Okubo 1986 PDF

Adv. Biophys., Vol. 22, pp.
1-94 (1986)
DYNAMICAL ASPECTS OF ANIMAL GROUPING:
SWARMS, SCHOOLS, FLOCKS, AND HERDS
AKI RA OKUBO
Marine Sciences Research Center, St at e University of Ne w York,
St ony Brook, Ne w York 1779d-5000 and Ecosystems Research
Center, Cornell University, Ithaca, New York 14583-2701, U. S. A.
By t he t erm "groupi ng" used in this article we mean a phenomenon such
as insect swarming or fish schooling in which a number of animal indi-
viduals are involved in movement as forming a group. Groupi ng of or-
ganisms in mot i on--fl yi ng or swi mmi ng- - i s a frequent l y observed pheno-
menon. I n particular many di pt erous insects and pelagic invertebrates
are known to form swarms and schools (Chiang (1-3); Downes (4); Sul-
livan (5); Koyama (6); Hamner and Carleton (7); Byron et al. (8); Ueda
et al. (9); Cl ut t er (10)).
Ther e are many references to t he grouping phenomenon in t he litera-
ture, ranging from simple records to more extensive functional and be-
havioral studies. Yet most studies have been incidental and descriptive
in a qualitative manner. Only a handful of precise studies of the move-
ment of individual organisms within a group have been made. These are
wi t h flies by Land and Collett (11), and wi t h midges by Chiang (3), Oku-
bo and Chiang (12), Okubo et al. (13), and Jansen (14).
I n this article an at t empt will be made to describe t he mot i on of
grouping individuals kinematically, to model t he grouping on t he basis
1
2 A, OKUBO
of dynamics of animal motion, and to interpret t he grouping from the
standpoint of advection-diffusion processes. Also will be discussed dy-
namical model s for t he group size distribution as a result of interacting
processes of group amalgamation and splitting.
Apologia
Being characteristic of any reviews this article suffers from some biases.
I t is biased t oward my favorite animals and against bacteria. Thus an
enormous amount of literature concerning bacteria aggregation primarily
due to chemotaxis will be almost entirely neglected in this review. May
t he reader pardon t he speciesist!
I. BASIC CONCEPTS FOR MATHEMATI CAL MODELI NG FOR GROUPI NG
The mathematical model for grouping devel oped here assumes first that
organism movement is more or less at random. I n this concept of random-
ness, organism displacement or any kinematic quantity takes irregular
fluctuations whose individual values cannot be det ermi ned by t he con-
trollable or "macroscopi c" conditions but rather t he average or statistical
properties of t he mot i on can be det ermi ned uni quel y by the conditions.
The degree of randomness in grouping mot i on varies not only with
species but wi t h certain physiological and ecological conditions for a given
species. Thus, for schooling fish, t he movement appears t o be domi nat ed
by t he deterministic element or organized motion, although one cannot
fully underst and or predi ct t he precise pat t ern of mot i on in schooling.
On the other hand, small insects like midges in a swarm apparently ex-
hibit disorganized movement s whi ch might be regarded as almost com-
pletely random. I n effect it is often difficult for us to make a clear distinc-
t i on bet ween deterministic and random movements.
I n this context of animal grouping we are not particularly concerned
wi t h t he migratory aspect of organisms, i . e . a mass movement of a group
of organisms as a whole, but rather concerned wi t h t he internal st ruct ure
of a group such as t he spatial pat t ern of t he organisms relative to t he
center of mass and also with the mot i on of organisms in a group. Even
t hough we restrict ourselves to this specific aspect of grouping, a simple
random walk or pure diffusion process itself never serves as a model to
account for t he phenomenon. Random mot i on alone makes a group of
animals spread out to occupy a larger area or vol ume as t i me progresses
- - t hi s t endency of spreading is simply a consequence of a bl i nd random-
DYNAMIC ASPECTS OF ANIMAL GROUPING
ness. Rainey (15) remarked that, despite the diffusive action of air t ur-
bul ence as welt as t he apparently random orientation of flying individuals
in swarms, Desert Locust s maintained their cohesion wi t hout appreciable
dispersion while travelling distances of hundreds of kilometers over many
days.
To deal adequately wi t h t he modeling of swarming, therefore, it will
be necessary to i nt roduce into t he theoretical model, a furt her complica-
t i on somewhat analogous to gravitational attraction or to intermolecular
forces. I n ot her words, t he mathematical model for animal grouping
also assumes that t here are certain regularities superposed upon t he ran-
domness in t he mot i on of organisms in a group. Being a deterministic
element of mot i on and having a nat ure of attractive forces, this regularity
plays a crucial role in maintaining animals in a group.
II. BIASED RANDOM WALK MODELS FOR ANIMAL AGGREGATION:
(ADVECTION-DIFFUSION EQUATION)
A random walk model has been considered as a starting point for the
analytical st udy of animal dispersal (Pearson and Blakeman (16); Brownlee
(7); Skellam (78); Alt (19); Patlak (20, 21) ; Kareiva and Shigesada (22);
Yasuda (23)). A passage from a random walk formulation to a cont i nuous
limit leads to an equation of diffusion ("diffusion approximation"). Skel-
lam (18) in his celebrated paper, used the random walk model and the
law of diffusion for t he st udy of spatial expansion and di st ri but i on of
animal populations.
Whet her or not an animal moves from a position x 1 tO an adjacent
position x 2 depends on many conditions. Thus, t he probabi l i t y that an
individual moves from x 1 to x~ in t he succeedi ng infinitesimal interval
of t i me depends on t he biological and physical conditions prevailing at
or bet ween t he two positions. Skellam (2d, 25) proposed t he equations of
diffusion appropriate for t hree distinct cases, i.e. t he movement is af-
fected by conditions at x 1 ("repulsive"), at x~ ("attractive"), and bet ween
the positions ("neut ral ").
In this section an at t empt will be made to generalize t he random
walk probl em in such a fashion that t he transition probability bet ween
t wo successive points depends upon t he conditions prevailing at any loca-
t i on bet ween (and including) these t wo points. Application of this random
walk model to the spatial di st ri but i on of grouping animal will be discus-
sed.
4 A. OKUBO
1. Basic Equation of Generalized Random Walk (or Flight)
F o r s i mp l i c i t y we c o n s i d e r a o n e - d i me n s i o n a l r a n d o m wa l k o f ani mal .
S u p p o s e t h a t at t i me i nt e r va l s r , 2r , 3 r , . . . , an a ni ma l e i t he r s t e p s a
di s t a nc e ~2 or r e ma i n s at t h e or i gi nal p o s i t i o n wi t h s pe c i f i e d t r a n s i t i o n
p r o b a b i l i t i e s .
L e t k+(i, i + l ; m ) b e t h e t r a n s i t i o n p r o b a b i l i t y t h a t i n an i nt e r va l r
at mr t i me , an a ni ma l mo v e s b y t h e di s t a nc e ~2 t o t h e r i ght f r o m a p o s i t i o n
i t o an a dj a c e nt p o s i t i o n i + 1. Li k e wi s e k-(i, i - - 1 ; m) d e n o t e s t h e t r a ns i -
t i o n p r o b a b i l i t y t h a t i n t h e s a me i nt e r va l o f t i me an a ni ma l mo v e s t o t h e
l e f t f r o m i t o i - - 1 . Al s o ko(i; m) d e n o t e s t h e p r o b a b i l i t y t h a t i n t h e s a me
i nt e r va l o f t i me an a ni ma l r e ma i n s at a p o s i t i o n i. T h e c o n s e r v a t i o n of t h e
t r a n s i t i o n p r o b a b i l i t i e s r e q u i r e s
k+(i, i + 1 ; m)+k-(i, i - 1 ; m)+ko(i; m) = 1 ( I I . 1 )
L e t p(i, m) b e t h e p r o b a b i l i t y t h a t t h e a ni ma l i s at i at mr a f t e r t h e
b e g i n n i n g o f wa l k f r o m i----0 at m = 0 . T h e p r o b a b i l i t y o b e y s t h e f ol l owi ng
e q u a t i o n s :
p(i, r e + l ) = k +( i - 1 , i; m)p(i -1, m)+ko(i; m)p(i, m)
+ k - ( i + l , i; m)p(i+l, m) (II. 2)
p(i, 0) = ~,0 (II. 3)
De f i n e 2- a n d / 2 - f u n c t i o n s s u c h t h a t
2( i +0, m) - k+(i, i + 1 ; m) ( I I . 4 )
p(i-O; m) =-k-(i, i - - 1 ; m) (II. 5)
wi t h 0<_0<1.
