. A', and NI. Rus, ds' 2004. Debtig Dsig:
Drulilo DN.
arnridg:amridg Univsit1,Prss.
-I.h
Gou]d, S. J. 1997. Dru.iniln funia-.rt.liss. N Ylrk Ruituf Books 44,
no.10: 4_37.
Kithr, P ' 2007 ' Liuig tl,tithDrtai: uolution, Dsig,
d th Futur of Fitb.
N''v Yrk: )forj Urrivrsitv Prss.
.W..
Plv, [1s02] 1s19. Ntttr.l!Tht',l!t,g1, (,,lltd works, vol. 4). Lndon:
Rivington.
Rus, . 2001. Dru.lii B hristi? Th Rltilnsbip
bt Scie
d Rligitl. arri1g:Crridg Univrsity Prss.
20. Dill'i d Dsig: Dlls ullutilt Hu Purpos?
r i ] t l i t f t l m 1 3 t l h l - r r i )a. anrridg,
: Harvrd LlnivrsityPrss.
762 Th Nature of Sltion

Thompson' D , \/.19\7 . Grotuth cldFlrm.amridg:mridgUnivrsitr,

Prss. Nandrtals
!illiams' G. . |966. dpttltild Nturl Sltiott.Printon,NJ: Prir"rtl
Urrivrsity Prss. -.R. onound Ny_z-d
ard as Nrrrdrtha
of 18.56.Nan]ris th
The Nture of SIection (||iottSor) form ws adopt.dy t|
posr' Johinl Num
Th Ntr of Sltion y lliott Sor is an important ook that addrsssa
wid rang of issus in th philosoplry of iology. It is on of th fw ampls hang of ] 901, lvl-r
r r D t i s s l d o r f 'r l t l r vi
of philosophil work that hs ttratdth attrrtionof biologists (most notaly
rnst Myr). Th ook inluds disussions f th tutology prolm, urritsof and th skltorrws f
a av lld th Fld}
sltion' adaptationism,han and proaility in volutionary thory' th vo-
might hav lngdtt
lution of altruism, ausation, and foundational issus onrning sintifi -
planation in th iologial sins.Hr w an disuss only a fw highlights. in thir tratmIrt,and
W not ()v(,(|. su
On of th rost important distintions that Slnraks in this ook is
whih th origiel Ni
tht twn sltio of and sltilfor,In his sipl ampl w rnsidr
a toy that ontins small grn ds and larg whit ads.l/hntlr ttly is C r o i . F r l t r t l' t I v 'l
fat, anothr spitnn
shakn' th small ads pass through a st of sm:rll hols and ar thry s-
of Giraltr irl t848,r
ltd. Ths ads hav two proprtis: thy ar th small and grn. But
ollg of Surglnsirl
thy ar sltdfor th proprty of ing small; th fat that grn adsar
Thos individuals hav.
sltdis mrly aidntal. So w hav sltionof small grn ads' ut
sltion for on| th propty of ing small. Nndrtal sklttrlI
Sor brings this distirrtioI-rto ar upon th prolnr of tlr r.rnitsof sl- t th ommun tlf S;l
logil OI1ttwith st
tion. Using a disussion in G. C. williams's dpttio d Nturl Sltio
(1'966I,h distinguishsrwnindividual and grup_lvladaptation in th nizd to b of th sr
.W of L Moustir' dug in
following way. suppos that a hrd of dr onsists of fast individuals who
ar al to sapprdators. As a rsult, tw things happn. Th first is that in. with th Nnlrtls;
dividual dr surviv and rprodu. Th sond is that th hrd survivs and dioativ lmts to t
rprodus.But austh survival of th hrd is mly a y-produtof th not possil until aft
individuals'survival, this is a as in whih thr js slctioof fast hrds.his M t t t l s t i nw l s r ' l t |
modrn-irpplrringhun
sltion of th fast hrd is in ontrast to te seltio for fast individuzrl dr.
In this way Sor argus that th dat aout goup sltionism hings upon lir than th Nadrt
in Java in 1891 and 1
th qustion whthr thr an e selcti<l/or groups. This disussion suggsts
that th prolm of th units of sltion is an mpirial qustion, whih is not Homrl) ertus,All of t
to sttld a priori. v o lu t i o n a r v p s p r
A thm that runs through this ook, as wll as th rst of Sor'swork, is th Strassurg itlll]t()m
with th tul mhil
that philosophial analysis an larify iologil onpts in sulr a wy s to
inform mpiial invstigation.On his viw thr r sustantivarrd dis- straightftlrrv:r rd pitur
j u s t a o u t t h l l l s tt i m
tintly philosophial qustions that onfront th iologial sins' ut thos
philosophial issusar dirtly upon how iologistsgo aout thir work. tionary prsptiv.
I n t h s u m m r < l f1 9
lBLloGRAPHY prists wr prrsuing
yr, . 2004. 80 yarsof whingtlrvolutionary sll,Y.
Si05 4647, h a p l l _ a u _ S l r i n ti ns
Sor,. |984. Th Ntl' lf Sltkl:uoltitlr1'.[.hor
i Phillslphil unovrd :l onr1.llt
Fot,ls.amridg'MA: N,{IPrss. atd thir 1islvry t.
l0illiams,
G. ' 1966, dpttitld Nturl Sltitltt.Printon,N.}:Prrntln Fran in Paris. A6 B
UnivrsityPrss.
rommndd tht th l
 |',[edrtlls 7 6