E q . ( 1 1 . 2 ) i s t h e n r e wr i t t e n as
p(i, m+ 1) = 2(i -1 + 0 ; m) p( i - 1; m)+ko(i, m)p(i, m)
+ / 2 ( i + 1 - 0 ; m)p(i +l ;m) (II. 6)
a n d ( I I . 1 ) is e x p r e s s e d b y
ko(i, m)+2(i+8; rn)+12(i--O; m) = 1 (II. 7)
We al so de f i ne
~(i +O; m) - 2( i +0; m)-/2(i+O; m) ( I I . 8 )
wh e r e / 9 r e p r e s e n t s a d e g r e e o f a n i s o t r o p y i n t h e r a n d o m wal k. Eq. ( I I . 6)
ma y b e s ol ve d s u b j e c t t o t h e i ni t i al c o n d i t i o n (I1. 3) a nd a p p r o p r i a t e b o u n -
d a r y c ondi t i ons .
DYNAMIC ASPECTS OF ANI MAL GROUPI NG
We can distinguish three types of random walk by the value of 0:
0=0, 1/2, and 1. For 0=0, the transition probabilities depend only on the
conditions at the point of departure and thus they are classified as being
induced by "repulsion" (Skellam (25)). For 0=1/2, the transition prob-
abilities depend on the conditions at the half way between the points of
departure and arrival, and thus are classified as "neutral." For 0=1, the
transition probabilities depend only on the conditions at the point of ar-
rival, and thus they are classified as being induced by "attraction" (Skel-
lam (25)). We could consider any intermediate cases with the value of
0, 0<0<1, but it is sufficient to represent the case by 0=1/2.
2. Diffusion Approximations
We now consider the random walk problem in a limit where the step
lengths and time intervals tend to zero, i.e., diffusion limit. Write i~7=_x
and mr =-t, so that p( i + 1, m+ 1) for instance implies p(x+~7, t + v).
Thus (II.6), (II.7), and (II.8) can be expressed as
p(x, t +r) = 2(x--~7+0~7; t)p(x--~7, t)+ko(x; t)p(x, t)
+ ~(x + ~7--0~7, t)p(x + ~7, t) (II. 9)
ko(x; t)+2(x+O~; t)+t~(x-O~7; t) = 1 (II. 10)
fl(x, t) = 2(x, t)--fz(x, t) (II. 11)
Expanding (II.10) in a Taylor series with respect to 07/ and making
use of (II.11), we obtain
ko(x; t)+2(x; t)+~(x; t)
92
= 1 -O~i~x~(X, t) - 1/202~72~X~X 2 {1 -ko(x, t)} +0(~/8) (II. 12)
A similar expansion of (II.9) with the use of (II.11) and (II.12) results
~Pt ~+0(~ )
= - ~Tx (~p)
2 a2 0 ~p
o~p
]
+(1 - ko)TZx~ j +o(v ~) (ii. 13)
where p,/3, and ko are evaluated at x and t.
We will take limits of ~--~ 0 and v--+ 0 under the following condi-
tions
6 A. OKUBO
l i m V~/r = U~(x, t) (II. 14)
~ ] , r - - - * 0
lira 722/2r = Do (constant) (II. 15)
~ 2 , v - - - * 0
whe r e Ua has di me ns i ons of vel oci t y and is a s s ume d t o be a s moot h f unc-
t i on of x and t, and Do has di me ns i ons of di ffusi vi t y. T h e l i mi t i ng pr oce-
dur e on (II. 13) r esul t s
at - 3x (U~p)+Do ( 1- 20) ( 1- k o) p
+- ~x {(1-ko) ~3~Px } 1 (II. 16)
We defi ne a vari abl e di ffusi vi t y D( x, t) by
D(x, t) ~ Do{1-ko(X, t)} = Do{2(x, t)+/2(x, t)} (II. 17)
Not i ce t ha t r a n d o m walks, wi t hout a non- c ons t a nt r emai ni ng pr obabi l i t y
cannot have a vari abl e di ffusi vi t y. Eq. ( 1 1 . 1 6 ) is t h e n r ewr i t t en as
- ~ x ~ l U ~ - ( 1 - 2 0 ) ~ x l p j + - ~ x ~ D ~ x ) (II. 18)
Z/ - =
T h e i nt e r pr e t a t i on of (I1.18) is clear. Ani s ot r opy i n t he t r ans i t i on pr ob-
abi l i t i es pr oduc e s a dr i f t Ua of ani mal , so does spat i al i nhomoge ne i t y i n
t he di ffusi vi t y D. However , t he di r ect i on of t he dr i f t due t o a vari abi l i t y
i n di f f usi vi t y de pe nds not onl y on t he gr adi ent of di f f usi vi t y b u t also on
t h e behavi or of ani mal i n maki ng a move, whi c h is par amet er i zed by 0.
Fo r 0 < 1/2, t he ani mal t e nds t o dr i f t t owa r d t he di r ect i on wher e t he di f-
f usi vi t y decreases, i.e. t he di r e c t i on wher e t he r emai ni ng pr obabi l i t y i n-
creases t h u s a degr ee of r a n d o m mo t i o n decreasi ng. For 0 > 1 / 2 , on t he
ot he r ha nd, t he ani mal t e nds t o dr i f t t owa r d t he di r ect i on wher e t he di f -
f usi vi t y i ncreases, i.e. t he di r ect i on wher e t he r emai ni ng pr obabi l i t y de-
creases t hus a degr ee of r a n d o m mo t i o n i ncreasi ng. For 0 = 1/2, t he dr i f t
vani shes aut omat i cal l y i r r espect i ve of t he vari abi l i t y i n D. Th i s behavi or al -
l y- dr i ven dr i f t pl ays a n i mp o r t a n t rol e i n mode l i ng ani mal di spersal
( Okubo (26)).
Fu r t h e r di scussi ons on t he advect i ondi f f usi on equat i on (II. 18) will
be ma de f or t he t hr e e t ypi cal cases of 0. Fo r 0 - - 0 (repul si ve), Eq. (II. 18)
can be expr es s ed as
~p _ ~ ~2
at Ox (U~P)+~ix~(DP) (II. 19)
DYNAMI C ASPECTS OF ANI MAL GROUP I NG
Thi s is a Fokker-Planck equation characterized by t he drift U~ and dif-
fusion coefficient D(x, t). I f we inspect t he contents of t he Fokker-Planck
equation model leading to (II.19), it should become clear that t he model
is based on transition probabilities that depend only on t he point of depar-
t ure and t hus t hey are classified as being i nduced by repulsion (Okubo
(26); Lapidus and Levandowsky (27)). Many models for animal dispersal
assume t he advection-diffusion equation of t he form of (II.19). The reader
may consult Shigesada and Teramot o (28); Shigesada (29); Nagasawa
(30); Dobzhansky et al. (31).
Heterogeneity in t he envi ronment can result in t he spatial variability
of D, and t hen t he behavioral response of organisms to t he stimuli may
produce a drift in one direction or another. For illustration purpose we
take a case where diffusivity varies linearly in space, and show t he t em-
poral behavior of organism concentration in a one-dimensional domain
bounded by x = 0 and L. At t he boundary we impose t he conditions that
no organisms can pass t hrough, i.e. reflective barriers. Isot ropy in t he
movement is assumed so that Ua- O. The diffusivity is given by D=
Do( l +k x ) , k>0. Initially we concentrate organisms near t he center of
t he domain.
Numeri cal results of Eq. (II.18) with t he specified form of diffusivity
and t he boundary condition are shown in Fig. 1 for t hree cases of 0. For
0=1/ 2 (neutral) organisms t end to distribute uniformly despite t he spatial
variation in diffusivity. For 0 =0 (repulsive) organisms t end to aggregate
where diffusivity is small, while for 0=1 (attractive) organisms t end to
aggregate where diffusivity is large.