amridg Univrsity
Lmridg:
Nandrtals
',!.Printon,NJ: Printorr
-.R. Pr<lntlundNa'-az-dr-tahland rrlwsplld Nandrtill, th nm first ap-
pard as Nndrthal whn th fa-rlus flssilr,vasdisovrdin th summr
of 1856. Nandr is th Grk rndition of Numnn (nw man). Th Grk
)r) form was adoptd th family of svntnth.ntury Gmn hymn om-
posr,Joahim Nut-nann.Thal was Gman for ..vlly'' fo th splling
mportantook that addrsss ll
lrangof 1901, w,ht.t it am Tal. Nirrrdrtalthn was N',vnan'svally
;y. It is on of th fw xampls nar Diissllorf, now in tlr indr-rstrilrll-rartof tlr rvstrn art tlf Grmanl,.
ntion of iologists(most notly
and th sklton rvas found y marl quarry workrs rvlro w laring ot
th tautology prolm, units Of
a av alld th F-ldhofrgrotto. Tlr workmn thouglrt that th sklton
. in volutionarythory' th vo-
might hav longd to a prhistori tvar, so thy wr not vry arful
l issusonrning sintifi -
in thir tratmnt'nd th mor fragil nd smallr parts of th fa nd tth
n disussonly a fw highlights.
Wr not rovrl.Susquntlyothr prsntativsof th plpulation to
at So maks in this ook is
whih th original Nirndrtal individul blongd wr fllunl in Blgium,
lris simplampl w lnsiir
lWhn th toy is roetia, Fratt,Italy, and Spirr,s rr",ll as frthr irstit-tlq nd Isral.In
; whit ads. ft, anothr s;litnnhad n disor,rdelow th llorth fa of th Rok
small hols and ar thrby s_
. r b o t h s m a l l a n d g r n .B u t of Giraltar in 1848, ut it rmaind r.rnstudid in th olltiorrsof th Royal
ollg lf Surgorrsin London until th first dadof th twntith ntury.
rll; th fat that grn ads ar
Thos individuals hav n alld Nr-rdrtals vr sin.
tin o/ srrrallgrn ads, ut
Nandrtal skltl matrial ws flutr1 in th Blgin provin of Namur
a t t o m m u n t l f S p , v( p r o n o u n dS p )i r r 1 8 8 6 i n s t a t i f i da r h a o -
r prolmof th units of sl_
logial olltt with stln tools :rnd nimal ons. Tlr tools Wr rOg-
dpttk d Ntrl Seltio
nizd to of th sm kind s thos found at th sourlrrvstrnFrnh sit
rd group_lvladaptation in th
of L Moustir, lug in 186. In honlr lf tht typ sit, tlr tools assoiatd
: onsistsof fast individuals who
with th Nandrtals ar lways rfrrd to as Moustrin. Th us of ra-
rings happn.Th first is tht in-
diotiv lmnts to stalish asoIut dating of prhisroi matrial was
lnd is th:rtth hrd survivs and
,rd is mrly a y-podut of th not possil until irftr World Wr II, although it was a|izd that th
o t l s t r i a n w a s l l ] tr n t h U p p r P l o l i t h i t r a d i t i t l n sa s s l i t dw i t h
:re ls sletilof fast,hrds. This
m o d r n - a p p r i n gh u m a n s s u h 2 r s( i r l _ M a g n o n ,f o t t r - ridn 1 8 6 8 . M u h r -
;ltiottfor fast individu:rl dr.
lir than th Nrrdrtalswas th ml-rrrialfound y a Drh army physiiarr
: group sltionismhings rrpon
in lava in 1891 nd 1892 that h allrdPiththrlptts(w nw all it
groups' This disussion Suggsts
Homo) rtus.All tlf this matrial Ws mtiulously studid and put into n
mpirialqustion'whih is not
volutionary prsptivduring th first dad lf th twlltith ntury y
th Strassug irnlrtlt-l-tist Gustav Sl-rrvl(1844_1916). H did not dl
ll as th rst of Sor'swork, is
witlr tlr atltirl tlllrnisnrsrvhr,lr:rrrg \1.aspodtl1r-rtprsntd.r
al onptsin suh a way as to
.thr a sustantivand dis_ straightflrwalpitur of ontirruity and hang tlrroLrghtim. This w:rs
just aout th lst tim that th hurrrilnfossil rord was rrtdin an volu-
hiologial sins,ut tlros
tionaryprsptiv.
riologistsgo aout thir work.
In th sr.rmrrof ] 908 two rothrs who had rntly n ordaind as
prists wr purstting rhallogial :rvation nar th viilag of La
hapll-au-Silltsill orrz in sorttlrrvstrnFrn. Or-rAugrrst 3 th
narvsry'Sitt5:46_47.
,rTherlri Phiklsryhil unlvr] tltrrp[t end wll-prsrvdNndrt:rltrtlri:rl.l-rytllrrrnuni-
atd thir disovry to A6 rrri Bruii, a profsslr t th ollg l
, I t i , ,Pl t,i n lN
n J. : P r i n r o n Fran in Paris. Ab6 Bruil was alrady wll known in :rrhallogy,and h
-Z..
rommnddth:rtth othrs out thir sklton into th hnds of his frind
764 |']drtls

a r l l i n B o u l ( 1 8 6 | - 1 9 4 2 I , a p a l r o n t o l o g i s tw h o w a s p r o f s s o ra t t h
Mus6um d'Histoir Naturll in Pris. Boul's monograph dsriing La
hapll-u-Saintsal ()ut in thr instllmntsin 191 1, 7912, and 191
and st th ton for th study of human ..volution''for th rmaindr of th
':::!:r,
:

twntith ntury. Thr was nothing vn faintly volutionary aout it. Gus-
tav Shwal wr()t ln of th rviws f Bor"rl'smonograph, noting th
anatomial rrors Boul had mad in his ronstrution ut, uriously
nough, pting th patntly ntivllutilnaryast of th intrprtation.
Th vry nt yar World .JrI rok out' and on of th asualtisof that
onflit Was th volutioner orinttion in th Grman tratmntof th hu-
man fossil rord.
Bfor th wr lth th Grman and th nglish litrtur tratd Nan-
drtals as n arlir stag in human volution that had om transformd
through tim into nttlmially mrdrn flrm. Undr Boul's influn th
Nndrtls wr privd in ariatur and rgardd as hving n r-
p l d y a n i n v a s i o n o f m o d r n - a p p a r i n gh u m a n s w h o a m f r o m . . t h
ast.'' Thr is no arhaologial or anatomial support for this intrprta-
tion, :rlthough it is th viw now hld y most studnts of humn ..volu-
tion.'' Various stimatsof Nandrtal statur show mals ranging from 764
t o 1 7 5 n t i m t r s( 5 , 5 , 't o . 5 . 9 , ' ) a n d f m a l sr a n g i r r gf r < l m1 5 2 t o 1 5 8 n -
t i m t r s( 5 ' 0 ' ' t o 5 ' l ' ' ) . M a l o d y w i g h t h a s n s t i m a t dt o r u n f r o m 9 0
t o 1 0 0 k i l o g r a m s ( 1 9 8 t o 2 2 0 p o u n d s ) . r n i a l a p a i t yw a s l a r g rt h a n t h This drawing showsho
modrn avrag ut prftly in proportiorr to thir gratr ody wight. th modrn stty a r
larly th Nandrtals wr haratrizdy a dgrof skltalrOustnss arhitturwhil th
.With unhangd.Th Nnd
and musularity wll yond th mor rnt human avrag. intll.
lf approimatly70,0
tual and linguisti apailitis proaly at modrn lvls' th Nndrtls
Palolithi ml from tl
qulify as an arhai form of Homo strpis.
yars ago' nd rn
Th onvrsion of Nandrtal into modrn human form was aomplishd
y rdution in thos manifstatitlnsof roustnssas a rsult of thnolog-
ial innovations that allowd thm tl gin susistnwith lss pnditur |LIoGRAPHY
of ffort. Analysis of tra lmntsshows that thir intak of animal protin
Brir, . L. 1995. I]il
was approimatly th sam s that whih hartrizsth wolf. Th us of
in th mrgnof
traps and snars' howvr, gratly rdud th anrount of ffort ndd to 7t1-72t.
ring food hom. Most mutations rdu th trait that thy ontrll, and 1 9 9 5 .T h e S t g
whn sltion that maintains r<lustnssis rlad, its manifstationswill P r n t i H | i .
show a rdtltion. Th oking lf food will rdu th intnsity of sltion 2.ultio
Prss.
tht maintains tooth siz. During th last glaiation Nandrtals wr using
Morin, . 2004. Lat Pl
arth ovns to thaw food that froze, and a rdutiln in tloth siz an volutionary aproa
shlwn fom 130,000 yars ago in a straight lin of 7o evr2,000 yars un. lf ihign, Ann A
til th nd of th Plistlnartlund 10,000 yrs ago. Trir.rkaus,., nd P. Shi
Prliminary mtDNA lmparislnsshow that th Nandrtals diffr mor clkid. Nr,vYol
from th ndition in living humilns thn th avrg diffrn twn
plpulations of th living. Th distintion twn Nandrtals and living
asatoshi (b
humans, howvr, is not as grat as tlr mtDNA diffrnstwn plpu-
ltions of himpanzs Or vn th diffrnswithin populations of him- N4astoshi Ni has
panZs. sity sin 1990.H o
 Nel 765

,t who was a profssor at th

l's monograph dsriing La
n t si n | 9 | 1 , 1 9 7 2 , n d 1 9 1 3
tion'' for th rmaindr of th
tly vllutionaryaout it. Gus-
ul's monograph' noting th
ronstrutionut, uriously
lry a5tof th intrprtation.
.d on of th asultisof that
Grmn tratmntof th hu-

nglish litrturtratd Nan-

that hd om transformd
r. Undr Btrul'sinflun th
J rgardd as having n r-
..th
humans who am from
al support for this intrprta.
,.volu-
lst studntslf human
show mals ranging from 1'64
l r a n g i n gf r o m 1 5 2 t o 1 5 8 n -
bnstimatdto run fom 90
al apaity was largr than th Tis drawing shows how Nandrtairaniofaialfom was onvrtdintl
to thir gratr ody wight. th mdrn staty a rdutionof th dntitionnd its supportingfaial
. a dgrof skltalrobustnss arhitturwhil th ontoursof th rain asrmaindf{tivly
unhangd.Th Nndrtalis th L Frrassi1 mal of southwstrnFran,
human avrag.\/ithintll.
of approirrratly70,000 yarsag' supimplsd ovr an arly Uppr
nodrn lvls, th Nandrtals Palolithimal from th zh Rpuli,Pidmosti ' of 25,000 to 29,000
yrsago' and a rntFirroIslandsmal.
human form was aomplishd
Jstnssas a rsult of thnolog-
rsistn with lss pnditur BIBLIOGRAPHY
rt thir intak of animal protin Br;r,. L, 1995a. Bio-ultural intrtitrnand th mhirnismof mosai volution
Laratrizs th wolf. Th us of in th mrgnof
..modrn'' mrphologv. mri Athologist 97, na.4:
:h amount of ffrt ndd to 7 1 1-721 .
h trit that thy ontrol, and 199.'. h Stgcsof Hum uolutirl..5thd. nglwlod liffs, NJ:
PntiHli.
rlaxd, its manifstations will
200. uolutirm itt Athropoklgil Vi' Wlnut rk, A: Altaira
duth intnsity of seltion
Prss.
r i t i o nN n d r t l sw r u s i n g population intrtion in wstrn urop: An
orin, . 2004' l,t Plist.lrr
rdution in tooth siz n volutionry approah to th rigins of modrn humans. PhD diss.,Univrsity
i n o f 1 % v r y2 , 0 0 0 y a r s u n - of ihign, Ann Arbor.
Trinkus, ., and P. Shiprnan. 1992. h Ndrtls: hgig the Img of
/arsag().
ktd.Nw Yok:AlfrdA. Knorf.
rat th Nandrtals diffr mor
th avragdiffrn twn
)twnNandrtals and living Ni, asatoshi (b. 1931)
)NA diffrnstwn popu-
:s within plpulations of him- Masatoshi Ni hs n van Pugh Profssor at Pnnsylvani Stat Univr-
sity sin 1990. H otirindhis PhD from Kyottl Univrsity in 1959. H was
766 Nei

ssistant profssor, Ky()to Univrsity, 19.'8_1962; gntiist,Nationl Insti-

tut of Radiologil Sins,hia' 1962-1969; assoiatto full profssor,
Brown Univrsity, 1969_IL)72;and profssor,Univrsity of Tas' HoLlston'
7972-t990.
In volution thr is a grat fflrt tl infr th volutilnay rltilnships
among organisms.his ranh of iology is ormonly known as systmatis.
In th 1960s volutionistsstartd to us iohnrialand molulr diltil, mainly
protin letrophortidata' to study systmatis;ltrophorsisis a thniqu
usd to masurth moility of a protin on a gl. |n |972 Ni dvlopd
mthod for stimating th gnti distans twn populations from l-
trophorti data. This mthod soon am th standard mthod in tlr study of ohta, Tomoko (.
volutionaryrlatilnshipsamong populationsrlr loslylatdspis,and th Tomoko Ohta has n
masur otaind y us of th mthod is known as Ni's gnti distan.Th tion for undrstandingr
rason for its popularity is that th mthod givs an stimatof th ntrmrof dp undrstanding of
amino aid diffrns twn two protins and thr-rsth dgr of protin- spis with reativ ml
squndivgntwn th two populirtions frorn wlrih th two protins fundamntal qustions:
wr takn. Thrfor, this nrasrrris as t<lundrstand.Tli mthtld alstl hs A studnt of nihr
ttr statistial proprtis thn prvious mthods. I 1979 Ni dvlopd a turnd to Japan aftr o
mthod fo stimating th numr of nulotid sustitutions pr sit twn mous ollaboration Witl
two DNA squnsfrom rstitiln-nzymdilta. l.risWi.lsanothr milstln Kimur'' in this volum)
in th study of gntidistan.l-atr dvlopdsvralllntmthodsfor of nutral allls(i..,all
stimating th numr of nulotid sustitutionspr sit twntwo DNA s- rirs). In 1971 seand Ki
quns from DNA-squn dta. In \987 h anl N. Saitou dvlopd th th nutrl thory ould
nighor-joining mthld for ronstruting vtlluti<lnar1, (plrylognti)trs. in natural populations, a
This has om on of th most popr.rlarmthods in phylogrrtironstru- ssays prsntd a ompl
tion sin th lat 1980s. Now lmost vry ssayon phylogntistudy ussthis potntial powr as an
mthod.In summary,Ni has n prominnt ladrin th study of molular work providd a thrt
systmatis(phylogntis)sin th 1970s. is rnthtldshav hlpd larify th protin variation to stl
volutionary rltionships among orgnisrs and gns. In addition, h hs l t i o n s i z ( a m a s u r o i
mad many sminal ontriutilrrsto th study of molular volution, shs gnration) to th us of
th nutral thory of molularvolution (dvisdy Motoo Kimura and asd In xploring th nutr
on th laim that muh mtllular hang is rrathth fr of ntural sl- out that ntural plpula
tion nd thus gnti drift is a major fatlr),as wll s to our undrstandingof prditd. o aunt f
th volution of major histoompatiility gns. F.or his lntrir'rtions,
Ni ws variants would slight
ltdto th U.S. Natilnl Aadmv <lfSinsln 1997. hypothsis ld to diffr
alignd ttr with os
|BLIoGRAPHY Ohta also did pion
Kimur,. 1983.Tb Nutrlhlrf lllulruoltitl,amridg: hmoglobins. Ths fam
ambridgUnivrsitPrss. ar major parts of th g
Ni, M. 1972.Gntidistantrvt-t
p<lpulirtilns.
mrittNturlist|6: ilis in volution and d
28_292. gnti drift would pro
Ni, ., nd.W.-.I,i. 1979.MthIltilro]l grrti
ftlr stul1,irrg vlrilrtitltrin
among opis within s
trmsof rstritiorr Prlcdigs
ndonulirss. lfth Ntitlldm-li
and among populations
'"'.*;.ill.Y;;.]J*IT;::*l;;.',:;;x'':*"!;h",il,i|:)ni.o^!,ii_o
SicesUSA 76: 5269_527 .
r of th Amrian A
forign assoiat of th
-w.-.L. nizd y th mpror of
 962; gntiist,Ntionl Insti-
)69; assoiatto fu11profssor'
, Univrsitytlf Tas, Houston'
th volutionaryrlaionships
l t n n t t l n lk1l l t l w n e s s y s t m a t i s .
nialand nroluladt, rninly
s; ltroph(lrsisis thniqu
gl. ln l972 Ni dviopd a
)twn popultitlns from l-
standrdmthod in th stdyof
lr loslyrlatdspis,and th
vn as Ni's gntilistn.Th
rsn stimi1tof th numr of
and thr-rsth dgr of prltin-
l n s f r t ' t l tr rh i h h t w o p r r l t i n s
tndrstnd. Th mthod also hs
hlds.In l979 Ni dviopd
l sustitutionspr sit twn
lata.This was anothr milston
pd svralllntlnth()dsfor
s pr sit t,ntwo DNA s-
r anJN. Saitou dvlopd th
volution;rry(phylogneti)trs.
hods rn phylogntironstru-
ly n phyllgntistudy uss this
t ladri th study of mrlular
s mthodshav hlpd larify th
and gns.In additi<ln,h hs
, of molularvlutitln,suh as
sd y Mtoo Kirnurir and asd
nathth for o{ ntur:rl sl-
s rvll as to our undrstanding of
s. For his lntriutitlns,
: si n 1 9 9 7 .
lr F'ullut itt. ar bridg:
lns.mri NturIist \06:.
. . lf o r l u d 1 i n gg' . n t iv r r i : l l l l il t t
gs ,lf th Ntill dem of
lg mthod:A n, Inthd for
\irltlgl d F't,llltirl4: 46-42"5.
Ohta, Tomoko (b. 1933)
Tomoko oht has n a lading figur in dvloping a thortial founda.
tion for undrstandingmolular volution. In hr work sh has omind a
dp undrstanding of pattrns of molular variation within and among
spiswith rtiv mathmatil modling to approh som of th most
fundamntal qustions about molular volution.
A studnt of Knihi Kojima at North aolina Stat Univrsity, ohta r-
turnd to Japan aftr omplting hr dotoral dg io t967 and gan a fa-
..otoo
mous ollaoration with Motoo Kimura (s th alphtial ntry
Kimura'' in this volum). Thir arly ssaysdvlopd modls for th dynamis
of nutral allls (i..,allls that do not onfr diffrnsin fitnsson thir ar-
rirs).In |971 seand Kimura pulishd vo landmark ssaysthat showd how
th nutral thory ould onnt mutation rats,lvls of protin polymorphism
in natural popultions, and long-trm rats of amino aid sustitutions.Ths
ssaysprsntda omplt pitur of th nutral thory and dmonstrtd its
potntial powr as an planation for smingly disparat phnomna. This
work providd a thortial foundation for svral lins of sarh'from using
protin variation to stimat popultion paramtrs suh s th fftivpopu-
lation siz (a msur of how many individuals atually ontriut to th nxt
gnration)to th us of molular variation in phylognti studis.
In ploring th nutral thory furthr, Ohta was among th first to point
out that natural populations harord mor rar llls than nutral thory
prditd. To aount for this pattrn' sh postulatd tht som molular
Ni was variants would slightly dltriousand othrs modratly d|trious.Hr
hypothsis ld to diffrnt statistil distriutions of fitnss ffts that wr
alignd ttrwith osrvddata on polymorphism and rats of volution.
ohta also did pionring work on volution in multign familis lik th
hmogloins. Ths familis ar sts of tandmly rptd opis of gns and
ar major parts of th gnom. ohta rognizdth importan of ths fam-
ilis in volution and dvlopd most f th arly thory that prditd how
gntidrift would promot divrgnin th DNA squnsof ths gns
among opis within a singl individual, among individuals in a population,
and among populations. In 1984 Tomoko Ohta was ltd a forign mm-
r of th Amrian Aadmy of Arts nd Sins.In 2002 sh was ltda
forign assoiat of th U.S. National Admy of Sinsand ws rog.
-w-H.L. nizd y th mpror of .|:rpanas a Prson of ultural rit.
767
768 On Crowth and Form

|BLloGRAPHY
Kimura, ., and T. oht. |971. TheortilAsptsof PopultioGtis.
Printon,NJ: PrintonUnivrsityPrss.
ohta, T. t98. uolutitld Vritilof ltltigmilis.Brlin:Springr-
Vrlag. -J.T.