A steady-state distribution of organisms can be obtained from Eq.
(II.18) by setting 3p/at =0 and integrating with respect to x. For t he case
of D=Do ( l +k x ) and U~=O, we integrate once to find
Do(1 +kx) oa~Px +(1--20)kDop = constant = 0 (II. 20)
The reflective boundary condition guarantees t he constant of integration
to be zero. Int egrat i ng (II.20) again results in t he organism concentration
20k
p = N (1 +kL) 2o- 1 (1 +kx) 2o-1 (II. 21)
where N is t he total number of organisms in t he domain. Therefore t he
asymptotic steady-state distributions for t he t hree cases of 0 are given by
A. OKUBO
0. 5
P
0
0. 5
P
0
0. 5
P
0
t=O
0.5
t =50
P
0
t=200
t =O
51
t = 5 0
t=200
t=O
_L
t=50
t=200
0 = 0
~=5
t =1000
0 = 1 / 2
t = 5
0.5
t=75
P
0 0. 5
/ =1000
P
0
8 = 1
f = 5
0..5
f=75
P
0
t = l O
t = 100
0. 5
t =4000
P
t =10
f = 100
t =4000
t ~l O
t = 100
f
t =4000
0 x 0 x L x
Fi g. 1. Te mp o r a l be ha vi or s o f or ga ni s m c o n c e n t r a t i o n di s pe r s i ng i n a h e t -
e r oge ne ous e n v i r o n me n t f or t h r e e t y p e s o f t r a ns i t i on pr oba bi l i t i e s : 8 = 0
( r epul si ve) , 6 = 1/2 ( neut r al ) , a n d 0 = 1 ( at t r act i ve) .
p = N in (1 +kL) (1 +kx) -1 for 0 = 0
p = N / L for 0=1/ 2
(
2k
( l +kx) for 0 = 1 ]
p = N (1 +kL)2_ 1 )
( I I . 2 2 )
Organisms orient themselves to t he effect of stimuli by taxis and
kinesis (Fraenkel and Gunn (32)). A great deal of effort has been made
for mathematically modeling chemotaxis of bacteria (Alt (79); Keller and
Segel (33-35); Nossal (36); Rosen (37, 38); Rosen and Baloga (39); Odell
and Keller (dO); Oosawa and Nakaoka (47); Lapidus (42, 43); Segel (,t4,
45)). The model t urns out to be an advection-diffusion equation essentially
t he same as (II.18), coupled with t he diffusion equation of t he chemical
attractant concentration. Bacteria's chemotactic behavior is incorporated
in t he parameters U~, O, and D appearing in Eq. (II.18). Also Oosawa and
Nakaoka (dl ) developed a model to describe t he tactic behavior of micro-
organisms in whi ch individual cells are treated as particles having internal
state variables and environmental conditions such as t emperat ure gradients
influence t he behavior of a cell t hrough its internal state. Lat er Lapidus
(43) showed that t he results of Oosawa and Nakaoka could be obtained
using only macroscopic quantities.
Many random walk models for animal dispersal do not take into ac-
count interference bet ween individual organisms, and t he parameters
pert i nent to t he model are i ndependent of t he animal density although
t hey may depend upon space. For an adequate model for animal aggrega-
tion it may be necessary to include density dependence in t he pertinent
parameter. Along this line of t hought Aronson (46) has recently gener-
alized t he random walk model in considering transition probabilities that
depend upon t he local population density. Thus t he resulting diffusion
equation becomes nonlinear since t he diffusivity is density dependent.
Shigesada and Ter amot o ( 28) developed a mathematical model of
advection and diffusion to explain t he spatial distribution of aPimal popu-
lations that are principally controlled by interference bet ween individuals
and ot her environmental conditions. Thei r formulation is based on t he
assumption that animals move under t he influence of t he following funda-
mental forces: (1) a dispersive force associated wi t h random movement
of animals; (2) an attractive force, whi ch induces directed movement or
drift of animals t oward favorable environments, and (3) population pres-
sure due to interference bet ween individual animals. The concept of t he
10 h. OKUBO
force associated with population pressure was originally proposed by
Morisita (47, dS).
The model equation of Shigesada and Teramot o is expressed in
t erms of an advection-diffusion equation for organism concentration
S ( x , t ) . I n a one-dimensional space t he equation is wri t t en as
~l-~x ) + a~x~[{a(x)+~(x)S}S]
OS O S (II. 23)
3t 3 x
where q~(x) denotes t he potential of t he environmental attraction, whi ch
induces advection velocity Ua(x)-----Oq~/Ox t oward favorable regions (see
also Chapt er V.5), and D = _ a ( x ) + ~ ( x ) S represents a virtual diffusivity,
dependent upon spatial inhomogeneity and population pressure. Thi s
form of t he diffusivity is consistent with Morisita' s semi-empirical formu-
lation. Shigesada and Ter amot o derived Eq. (II.23) from a biased random
walk, where t he transition probabilities depend upon t he conditions at
t he point of departure, i . e. repulsive transition, so that t he diffusion t er m
is expressed as a ~ ( D S ) / a x ~.
Anot her model for animal aggregation similar to Shigesada and Ter a-
mot o but formul at ed in discrete spatial units has been developed by Tsu-
baki and Yamamura (49). I n t hei r model one-dimensional space is divided
into habitat units, and movement s of individual animals are considered
as density dependent.
Let n ~ ( t ) be t he number of individuals in ith uni t at t i me t ( i = l , 2,
. . . ) . The numbers of individuals moving from t he ith unit to t he adjacent
( i + l ) t h uni t and ( i - 1 ) t h unit in a t i me interval ( t , t + 6 t ) are represent ed
by D ( i , i + l)n~(t)6t and D ( i , i - - 1 ) n ~ ( t ) S t , respectively, where D ( i , i + l )
and D ( i , i - - 1) denote transition rates of an individual from t he ith unit
t o ( i +1) and (i --1) units, respectively. Movement s of an individual from
t he i t h uni t to ( i +m) t h units (m>2) are assumed to be of t he order of
(6t) 2. The n t he t i me rate of change of h i ( t ) is wri t t en as
dnl
at = D ( i - - 1 , i)n~_l--D(i, i - - 1 ) n ~ - - D( i , i + l ) n i + D ( i + l , i)ni+l , i > 2
dn~ (II. 24)
- D(1, 2)n~+D(2, 1)n~
at
We now assume that each unit has a carrying capacity for organisms,
K, and t he rate parameters are expressed as
D( i , i +1) = a i ( 1 - n ~ + l / K ) for n~+l<K ]
= 0 for ni +i >_K I ( I I . 2 5 )
DYNAMIC ASPECTS OF ANIMAL GROUPING 11
D( i + l , i) = a2( 1- nJK) for n , < K
(II. 26)
/
= 0 for n~>>_K
where ~r 1 and % are constants. Eqs. (II.25) and (II.26) state that a uni t
occupi ed by individuals more than K cannot be moved in by another in-
dividual and that only when a uni t is occupied by individuals fewer than
K, t he uni t can accept individuals from t he neighboring units at a rate
proport i onal to t he difference bet ween t he carrying capacity and t he
number of individuals already occupi ed the unit. It is wort hy of noting
t hat t he individual movement specified by (II.25) and (II.26) is categorized
as being attractive.
The existence of an equilibrium distribution of animal in t he units
depends upon t he ratio of % and %, i.e. t he relative intensity of t he for-
ward and backward transitions. I f a=/~1>1, t he organism di st ri but i on
attains an equi l i bri um n** that is given by
K
n~ * = 1 + 2*- X( K/ nl - 1) (II. 27)
where ,t=a2/~r 1 and t he value of n 1 is det ermi ned by t he condition that
~. n ~ = N (total number of animals). Thus t he equilibrium di st ri but i on is
/=1
approxi mat ed by
where
n~* = i ~ 2 {ln (A+2-i+l)--ln (A+2-i)} ( i i . 2 8 )
A = {2 ~v/~- 1} -~ (II. 29)
Animals t end to aggregate t oward t he first unit. The aggregation is more
conspi cuous for large values of 2, and the first few units are occupied by
a full capacity of individuals, i . e . K . If, on t he ot her hand, ~ r J c q = 2 < l ,
all t he individuals disperse t oward infinity (i --. co) and no equilibrium
can be attained.