Groth nd Form (D'Ary Wntwrth Thompson)

Groth d Form' first pulished in 1.917,is still in print as an arvi-
atd vrsion in paprak' with an introdution y StphnJay Gould. This
in itslf marks th ook as somthing rmarkal.It is hard to think of lthr
iologial txts of a similar vintag that hav a omparal nduring sin-
U
tifi rlvani apart from volums rprintd primarily for thir historial in- str

trst' on thinks of Darwin's works and not muh ls.Th ok is and has
always bn sn as th distintly idiosynrati opus of a highly nti i.
.sntworth
ologist, D'Ary Thompson (profssor of zoology at St. Andrws
Univrsity in Sotland, 1917_1948),who is fftivlyrmmrdsolly on
th strngth of this ook. Why has it survivd?
Th ook prsnts th as for an aorrnt of iologial struturs asd
on mhanial and physial prinipls. It aspirsspiallyto mathmatial
mthodology; hmistry hardly fatursat all. Using an stonishing srisof Diodon
ampls aross th whole rang of phylogny and of salsof magnitud,it
shows how various typs of iologial ..form'' an potntilly plaind
Left: out|ine of th poru
y various typs of physial fors. Topis ovrd inlud saling, ratios o{ ( rt h glriscus mll).Nt
surfa to volum, growth urvs' hromosom pattrns' ll strutur and any knowldg of th ma
ll division (xplaind in trms of surfatnsion),multillula arrays (asin rpliathis findingsnd
honyoms), shlls, horns, flowr pattrns (ana|zd in trms of Fionai fom th fish form on th
sris),zra striping, and shaping of ggs and on strutur (plaindas 1948,1064.1
adaptations to imposd mhanial strsss).Th losing haptr is undout-
dly th st known and th stion most dirtly rlatd to volution. It
provd to b ..wrong'' (
dals with Thompson's ..transform'' mthod-a dpition of diffrns in
sis was saltationist.Th
morphology of rlatd spis (.g.,fish) or whol strutllrs'suh as skulls,
s o n ' s r l ' m i s s i o nw a s -
in th form of oordinatd gomtri dformations (familiar nowadays in th
stil1 prvalnt-to show
guis of omputrizd morphing) (s figur).Thr natur and vlu of this
..mthod'' is disussd l i n s o f t h a l r d ys i
lswhr(Hordr 2005); it is atually no mor than a
physis.
dsriptiv dvi and is not asd on' or an indiation of, any dvlopmn-
How is it possil th
tal or volutionary mhanisms as suh. Among prsnt.day apprahs to
ago? On fator might r
volution, th losst omparisons ar morphomtris and spia1lyallom.
with som instantly stri
try' whih, through J. S. Huly' was th most dirt and pliit dvlop-
up Thompson's mssag
mnt that aros out of Thompson's ook.
Proaly mor signifia
Th survival of this arhtypially lassi book is ll th mor puzzling -
ologists wr inspird
aus it is distintly datd in oth styl-it is writtn in sholasti and ven
Goldshmidt, Julian Hu
litrary mod-and ontnt. It dats from a tim whn gntis was in its
dan, and tr Mdaw
infany, Darwinian volution Was ativly ing qustiond in favr of La-
a symol and idal of l
markism, and thr wr many gaps in undrstandingof asi ll iology.
prolms of iology. T
or surprising still, Thompson adoptd positions on thsthmsthat hav
ious swp and rang
 on Gowth and Fom 769

ts of Popultio]tttis.

e Filis. Brlin:Springr-
-J.'

worth Thompson)
,' is still in print as an abrvi-
n StphnJay Gould' This
l. It is hard to think of othr
a o n r p a r a l n d u r i n g s i n -
lrimarily for thir lristorial in-
nuh ls.Th ook is and has
opus of a highly Irtii-
ssor of zoolgy at St. Andrws
.ftivlyrmmrdsolly on
?
of iologial strr"rturs asd
:sspiallyto a mathmatil
. Using arr astonislringsis of Diodon
y and of salsof magnitud, it
t'' an potntially plind
Left: out'|tneof th porupin fish (Diodo) , Rigf: outlirrof sunfish
tvrdinlud saling, ratios of (rthgorissmol). Not tht Thompson was working intuitivly, without
m pattrns' ll strutur and an1'krrorvldgof tl-rmtlrmatisinvolvd. Today' with omputrs, w n
sion),multillulararrays (as in rpliathis findigsand hav a full undrstandingof the movsneddto go
ana|zedin trms of Fionai frorn th fislr form on th lft t th fish {orm on t right. (FrlmThompson
rd on strutur (eplaind as 1948,1064.)
h losinghptr is undout-
lirtly rlatd to volution. It
provd to ..wrong'' (sHordr 2005). is position rgarding phylogn-
- a d p i t i t l no f d i f r n si n
sis ws slttionist.Th ook is in vry wy a produt of its tim. Thomp-
vhol strutrs,suh as skulls,
son's ral mission Was-as a ountr to th vauous vitalisti xplanations
rtions(familiarnow:rdaysin th
still prvalnt-to show that iologial phrromnw plibl along th
). Th natur and valu of this lins f th alrady sintifiallyhighly dvloped disiplin and mthods of
l5);it is atually no mor than
physis.
indizrtionf, any dvlopmn-
ow is it possil that this ook is as influntial now as it was 90 years
ong prsnt-dayapproahs to
ago? On ftor n-rightwll th illustrations.Th ook is rihly illustratd
omtrisand spially-rllom-
witlr som instantly striking nd mmoral piturs tht in thmslvssum
ost dit and pliit dvlop.
up Thompson's mSsag;th an ' and hirv oftn n, radily rprodud.
Praly mor signifiant is th way in whih a stady sussionf ky i-
ook is all th mor puzzling -
ologists wr inspird y th work. Aross svral gnrations (Rihard
; writtn in sholsti and vn
Goldshrrridt,JLrlian Huxly, vlyn Huthinson, Alan Turing, J. B. S. Hal-
tim whn ]ntiswas in its
dan, and Ptr dwar to StphnGould and John Bonn) th ook was
ing qustiondin fvor of L-
a symol and idal of a rigorous sintifi approah to many of th major
:rstandingof asi ll ilogy.
prolms of iolgy. Th ook appald all th mor aus of its auda-
itions or-rths thnrs that hve
ious swp and rarrg. Tlris list inluds th nms of som of th most
on th origin of Spis

influntil biologists of th twntith rrtury,many of rvhor mad Thomp. naturl s|tilnpr<lvi

son widly kr-rownthrough thir popular writings. Th ook nright appar to ths asonfir th l
of most dirt rlvanto dvloprnntaliologists,ut thy hav rarly ular agunrnts on t
..holists''(morphologistsnd spially
takn it up; it has appald mainly to as a osilie of idtt,
palontologists)or on mthodologial grounds (iorrrathmatiins). In an sin Willinr .Whr'v
ag domintd y molular iology, gntis' and p<lpulationtlrinking- its inrplt:lrrand plrr
whn nrophology is lss nd lss tauglrt in univsitis-this btrok stands to- Thr wr majtl h
day as a rmindr of ultimat volutinary prlms and planatory D:rrwin pulishl,vr1
hallngs posd y whol organisms. Th afftion with whih it has n th truth of volutior..
hld through th gnratiorrsis inditd y th fat that D,Ary $7rrtworth strjtdtl stringnts
Thompson is th ddiatof Hul's Prollntstf Rltiu Gruth ||9z) ous. Tlr 6rst was tl-rtI
and Gould's ntoge d Phlog1'fl977). today ll,luld ll a tht
o v i r r sr h n t n n l h n
|LIoGRAPHY ond ojtion,sd
Gor'rld'S. I.1'977.tlgl'dPhlklgr'.rnridg. lv,1:
BlknpPrssof hting po\\'rtlf th sr
rvarJUnivrsityPrss. mhirtlistns sltr
Hordr, T. J.2005. haptr64.II. Gron-Guinnss, d.,Ldmrk Writigs As w now know, th.
i Wstrthmtis:s Studies,1640-1940, 823_82.Amstrdm: ndl's thory lf h
lsvir.
ountrl.h srlndr
Huly,I. s. l932. Problmsof Reltiurrlu,th.London:lVlthun.
Klir' . F. 2002. kig Sso| Lii: plining BitllogilDulrlptnttuith f t o f r d i o a t i v d
odls' etphors,d his.amridg'A: Hrvard UnivrsityPrss. Was not thn kn<lwn.]
Thompson,D. \. [1917l 1948. Groth d Form.2nd d. Carnridg: lms, nd to ountr th
amritgUnivrsityPrss. tay mhnisms ttl sl
1992. Groluthd Form.Aridgdvrsion.Clrrrrhridg: ambrilg
-.J,H' inlrritn of aqtrird,
UnivrsitPrss.
Unfltunatly,th rr
gant tndd ssayrr
that tod1.'sradrsgn
the rigin of Species (Charls Darwin)
hrls Drwin's mjor wrk, th ()rigilt rlf Speisl s tlf NturI BIBLIOGRAPHY
Sletio, or th reserutio of Futred Rs itt th Strggl for Lif' was Drwin' . l859' () thc
pulishd in Novmr 1859. Thr wr 1'250 opis, and it was sold out P r srut itltt llf F ullt'
on th first day (mor prisly,th pulishr sold ll f th opis to ook- urit\..
sllrs at his innual sal). Darwin startd immdirly to prpa a sond I e S e .f 1 1 n
, 'r 1 t 1
dition, and in ll thr wr six ditions, of whih tlr last appadin 7872. P k h a m, d . P h i l d
Ril.r:rrds. ll. .|',nd . Rt
Th work is dividd into two parts. In th first prt Darwin prsntd his
of Sp:is.'' mid
mhanism of natural sltion and offerd rasons to think it plausil.H
did this first through analogy with artifiial slti<rn' pointing out th su-
sssof niml and plant rdrs' tlrn Wnt on to show (in an argu-
mnt modld on tht of Thomas Rort Malthus) how a stuggl for
ontogn nd Ph
istn is a onsqun of th growth of populations to a lvl gratr In tl3i1,d h1,|
than vailal supplis of food and spa an support, and thn how, givn growth o th irdividu
naturally ourring variation' th struggl lads to diftrntialsurvir,l and lristor1l sttrlvtlf rll
prodution, whih Drwinalld tual sltion. twn stagsin dv|op
In th sond part of te rigiz, Darwin applid his mhanism aross th Th anlogy twl
sptrum of iologial aras of inquiry_instint, biogographi distriutions, ing'' was familiilr to Al
palontologyt anatomy' lssifition, nrryolgy, and othrs-showing how this il r,rs
rslrpdi