III. KI NEMATI C DI STI NCTI ON BETWEEN DI FFUSI ON ( RANDOM WALK)
AND GROUPI NG MOTI ON
Imagi ne a group of organisms, t he total number of individuals of which
is N. All individuals are assumed to start grouping mot i on at a point
0(x). Aft er a certain t i me has elapsed some of t hem will have acquired a
12 A. OKUBO
resultant displacement in one direction, others in another direction. The
process is considered as stochastic.
Let x , ( t ) and v , ( t ) be respectively the displacement and velocity vec-
tors of ith individual in the group at time t ( i = l , 2 , . . . , N ) . Starting at
the origin (xi =0) at t =0, the distance travelled by the ith individual after
time t reads
f
t ~ ( t ' ) d t '
x , ( t ) = (III. 1)
We consider an ensemble average over a large number of grouping
individuals (N ~ oo) and denote the average by an overbar. Thus
1 1 N /
x ( t ) _= lim _~= ~, x , ( t )
N ~ c o n / i = 1
N
v ( t ) ~_ lim _~_ 5 2 , v , ( t )
N--. c~ -/V i = 1
(III. 2)
Actually the number of organisms in a group may not be so large as to
approximate the mathematical requirement of lim. For a small finite
number of individuals, the averaged quantities themselves will suffer
significant statistical fluctuations. In practice, however, a number such
as 100 individuals may be regarded as "large" so that the statistical fluc-
tuation will not become a serious problem. Instead of a large number of
individuals, the average values can be defined on a large number of re-
alizations of a single organism in a group under identical macroscopic
conditions. In this connection there is an interesting observation by
Bassler that a swarm could be reduced to a single individual of mosquito,
C u l e x p f p f e n s which yet continued to show the characteristic behavior of
swarming dance (Clements ( 5 0 ) ) . Also Goldsmith e t a l . ( 5 7 ) studied by
high speed motion photography the swarming behavior of midges, A n -
a r e t e p r f t c h a r d i Kim, and noted that a single and a few midges did show
movements characteristic of swarming by a large number of animals.
From (III.1) and (III.2)
x ( t ) = f t v ( t , ) d t , (III. 3)
J 0
Since our primary concern is the motion relative to the center, we con-
sider the deviations of displacement and velocity from their average
values ;
DYNAMI C ASPECTS OF ANI MAL GROUP I NG 13
x , ' ( t ) = x , ( t ) - - x ( t ) )
v ~ ' ( t ) =_ v , ( t ) - - v ( t ) I (III. 4)
wi t h
xl' = vl' = 0 (III. 5)
We now assume that t he random velocity field is statistically sta-
tionary. I n ot her words, any statistical properties associated with t he
velocity field remai n unchanged no mat t er when t he grouping begins.
The process v ( t ) may not be stationary in t he very initial period after t he
mot i on starts at x = 0 at t = 0 . However, it is reasonable to assume that
v ( t ) approaches stationary for later times. Here we are not particularly
i nt erest ed in t he initial stages of grouping but in t he later stages.
We now proceed to comput e t he variance of t he organism displace-
ment , x ~ t 2 ( t ) . The st andard deviation, i . e . t he square root of t he variance,
is a measure of spread of t he organisms, namel y a dimensional extent of
t he group ("group dimension"). We calculate t he variance as follows
(omission of primes in not at i on is assumed);
x - 2 ( t ) = f * o f ; ~ d t ' d t " = 2 f f o d t ' f 2 v ( t ' ) v ( t " ) d t ' t (III. 6)
For statistically stationary velocities t he velocity autocorrelation
funct i on " v ( t ' ) v ( V r ) is a funct i on of ] t " - - t f l = r only and v-2(t) is invariant
with time. I nt r oduce t he velocity correlation coefficient:
R ( ~ ) = V ( t ) v ( t + r ) / ~ (III. 7)
R(r) = R(--r) (III. 8)
Combining (III.7), (111.8) wi t h (111.6) we obtain
- ~ ( t ) = 2 ~ f t o d t ' f 2 R ( r ) d r (III. 9)
Thi s is t he relation originally obtained by Tayl or ( 5 2 ) for t urbul ent dif-
fusion. Eq. (III. 9) may be t ransformed into a more convenient form by
integration by parts (Kamp6 de Fdriet (53)):
x ~ ( t ) = 2 v - S t ,j't oR( r ) dr - 2~ f r o
rR(r)dr (III. 10)
Eq. (III.10) is t he starting point f r om whi ch one can underst and (physi-
cally, not biologically!) t he difference bet ween diffusion and grouping in
t erms of variance.
14 A. OKUBO
1
. g
\\
\ \ \ \ \ ~
Fi g. 2. Vel oci t y aut ocor r el at i on coefficient R(v) vs. t i me lag v (pure di ffu-
si on)
For an ordinary diffusion process we anticipate t hat an initially
strong velocity correlation weakens as t he t i me lag r is increased. I n
ot her words, t he velocity loses its statistical dependence on past values as
random movement continues. Thus R(r) woul d look like Fig. 2, where
t wo fairly idealized typical curves are shown. In either case R approaches
zero monot onousl y at large values of v. Thi s behavior of R ( r ) is charac-
teristic of diffusion, and makes
approaching constant values as t becomes large. However, for large t i me
t he first integral in (III. 10), which is proportional to t, will exceed t he
second integral, whi ch is essentially constant. Hence, for ~( t ) we obt ai n
where
( ; o R I . I )
-~(t) = 2 ~ t ~ 2Dt ( I I I . 11)
S
oo
D = ~ R(r)dr - ~ T (III. 12)
o
T = f : R( r ) dc (III. 13)
T is an integral time, which is a measure of the longest persistence t i me
for t he motion: and D is an effective diffusivity. Clearly (III. 11) is the
relation for Variance found for purel y random dispersal: Fickian diffu-
sion. The variance increases linearly wi t h t i me; t he standard deviation
b
Fi g. 3. Vel oci t y aut oeor r el at i on coefficient R( r ) v s . t i me lag v (groupi ng).
a, unde r da mpi ng; b, over dampi ng.
increases wi t h t he square root of time. Organisms initially concentrated
at a point t end to spread indefinitely.
To describe grouping kinematically, t he variance must become in-
dependent of t i me for steady-state group maintenance. It is sufficient to
find that t he first integral in (III. 10) vanishes asymptotically; therefore
t he correlation coefficient R ( ' c ) , which takes positive values for small z"
must take negative values for at least some large values of ~'. As shown
in Fig. 3a, t he R curve may oscillate about t he zero value, and t he oscilla-
t i on must be such that t he positive areas of t he R-z" curve cancel t he nega-
tive areas for sufficiently large values of z'. Anot her possibility is t hat t he
R curve crosses zero at a certain t i me lag, proceeds into t he negative re-
gion, and remains negative by approaching zero asymptotically in such a
manner that t he positive areas also cancel t he negative areas for suffi-
ciently large values of z" (Fig. 3b).
I n t he former case (Fig. 3a) the individual mot i ons appear to resemble
a pendul um-l i ke swinging about x =0 , and in t he latter case (Fig. 3b) t he
16 A. OKUBO
individual motions resemble a centrally-biased random walk. Experi-
mental confirmation on this kinematic model is to determine the velocity
correlation coefficient of individual animals in a group. This will be dis-
cussed in Chapter VII.
For either case of Fig. 3 the first integral of (III.10) vanishes and
the second intergral approaches a negative constant value, which we shall
denote - - L ~ / 2 . We then find
x ~ = L 2 (III. 14)
The variance thus attains a constant value; in other words the spatial
extent of group, i . e . group dimension becomes stationary.
The discussion based on (III.10) is purely kinematical. No explana-
tion is given as to how the requisite shape of the velocity correlation
coefficient arises from more fundamental mechanisms or processes. We
now proceed into dynamical modeling for animal movement.