many of whoin mad Thomp-
rgs.Th ook might appar to
liologists,ut thy hav rarly
(morphologistsand spially
Js (biomathratiians).In an
:s' nd ppulin rhinking-
ivrsitis-this ook stnds to-
y problms and planatory
tionwith whih it has n
.s7ntworth
r ft that D'Ary
ms of Rltiue Grotb (192)
lridg': tslknapPrss of
nnss,d.' Lndmrk \Yritigs
|940, 82-832. Amstrdam:
London:Nlthun.
,g Biolgi,lDUk)pmt u)ith
, MA: rvrd Univrsity Prss.
orm' 2nd d. amidg:
rsion. .,-o'jfl/.'.
amrid*.'
arwin)
of Spisb es of Ntrl
:s i th Struggl for Life, was
l50 opis,and it was sold out
sold all of th opis to ook-
nmdiatlytO prpar a sond
,hihth last appard in 1872.
first part Darwin prsntd his
]asonsto think it plausibl. H
;ltion,pointing out th su-
wnt on to show (in an argu-
althus) how a struggl for
populations to a lvl gratr
l s u p p o r t . n d t h n h o w ' g i v n
ads to diffrntial survival and
ltion.
lplid his mhanism aross th
lt' iogographi distriutions,
logy, nd othrs-showing how
Ontogny and Phylogn 771
ntrrralsltitlnprovids plllatilnsin all tsras and how, onvrsly,
thsarasonfirm th tuth nd plwrof sltion.This is not so muh a ir-
ular argulnnt s ()l1tht is rirrfoing.Th argumnt is known thnially
as a osilill of idutios. |t Was pom()tdy th nglish philosophr of
sinWillianr lVhwll, and it was from this sour tht Drwin larnd of
its implrtan r]d powr.
Thr wr m:rjor hngs in th sussivditions of te rigin. fter
Darwirr pulishd, vry fw in tlr sintifi tlmtnunity ontinud to dny
rh truth of vtllution. Htlll,vr, Darwin's thory of natural sltion was
sujtdtO stringntsintifianalysis,and two objtions smdvry sri-
ous. Th first w:ls that Darwin had no adquatthory of hrdity-wht w
today would all a thor1.tlf gntis-and without suh a thory lt Was not
lviottsthat a mhanisrnlik sltionluld lrav a lasting fft.Th s-
ond ojtion, asd orr physiists' lultitlns aout suh things as th
hatingpowr of th sun, ws tlrat th arth was far too yung for so lisurly
a mhnism as sltion tr hav takn fft.
As w now know, tlr first ljtionwas wll takn, nd it was not until
Mndl's thory of hrdit'Was rdisovrdthat this ojtion ould
ountrd.Th sond ojtilnWas not wll takn, austh hating f-
ft of radioativ day (and th tlnsquntmuh oldr ag of th arth)
Was n()t thn kn<lwn.Hlwvr' Drwin flt oligd to spak t oth pro-
lms, and to ountr thm h st:rrtdto rly mor and mor on supplmn-
tary mhanisms to sltion, most partiularly stl-alld Lamarkism, th
inhritenof aqr.rirdharatrs.
Unfortunatly, th nd produt of tlr various hangswas no longr an l-
gant tnddssayut lngthy and umrsonltrat. It is for this rson
that today's radrsgnrally prfr th first dition of te origi.
BIBLIOGRAPHY
D r w i n , . l 8 5 9 ' o t h r i g i r l fS p c i s b 1 ' sN
t lf t u r l S l t i o , l r t h
Prcsrutiltof l:t,lurdI|si th Strgglfor l'if. Londorr:Jol-rn
Mury.
1'959..hrigi f Spis|,htlrlsI)rt'uitt: Vrioru ?r..
Pkham,d.Philallphie: Univrsitvof Prrrrslvnia Prss'
Rihrds,R. J., nd . Rus,ls.2008. h mbridgcrlm1lil t th ,.rigi
rbilg: rrilg UnivrsityPrss. -^4.R.
of Sis.''
ntogen nd hIoge (StephnJay Gould)
|n toge d P,logya dtaild :rnalysisof th rlirtionshiptwn
growth of th iniividual<lrganismand its anstrYis tightly intgratdwith a
historial study of rpitr.rlationand othr pattrns of orrspondn-
twn stagsin dvlopmntand thos <lfvolutiorraryhistory.
..grathain lf -
Th anal<lgytwn individul dvlopmnt and th
ing'' ws fmiliar ttl risttltl :rnd othr lassial authors. Gold shows how
this id w:rs rshpdir-rth ()nttof nintnth-nturyGrtnan idalism
772 ontogny nd Phylgny

Tird, using a lok m

to intrprt studis of th dvloping nrryo. Karl rnsf vn Bar:saw in-
ar nd mmrs in a
rasing divrgn from omlon lnitial odv plrr as vrtratslvlop
raiyzd quantitiltivl)
distint adult lrms. rnst Hakl ptdD:lrwin's vidnfl volution
r r n sa n d p r i m n r i' l
ut not th adquy of his mhnism. infrrd that strr.rtrl innova.
volvd. In short, lr lid
tion involvs th addition of nw trminal stagsto Ontogny.Th rsulting
dvlopd mrrflmlI
rpitulation of abbrviatd anstral ftlrms t-t-l o]ifidas th ioge-
Th most ritial lf r
nti laW' from whih anstralrlationshipsould infrd and lost forms
l t r t i o n n r 1i.m p l i i r l r l sl .
prdited, thry ovroming dfiinisof tlr fossil rord.
mlrphi' juvnilizdds
In th bsnof a satisfatorythory of individual vriatilnand hrdit,v,
stonglv in favor tlf r
akl's idas gaind wid urny in th post.DaWinian ra. Hkl in-
ris to novl ombinatr<
sistd that rmirraladditilrrand apittrlation.rth rnharrism,th ffi-
dvlopmnt' This pro
int aus of volutionary hang. Gould shows that this th<lrydpndd
spializzrti<ln, and popr
on inhritan of aquird lrirratsand an trrrklowldgd vitalisr tlrat
has a uniqu potntialt(
wr haratristiof most volutionary thought irt that tirn.
nrv highr ta' Goul
It is oftn supposdthat rpitultion fll from gra ausvidn-
many othrs hav sin
umulatd that volurionrvnovltisn ppar at anv stgin th lif yl.
tnds th tim alltlttdt
In fat, suh insrtiorrshad n aommdatd as prmissilptilnsto
r a p i d . l r i s i m p l i t di r
th iogntilaw,,vn by Hakl hirnslf.Gould rgusthat it was work in
inid rins, inrasdstl
primntal mbryology and th mr]nof ndlian ]ntisthet mad
Gould infrrd tlrat h.
th thor.untal' Ntural sltionan and dos at on all stgsof tlr
w i h h a d b n o n i v
lif yl. Thrftr, rft of rhanism and prditiv powr' th pattrns
d Pbllgy.His pr
of rlationship btwn stagsin dvlopmntand thos of volutionary his-
stimulat produtiv int
tory wr subjtdt sussionf parlyzing lassifiations'Ths inhi-
ary iollgy lras n ar.t
itd furthr work on what hd om to alld htrohrony.
not n limird t<lthis I
In tlgn d Phlogy Gruld rnd thr mjor olrtirrtistlrat r.
has n itd in artils
vivd htrohronyas a vigorous fild of inquiry. First, h shiftdth fous of
study from mapping Pattrns to th analvsis of dl,lopnlntalmhanisms
|LloGRAPHY
that an studid xprimntally in living organisms and infrrd from nal-
Ogolls pattrns in fossils of tint forms' Sond, h stalishd simpl A l r h ,P . ' S . J . G l t r l dG, .
onto!]nyand phyllg
modl in whih th timing of ovrall dvl<rpmrrt and th onst f r;lrodu.
G o u l d , S . J . 1 9 7 7 . t g a
tiv maturity n b advat-rdor rtardd in ral trms' against lok tir. Prss.
This shows that th two fundamntl pattrrls of htrohl1yI1lr aris N a m r a 'K . ' | . 1 9 9 7 S .l
in two ways' with radially diffrnt dmogrphi and ologial ons- Deuloplnt.Bltim
quns for th organisms invrlvd,as shown in th ft>llolvingta[rl.

Rprodutiv
oparin, Alksandr
Somti dvlopnrnt mturirtion Shift in attrn Alksandr l. oparin, a l
lrtio fluntiaI pionrs of th l
N o h r r g i n t i m i n g Rpitulcttirl
( lration) in th Sovit Union in 19
No hngin timing lrrttil Pdomorphosis ( 1 9 6 ) ' ) p a i n p r s n t
(y truntiol-r)= P gsis a r t h . O p r i n . s i d si n i
Rtrdtio No harrg in timing Pdllmorphrlsis mtal in stlishigrn
(y rtadatit>n)= N otlt- Ralizing th nrml
No hangin timing Rt'trdtil Rpitultil fild of iohmistry, o
(v polongation)
of th origin of lif fror

l. Karl nst von Bar saw in-
'dv plan as vrtratsdvlop
)rwin'svidrrfo volution
r.rfrdthat strutur:rltnnov-
'gsto ontogny.Th rsulting
amodifid as th iog-
luld infrd and lost forrr-rs
th fossil rord.
]ividual vriation and hrdity,
ost-Darwinir-r ra. Hakl irr-
n w th mhanism, th ffi-
Lowstht this thory dpndd
unknlu.ldgd virirlism tlrt
-' ^+.^t r( iir -I .r L .
dL lIl r L
| t l
ror gra ausvi1na_
irrJt n}'st.tgin h lif 1.l.
, da s r m i s s i l x
iould argusthat it was wok in
lf Jr4ndlin gntistht mad
nd dos at on all stgsof th
c prditivpowr' th pattrns
t and ths tlf volutior-rirry
lirrglssifiations.lrs inhi-
ld htrohrny.
trmajor ontriutirlnsthat r-
ir,v.First, h shiftd th fous of
; of dvlopmntalmhanisms
ganismsand irrfrrdfrom anl-
iond,h stalishda simpl
lnt and th onst of rprodu-
t r:rltrms, gainst lok til.
s of htrolrronyan h ris
lgraphi and olgial ons_
in th followirrgtal.
Shift in pttrn
Rpitltkln
(y lrtion)
Pdomorphosis
(by trunationI = Prlgtlsts
Pdomorphosis
(y rtardation) = N ote
Rpitultit-l
(y prlongatiorr)
pri 77
Tlrird, usirrga lk mldl, h shorvd tht rh tgoris of htrtlhrony
ar nd mmrs in a ontinuum of potntial volutiony hang that an
analyzd quantittivly, using mthods of allomtry to study vllving pt-
trns and pimntsin dvlopmntaliology to iuidt th posssin-
volvd. In short, lr laid th groundrvork for a alulus of htrohrony, latr
dvlopdmo fi>rmallyin onjuntion with a group of ollagus.
Th most ritil of th distintior-rsGould ltridtd,it-ltms of its vl-
lutionary impliations, is tht twntwo prossstlrat giv ris tO pd<l_
morphi' ;uvnilizddsndants.Prognsisours whr sltionats mor
strongl in favor of arly rpr<ldutionthn it .loson rrrorpho|lgv,givlng
ris to nlvl onrinations of haratrs as rsult of th shift in timing of
dvlopmr-rt. This pross involvs a shak-up in morphology, ltlss of adult
spialization,and popultion growtlr alrtdy th short lif yl. lt
has a uniqu p()tntialto gnratraid hrrgnd tlr suddn tnrgn<lf
nw highr ta. Gould itd wll.known xampls frtlm rlir litratur;
many othrs hav sin n doumntd. Nt.ltny'On th oth hand, -
tnds th tim llottd to aly stgsof dr,ltlpmnr,whn g()Wthis mlst
p t i o n st o
rapid. tt is impliatd in rapid siz inras,notaly inluding that of horn-
inid raills, inrasdsoializtiln,and othr forms of spializtion.
Gold infrrjthat htrlhronyis undr th tltrtrolof rglll:rtorygl1s'
whih had n onivd ut not yt dourntdwhn h wrote togtl
d Pb,logy.llis prdition tht rnwd intrst in htrony would
his_
stimul:rtprodr-rtivintrtion twll mtllular gltisnd volution-
ary iollgy has n arpl1,orn oltt. But th influnof Gould's lok has
not n limitd tO this fild. Its road sop rrd signifianar suh tht it
lras rritd in tilsptrlllishdin ovr.500 sintifijounls.
BIBLIOGRAPHY
A l b e r hP, . ' s . J . G < l t r l Gl ,. . O s t r , n d D . B . W a k . l 9 7 9 . S i z n is h a p i r r
ont()gny and phylogll', PIobiollg 5: 296-17,
Gould' s. J. 1977.()togd PhIog. mridg,MA: arvadUnii riti'
Press.
Nrirr,K. J. |997, Shpsrlf illtt':Thc t,oltioof G rothtut,!
D t ' l o p m t . B a l t i m o r : . | o h n s H p k i r - r s U n i v r s i t 1 . P r s-sR.. D . . .
oparin, AlksandrI. (1894-1980)
AlksandrI. oparin, a l1ing R u s s i n i o m i s t ,W a s a m o n g t l r m o s t i r l -
fluntil pionrslf th stLrdyof tlr mrgnlf lif. Irr ooklt pulislrd
in th Sovit Union tn 192-4,and latr in a to dtaild ook, rigi of Lif
(196|' )parin psntdhis raktrugh tlrory ()n th origirr of lif on
arth. Oprin's ids insird lan sarlrrsworldwid arrlwr itrstru-
-l
mntl in stalishingin th 950s a sintififild dvotd to th sujt.
RaIizing th normous omplxity f th living il, valdy th rising
fild of iohrnistry,Opzrir-r rjtd rrhsirplisti Inlranistisnaritls
of th origin of lif from lrlattr and vitalisti laims of an unridglgp
774 1lri (

twnmattr nd Iif. Instd,rling oll nW sronomilnd gllogil |BLloGRAPH\

tlroris irnl lta, Il formtrlatd t.ttualistivolutiOnav sllrio that
Fr, I. 2000. Th mrg
postultd spifi physiohriirlonditions' onduiv to lif, On th pri- Nw Brunswik,N.