IV. ANIMAL DYNAMICS
Animal dynamics deals with the spatial distribution of animal and the
motion in animal systems. The term "animal system" defines any bio-
logically significant aggregation of animals. Thus the study of the move-
ments such as in animal grouping, in predator-prey, prey escaping from
predator may be included in animal dynamics. A specific emphasis is on
the interpretation of the characteristic features of an animal system in
terms of the "forces," which are supposed to operate on the animal in-
dividuals and to govern their motions in the system.
We start with assuming that Newton' s first two laws of motion also
apply to animal dynamics:
1. The forces acting on an organism at rest or moving with constant
velocity are in balance.
2. When the force balance is broken, a time change of the momen-
tum of an organism occurs in the direction of the total force. The rate of
change of momentum is equal to the total force. If we consider times that
are small compared with those required for noticeable loss or gain in
weight of the organism, the change in mass can be ignored, and the net
force may be equated to the product of mass and acceleration of the or-
ganism.
Let P~ and F~ be respectively the linear momentum of an animal
individual identified by the index i (/-animal or/-individual) and the total
force acting on the animal. Newton' s second law is expressed as
iJ~ = F~(p~ =_ d p d d t ) (IV. 1)
The linear momentum is defined by
p~ = mw~ = m, r i ( I V. 2)
where ms, vs, and r, are the mass, velocity, and displacement of/-animal,
respectively. The law may also be written as
F~ = rn,a, : m, 6, = m,r~ (I V. 3 )
where as is animal's acceleration and the mass is assumed invariant with
time. We also define the kinetic energy of/-individual by
Ki = 1/2 r ni v, 2 (IV. 4)
In application of Newton' s law to animal motion we must realize
that the nature of the exerted force may be quite general. It may be of
physiological and behavioral origin, e . g . the optimal light intensity, or of
physical origin, e . g . wind stresses. It may be of ecological origin, e . g . the
"population pressure" for dispersal (Morisita ( 5 4 ) ) . It may even be of
sexual origin, e . g . attraction to the other sex. From a purely mechanical
standpoint a fish can swim forward by pushing its environmental water
backward; the surrounding in turn reacts to provide a driving force to the
animal. Newton' s third law of motion is valid in this case, i . e . when or-
ganism A applied a force to its environment B, B reacts by applying a
force of the same magnitude in the opposite direction on A. However,
we must stipulate that when B is also an organism, this action-reaction
law may not necessarily apply (Okubo (26)). Of course this is a postulate,
and common sense tells us that the forces which two animal individuals
exert on each other are generally not equal and opposite, nor lie along the
line joining them. Remember a predator chasing a prey! This peculiarity
in the force of biological origin plays a curious role in animal dynamics
as will be seen in the following.
Consider a group of N animal individuals (i =1, 2, . . . PC). Quite
formally we can divide the force Fs exerted on/ -ani mal into three com-
ponents
N
F~ = F,(~) +F,(~) + Z F~ (IV. 5)
j =l
where Fs e) stands for an external force acting on/ -ani mal due to sources
18 A. OKUBO
out si de t he group, F~ tr) for t he fri ct i onal f or ce act i ng on t he ani mal as it
moves, and Fi j is an i nt ernal force on / -ani mal due to j - ani mal i n t he
group. Nat ur al l y
F , - 0 (IV. 6)
As ment i oned previ ousl y, F, j as wel l as Fi ~e} does not necessarily obey
Newt on' s t hi r d law of act i on and reaction.
The equat i on of mot i on f or / - ani mal t hus reads
N
m,r, = F, I*) + F , ~r' + 22 F, j (i, j = 1, 2 . . . . N) (IV. 7)
j = l
Summi ng over all i ndi vi dual s i n t he gr oup and defi ni ng t he cent er of
mass of t he syst em by
N N N
R =-- ~. m~rd ~ ms - 22 m~rdM (IV. 8)
i = 1 i = 1 i = 1
we r educe (IV.7) t o
M/~ = FC~) +FC~l +FCa~ (IV. 9)
The first t er m on t he ri ght is t he t ot al ext ernal force oper at i ng on t he
group, t he second t er m is t he t ot al fri ct i onal force, and t he t hi r d t er m
represent s t he r esul t ant of all t he i nt er nal forces:
N N
F ~ = ~ 22 F~j (IV. 10)
i = 1 j = l
We shall call F(a) t he r esul t ant i nt er nal gr oupi ng force. For physi cal
part i cl es t he t hi r d t er m of (IV.9) vani shes as a resul t of Newt on' s t hi r d
law of mot i on.
For a fri ct i onl ess and free (no ext ernal forces) syst em,
M/~ = F<g~ (IV. 11)
wher e F{a~ may not vani sh. Thus , t he cent er of mass of an ani mal group
can move ar ound even i f it is i ni t i al l y at rest and nei t her fri ct i on nor
ext ernal st i mul i exist.
We next consi der t he t ot al angul ar mome nt um of t he ani mal sys-
t em, A.
N
A - 22 r~ X Pi (IV. 12)
i = l
Oper at i ng t he cross pr oduct by r~ on (IV,2) wi t h (IV.7) and s ummi ng
D Y N A M I C ASPECTS OF A N I M A L G R O U P I N G 1 9
over i, we obt ai n
where
-d = T (e) + T (~ + T (~) (IV. 13)
N
T (~) =_ ~ r i F , ( e ) : external t orque (IV. 14)
i =l
N
T (~) -- ~ r ~ F , (r) : frictional t orque (IV. 15)
i =l
N N
T(g) = W. ~ r ~ F l j : internal groupi ng torque (IV. 16)
i =1 j =l
Agai n t he i nt er nal t or que does not necessari l y vani sh, whereas i t does for
physi cal particles The t ot al angul ar mome nt um is not necessari l y con-
served for a fri ct i onl ess and free syst em of animals The whol e gr oup can
rot at e as t he cent er of mass can move ar ound.
Fi nal l y l et us consi der t he equat i on of mechani cal energy. Mul t i pl y-
i ng ri by (IV.7) we obt ai n
d / 1 3\ . . 2v
K ' = - d ~ 2 m ' v ' ) = r " F" *' + r " F*'r' + r ' }2 F'~ j =l (IV. 17)
where t he use is made of (IV.4)
The t ot al ki net i c ener gy of t he syst em K is
N 1 N . 1 N
i =l i =1 2, i =1
(IV. 18)
Summi ng (IV.17) over i we fi nd
N . N . N N .
R = W r~.F~ (e) + ~ ri.F~(~) + W W r ~. Fi : (IV. 19)
i =1 i =1 i =1 j =l
V. DYNAMI CAL MODELS FOR ANI MAL GROUPI NG
7. N - b o d y Dy n a mi c s
The dynami cal t heor y of ani mal gr oupi ng begi ns wi t h t he fol l owi ng
premi ses.
1) A gr oup consi st s of a great numbe r of i ndi vi dual s (N>> 1),
2) Th e mot i ons of gr oupi ng organi sms are obeyed by Newt on' s
laws of mot i on wi t h a possi bl e except i on of t he t hi r d law.
3) The i ndi vi dual organi sms are t r eat ed as mat eri al poi nt s; i n
real i t y organi sms are supposed t o be har d bodi es of fi ni t e "col l i si on vol-
20 A. OKUBO
: <
J
v
C M
r,.
0
0
F i g . 4. A s y s t e m o f g r o u p i n g a n i ma l s .
ume" or a sphere of influence of vol ume vc so that we require t he total
vol ume of groupi ng domain V to be much larger t han N v c .
Our main interest is t he mot i on of grouping organisms relative to
their cent er of mass (Fig. 4). Usi ng Eqs. (IV.7) and (IV.9) we will derive
t he equation of t he relative motion. Define t he displacement vect or of i-
animal from t he cent er of mass by
re' = r e - R (V. 1)
Fr om (IV.Y), (IV.9), and (V.1) we obtain
. rg/. N N N
m, r , ' = F, (~) _ 2 ~ Yl, F, (~) + Fi (r) - m~ ~ F, (r ) + Yl, F ~ :
i=i M i=i j=i
N /V
m,, ~ W, F ~ : (V. 2)
M i=l j=l
We will express t he t hree kinds of forces more specifically. First, t he
external force acting on/ - ani mal depends only upon t he position of t he
organism at t i me t.
F~ (~) = rn~A~(~)(R +re' , t) (V. 3)
Second, t he frictional force is assumed to be linearly proportional
to t he velocity of organisms
F , ( r ) = _ m & ( i ~ + ; , ' ) ( v . 4)
where k, is t he frictional coefficient. Strictly speaking t he relation (V.4)
should be applicable only when t he Reynolds number of t he movi ng
organism is small compared with uni t y (Vogel ( 5 5 ) ) . Reynolds number s
associated with small insects and zooplankters range from 0.1 to 100.
Larger organisms have usually larger Reynolds numbers. A more ap-
propriate formula for large Reynolds numbers woul d be quadratic with
respect to t he velocity (Vogel (55); Al exander ( 5 6 , 5 7 ) ) . For simplicity
in mathematical analysis we assume t he linear law, but when we compare
t he t heory with data, we estimate t he value of k~ in such a way that t he
frictional coefficient depends upon t he mean speed of organism duri ng
t he groupi ng mot i on (see Chapt er VII).
Thi r d, t he i nt ernal force depends only on t he relative distances be-
t ween two individuals; its value is invariant under a translation of t he
origin of t he coordinate system.
F ~ = m l A i j ( r ~ - - r i ) = m ~ A i j ( r / - r ~ ' ) (V. 5)
For groupi ng animals t he i nt ernal forces are generally attractive in na-
t ure except for being repulsive in t he very vicinity of individuals. I f t he
i nt ernal force were a t ype of Newt oni an gravitational attraction, a uni f or m
spherical distribution of particles woul d attract an i nt ernal point as t hough
all t he masses inside t he point were concent rat ed at t he cent er of sphere,
and t he resultant attraction woul d be directly proportional to t he distance
from t he cent er (Ramsey ( 5 8 ) ) .
By a mere analogy to t he Newt oni an attraction we speculate that t he
resultant internal attraction of grouping animals produces, on t he aver-
age, a centrally attractive force acting on an individual. Partly because
of a limited number of individuals in grouping and partly because of
nonuni formi t y in their spatial distribution, t he resultant internal force
on an individual also produces a fluctuating force whi ch arises chiefly
from chance encounters with individuals who happen to be in t he neigh-
borhood of t he individual in question. We will t reat this fluctuating force
as a random variable. Therefore
~. F ~ = m , Y~. A ~ j ( r / - - r , ' ) = m , f ~ ( r , ' ) + m i Z ~ ( t ) (V. 6)
J J
where m ~ f ~ ( r ~ ' ) is t he centrally attractive resultant force and r a Z e ( t ) is t he
random force. The former force is satisfied by
r , ' f , ( r , ' ) = 0 (V. 7)
22 a. OKUBO
We also assume that the resultant internal grouping force is random
as the positions of grouping animals varies with time in a complex man-
ner. Combining this resultant internal grouping force and the random
force in (V.6), we define m~W, to be the random resultant force operating
on z-animal.
2. Basi c Dy n a mi c a l Model s f o r t he Mai nt enance o f Groupi ng and Cor-
rel at i on Funct i ons
In summary the equation of the relative motion can be expressed as
~ ' ~ + k i ~ - f ~ ( r ~ ) = w~(t) ( v . 8)
where the frictional coefficient is assumed to be the same for all the in-
dividuals, and the external force is momentarily neglected. We will con-
sider only the radial motion for r = [r[, so that
r + k r - f ( r ) = W( t ) (V. 9)
where for simplicity the subscript sign has been suppressed. Eq. (V.9)
is subject to the initial conditions that
at t = 0, r = ro, r -= vo (V. 10)
Since the non-random force f is attractive in nature, we may express
a s
f (r) = --wZr--b(r) (V. 11)
where w is the frequency of harmonic component of the attractive force,
and b(r) is the acceleration due to anharmonic component of the attrac-
tive force. Except in Chapter V.5 our main concern is the dynamics of
group maintenance around a single attractive center, so that b(r) is an odd
function in r;
b(r) = - - b( - - r ) (V. 12)
Eq. (V.9) combined with (V.11) becomes
r + k r +~oZr +b(r) = W( t ) (V. 13)
Furthermore we assume that the random acceleration W( t ) represents a
stationary process with zero mean
W( t ) = 0 (V. 14)
For the nonlinear an_harmonic function b(r) it is not possible to ob-
tain an exact analytical solution to (V.13). One approach that has been
used to obtain approximate solutions of t he oscillatory behavior of (V.13)
is to linearize it, i.e., to replace (V.13) by an equivalent harmoni c oscillator
wi t h frequency we.
r' +kr +eoe2r = W(t) (V. 15)
The frequency we is chosen in such a way t hat t he error made by t he
repl acement of w' r+b(r) by we2r is mi ni mi zed (Bulsara et al. (59); Caughey
(60); Spanos and Iwan (67); Zi mmer man (62); Indi ra et al. (63); Crandall
(64)). The solution of (V.15) is t hen used as an approximate description
of t he oscillator (V.13), and statistical characteristics such as correlation
functions are calculated from t he approximate solution.
A st andard met hod is available to solve (V.15) for r and v = r given
t he initial conditions (V.10). For example, see Uhl enbeck and Ornstein
(65) and Chandrasekhar (66). Instead, we will calculate directly t he cor-
relation functions r ( t ) r ( t +v ) and v ( t ) v ( t +r ) .
Consider (V.15) as referring to times t = t r and t = W and correspond-
ingly express r ( t l ) =r ' , W ( t ' ) = W ' and r ( t " ) =r " , W ( U ) = W 'I. Multiply-
ing bot h equations t oget her and averaging t he resultant equation, we
obtain
( d 4 / d a d a \ d ~
_ _ - - - - k2.
+
d d d 2 d 2
+ k)e~(dt ' + dt " )+' e~( dt '2 + d t "2 ) +c~4} r' r" = WT-W-n (V. 16)
Since Eq. (V.15) states t he linear relationship bet ween r(t) and W( t ) , it
is clear that a stationary random variation of W( t ) will produce a sta-
t i onary random variation of r(t ) after t he effect of initial conditions de-
cays, so that rt r I/ and W t W ~ are functions of v = t ~ - - t t only. The as-
sumpt i on of stationarity also allows t he differential operator on t he
stationary functions to change
d d d d
d t ' - dr a n d - d U = d-~-"
Consequently (V.16) results in
~ + c 0 e ~) - = Wr-W " (V. 17)
Define t he Fouri er t ransform of rt r ~ and W t W ~ by
24
~
oo
CO(f) = (2n)-* r-r~r"(r)e-~f~dr
V( I ) = (2u)-* WT Wa ( r ) e - ' f : d v
- - oo
A. OKUBO
(V. 18)
(V. 19)
r e s pe c t i ve l y. Ta k i n g t h e F o u r i e r t r a n s f o r ms o f (V. 17) a n d u s i n g (V. 18)
a n d (V. 19), we o b t a i n
q)(f ) = {(f 2--we~)z+k2 f Z } - l ~ ; ( f ) ( v. 2o)
No t e t h a t Ma t s u o ( 67) a p p l i e d a p r o j e c t i o n o p e r a t o r me t h o d t o t h e n o n -
l i ne a r s y s t e m (V. 13) wi t h b ( r ) = r 3 a n d s h o we d t h a t t h e me t h o d p r o v i d e s
t h e s pe c t r a l r e l a t i o n wh i c h is t h e s a me as (V. 20).
T h e i nve r s e F o u r i e r t r a n s f o r m of (V. 20) gi ves
Si nc e
a n d
wh e r e
f ~ o o ~ ; ( f ) e , f ~ d f = W ' W " ( ~ )
= ~l(ko)eO)l)e -1/2<~1 sin (o~ 1 I v I +s i n -1 oJU% )
(V. 21)
(V. 22)
(V. 23)
Z( r ) = W( t ) W( t + r) (V. 26)
We n o w s p e c i f y t h a t W( t ) is s t a t i ona r y Ga us s i a n, a n d t h e c or r e l a -
t i o n coef f i ci ent p ( r ) = Z ( r ) / W 2 f ul f i l l s t h e f ol l owi ng f u n c t i o n e q u a t i o n
p( t a- - t l ) = p( t a- - t 2) p( t 2- - h) , t a >t 2 >t , (V.27)
T h e onl y n o n - s i n g u l a r s ol ut i on o f (V. 27) is
wh e r e
a,, - (~o~2-k~I4)*/2 (V. 24)
t h e us e o f Pa r s e va l ' s f o r mu l a t o (V. 21) yi e l ds
r ( t - ~( t +r ) = (2ko)~oJl) -I Z ( v - ~ ) e -1/2<~' si n (oJ I 1~ I
+ s i n -1 o)l/o)~)d~ (V. 25)
D Y N A MI C AS P ECTS OF A N I MA L G R O U P I N G 2 5
p ( r ) = e - ~
o r
Z( r) = W2e- r ~ (V. 28)
where i "-1 represent s a charact eri st i c t i me of correl at i on.
A l i mi t i ng case of (V.28) is of consi derabl e i nt erest . Suppose t hat
).-1 __~ 0 or 2" --9 oo i n such a way t hat
lira W~r-1 = B (a finite positive number) (V. 29)
T~co
Not e t hat B has di mensi ons of L 2 T -~, i. e. t he rat e of di ssi pat i on of t he
power of W( t ) . For t hi s l i mi t i ng case,
W( t ) W( t + r) = l im B r e - ~ = 2B~(v) (V. 30)
~'---~oo
wher e O(r) is a Di rac f unct i on.
The cor r espondi ng Four i er t r ans f or m of Z(r) becomes
~b(f) = B/~r (V. 31)
whi ch means t hat W( t ) is whi t e noise.
Subst i t ut i on of (V.30) i nt o (V.25) yi el ds
r ( t ) r ( t +r ) = B ( k o ) e O J l ) - ~ e - 1 1 ~ sin (o)ir +s i n -1 collo),), r > 0 (V. 32)
The vari ance of t he di spl acement is f ound f r om (V.32) by set t i ng r = 0 :
7~ = n / (k~o, ~) (V. 33)
t hat is, t he vari ance approaches const ant as t --~ ~ .
I f we define t he Four i er t r ans f or m of v t v ~t by
X2(f) = (2z) -~ v' v"(r)e-~S~dr (V. 34)
~oo
i t is rel at ed t o t he spect ral densi t y of r ( t ) by
~2(f) = f z g ) ( f ) (V. 35)
o r
d 2
v ( t ) v ( t + ~ ) - - - 2 ; 2 - ~ r ( t ) r ( t + ~ ) ( V . 3 6 )
Di f f er ent i at i ng (V.32) t wi ce wi t h respect t o r and usi ng (V.36) we
obt ai n
26
and
v ( t ) v ( t + r ) = ( B / k ) e - 1/ z ~ (cos w l v - k / 2 o ) l sin o)lv),
v 2 = B / k
A. OKUBO
r>O (V. 37)
( v . 38)
f : R ( r ) d v = 0 40) (V.
f : r R ( v ) d r = -- (1/~o~ 2) (V.41)
Subst i t ut i ng (V.38), (V.40), (V.41) into (III. lO) we obt ai n as t --* ~o
r ~ = B/ ( k oJ ~)
whi ch is only a rediscove W of (V.33).
The behavior of R ( v ) against r looks as illustrated in Fig. 3. For
o)~2>1/4k 2 (underdamping) t he correlation coefficient oscillates about
t he zero value wi t h decaying ampl i t ude (Fig. 3a). I n physical t erms t he
individual mot i ons appear t o resemble a random pendul um-l i ke mo-
t i on about t he origin. In midge swarming, animal individuals fall back
and fort h t hrough t he swarm center (see Chapt er VII). For o)e2<1/4 k 2
(overdamping), on t he ot her hand, t he frictional force dominates t he at-
tractive force and t he correlation coefficient becomes aperiodic, i . e . R ( v )
decreases monot onousl y, crosses t he zero line and goes into t he region
of negative value and approaches t he zero line asymptotically (Fig. 3b).
I t does not oscillate about zero.
I f we consider a limiting case of O)e -+ 0, i . e . no attractive force, our
expectation is to reveal pure diffusion. As a mat t er of fact,
lim R ( v ) = e -k~
oJ--+O
We t hus obtain
f
R d r = 1 / k
0
I t is found
Hence t he velocity correlation coefficient R(v) is given by
R(r) = e -1/2~ (cos o)lv--k/2col sin colv), v >0 (V. 39)
It can be seen t hat this correlation coefficient represents t he feature
of grouping. To this end, comput e
R ( r ) d v and r R ( r ) d v .
0 0
DYNAMIC ASPECTS O F ANIMAL GROUPING 27
f o r 1/ k
R d v = 2
The r e f or e , as t --~ co
"~ = 2v-~k-it = 2B/ k 2t
wh i c h i dent i f i es t he di f f usi vi t y D by
D = B / k 2 (V. 42)
So f ar we have deal t pr i ma r i l y wi t h an equi val ent l i near dynami cal
mo d e l f or ani mal gr oupi ng. A var i et y of nonl i near mode l s ma y be con-
s t r uc t e d. Th i s i ncl udes a quadr at i c f r i ct i onal l aw a nd a nha r moni c at -
t r act i ve f or ces, e. g. by r et ai ni ng hi ghe r - or de r t e r ms i n t he power seri es
expans i on of f ( r ) . Some ma t he ma t i c a l me t h o d s expl or ed i n t he nonl i near
dyna mi c s ys t ems and nonl i near osci l l at i on pr obl e ms are available (for
r ef er ences, see Ma t s uo (67); Boyd (68); Wu and Li n (69); Mor i guc hi
a nd Na k a mu r a (70); Cr andal l (77); Mo r t o n and Cor r s i n (72); Hs i e h (73);
Nayf eh and Mo o k (74); St oker (75); Mi nor s ky ( 76) ) . We are seeki ng f or
onl y t h e mode l s t ha t p r o d u c e an as ympt ot i cal l y st abl e fi ni t e vari ance,
i . e. l i m r ~ conver ges t o a fi ni t e val ue, whi c h can be expr es s ed i n t e r ms of
t, --~co
t he pa r a me t e r s pe r t i ne nt t o t he model . I nt ui t i ve l y we expect t ha t by and
l arge t he de t e r mi ni s t i c f or ce mu s t be at t r act i ve i n nat ur e, especi al l y at
l arge di s t ances f r o m t h e cent er of gr oupi ng.
We finally cons i der t he cor r el at i on f unc t i on of accel er at i on
a' a " = a( t ) a( t + r) (V. 43)
and defi ne t he spect r al f unc t i on by t he Four i e r t r a ns f or m of (V.43)
F
o( f ) : (2~) -1 a' a"(r)e-' =dc ( V. 44)
- - t o
whe r e s t at i onar i t y is as s umed.
By anal ogy wi t h (V.35) and (V.36) t he cor r el at i on f unc t i on and
spect r al dens i t y of a(t ) are r el at ed t o t hos e of v(t) by
d ~
a( t ) a( t + ~) = - - ~ v ( t ) v ( t + r) (V. 45)
w( f ) = f 2~2( f ) (V. 46)
Su b s t i t u t i o n of (V.37) i nt o (V.45) yi el ds
a( t ) a( t +~) = B@o~2-k~)/ke - 1 / ~ [cos o~lr
- {k(4~o12-oJ~2)/2oa(o4~-le~)} sin oJ1r] (V. 47)
28 A. OKUBO
and
a 2 = B(o), ~-k~)/ k (V. 48)
3. Di st ri but i on o f Organisms wi t hi n a Group: Fok k e r - Pl anc k Equat i on
I n order to bui l d a mathematical model appropriate for t he spatial dis-
t ri but i on of grouping organisms, we will make use of a Fokker-Planek
equation derivable f r om t he basic dynamical equation, Eq. (V.9).
Let p~(x, v, t) be t he probability density funct i on t hat an organism is
found in t he phase space (position x and velocity v) of t he range (x, x +
dx) and (v, v + d v ) at t i me t. Assuming that t he process described by t he
probability density funct i on P2 is Markovian, we obtain (Risken (77);
Van Kampen (78))
at ~=1 ~ \ ~ x O~v ] {M~(~)P2} (V. 49)
where Mx v (n) denotes t he rate of change of t he nt h moment in t he phase
space. Eq. (V. 49) is referred to as t he Kramers-Moyal expansion.
A formal solution of (V.49) subject to t he initial condition t hat P2
(x, v, 0) =/ 2( x, v) is obtained by (Risken (77))
p 2 ( x , v , t ) = 1+ o dh o dt2"'" o duL~zM(x,v, tl)
LEa(X, v, t 2). . . L~e(x, v, t~) l f 2(x, v)
2
where
t he Kramers-Moyal operator.
I f t he Kramers-Moyal expansion (V.49) stops after t he second-order
t erms, we get t he Fokker-P1anck equation or forward Kolmogorov equa-
t i on
a t Ox (M~p~) - (M~p2) + ( M~p2)
+ 2 ~ v (NIx~p2) + ~ (M~p~) (V. 50)
Not e t hat t he Pawula t heorem states that if t he expansion (V.49) does not
stop after t he second-order t erms, it must contain an infinite number of
t erms.
I n ( V. 50) Mx, My, Max, Mxv, a n d Mvv c a n b e e v a l u a t e d f r o m t h e b a s i c
s e t o f e q u a t i o n s f or x a n d v. Be i ng r e wr i t t e n Eq. ( V. 13) r e a ds
dx/dt = v (V. 51)
dv/dt = - k v - o~2x - b( x ) + W(t) (V. 52)
wh e r e we a s s u me t ha t b(x) is an o d d f u n c t i o n i n x a n d W(t ) r e p r e s e n t s a
Ga u s s i a n r a n d o m va r i a bl e wi t h z e r o me a n a n d a c or r e l a t i on,
w ( i i = o
W(t) W( t + r ) = 2Ba(r) / (V. 53)
wh e r e B is t h e i n t e n s i t y o f t h e va r i a nc e of W(t ) as gi ve n b y (V. 31).
F o r t h e n o n l i n e a r L a n g e v i n e q u a t i o n s ( . 5 1 ) a nd ( V. 52) wi t h (V. 53)
we c a l c ul a t e t h e Mrs b y
M~(x, v, t) = l i m "x(t+At)--x(t)
zlt--*O A t
M~(x, v, t) = l i m v( t +dt ) - - v( t )
3t--,o A t
2 d ~ ( x , v , t) = l i m {x(t+dt)- - x(t)}2
At~O At
Mx~(x, v, t) = l i m {x(t + At)--x(t)] {v(t + At)--v(t)}
At~O At
Mw(x, v, t) = l i m {v(t+At)--v(t)}2
zt~o At
(V. 54)
I t r e s u l t s ( Ar n o l d ( 79) ; S o o n g (80))
M x ~'O l
M~ = --kv--~o~x--b(x)
M~ = 0
/
M~ = 0 !
)
Mo~ = 2B
( v . 55)
T h e F o k k e r - P l a n c k e q u a t i o n ( V. 50) i s t h e n wr i t t e n as
@2 @2 a 2 a2P 2
Ot - - Vax+~- vv [(kv+J x+b) pd +B. Ov z
( v . 56)
wh i c h is o f t e n cal l ed t h e Kr a me r s e q u a t i o n ( Kr a me r s ( 81) ; Vo l l me r a n d
Ri s k e n (82)). I t i s n o t e d t h a t f or t h e s y s t e m ( V. 51) - ( V. 53) al l h i g h e r c oe f -
30 A. ommo
ficients of M~)(v_>3) are zero and t herefore t he Fokker-Planck equation
(V.50) becomes t he exact expansion of t he Kramers-Moyal equation.
Ther e are two i mport ant marginal probability density functions de-
fined by
p ~ ( x , t / = ~ _ p 2 ( x , v , , ) d v ( v . 57)
p#, 0 - ~o2~( x, v , , > a x ( v . 58>
whi ch are t he probability density funct i on of position and that of velocity,
respectively.
I n view of t he properties that P2 is an even funct i on of x and b ( x ) is
an odd funct i on of x, integration of (V.56) over x leads to
= k ~ 7 ( v p ~ ) + B a2p* (V. 59)
at av~
where p = ( x , v , t ) converges to zero fast as Ix] -+ co. The solution of (V.59)
subject to p v ( v , 0) =a(v--v0) is found to be
p v ( v , t ) = - k j exp (V. 60)
2roB {1 - exp (--2kt)} ~ J
For steady-state or t>>k - 1 Eq. (V.60) becomes
p~*(v) = (k/2~rB)l/Ze - ~ / 2 B (V. 61)
whi ch is t he Maxwellian velocity distribution.
The equilibrium (steady-state) solution of (V.56) is given by
p 2 * ( x , v ) = Co e x p { - - - ~ - ~ - + - - f x 2 + J b ( x ) d x ) ) (V. 62)
where
k v 2 oJ~
(V. 63)
Solving exactly (V.56) with arbitrary b ( x ) is a very difficult task. The
only known exact solution of (V.56) subject to p ~ ( x , v , O ) = f o ( X , v ) is for
t he harmoni c force, b_=0. For high friction t he Kramers equation is
reduced to t he Smoluchowski equation whi ch is a special Fokker-Planck
equation for t he density funct i on for t he position only, i . e . p x ( x , t ) .
The limit of large k should result in very rapid relaxation of t he
mo me n t u m equat i on (-7.52) t o a quasi - st at i onar y st at e i n whi c h d v / d t = O .
Hence, we as s ume t ha t f or l arge e nough k, (V.51) and (V.52) are r e duc e d
t o
d x / d t = - k - l ( o J 2 x + b(x) ) + k - l W (t) (V. 64)
T h e c or r e s pondi ng e qua t i on f or t he pr obabi l i t y dens i t y f unc t i on p x ( x , t )
becomes t he Smol uc hows ki equat i on:
ap~ _ k - l ~ {(JZx + b ( x ) ) az
+ B k - I T ~ x ~ p ~ (V. 65)
Ga r di ne r ( 8 3 ) p r o p o s e d t he f ol l owi ng equat i on f or p x ( x , t ) whi c h is val i d
not onl y f or l arge t b u t also for smal l t :
~P~ ~ I f - - ~ ] f t e x p ( k ( t ' - t ) ) p ~ ( x , t ' ) d t ' (V. 66)
a t - ~ x ( w e x + b ( x ) ) + B J o
T h e ker nel exp ( k ( t t - t ) ) is si gni fi cant l y di f f er ent f r o m zero onl y f or it t -
t l N k - 1 , a nd on a t i me scale mu c h l arger t h a n k - i , (V.66) is agai n ap-
pr oxi ma t e d by t he Smol uc hows ki e qua t i on (V.65).
Van Ka mp e n ( 7 8 ) s ol ved t he Kr a me r s equat i on (V.56) appr oxi mat el y
f or l arge k by means of an expans i on i n power s of k -1. I nt e gr a t i on of t he
r e s ul t a nt s ol ut i on over v yi el ds t he appr oxi mat e equat i on f or p x ( x , t ) , whi c h
satisfies t he Smol uc hows ki equat i on wi t h cor r ect i on t e r ms of or de r k -2,
/.e.
opx
k - l f - - ~ { ( o J 2 x + b ) p x } - B k -~ O2P~ = 0(k -2) (V. 67)
~t ax 2
I n vi ew of t hi s l i mi t a t i on we are pr i mar i l y i nt e r e s t e d t o deal wi t h t he
st eady- st at e n u mb e r - d e n s i t y di s t r i but i on of gr oupi ng or gani sms, p x * ( x ) by
s et t i ng ~ p x / ~ t - ~ 0 i n ( V . 6 5 ) .
f oj2 ~ r b ( x )
p ~ * ( x ) = po e x p l - ~ x - - J - - B - d x ) (V. 68)
wher e Po is t he dens i t y at t he gr oup cent er . T h e non- Ga us s i a n dens i t y
di s t r i but i on arises f r om t he a nha r moni c at t r act i ve force. Si nce x b ( x ) > O ,
t he dens i t y di s t r i but i on t e nds t o be pl at ykur t i c, i . e . t he kur t osi s of t he
di s t r i but i on is less t h a n 3.0.
For a l i near s ys t em, i . e . b - O , Bul sar a e t a l . ( 5 9 ) de mons t r a t e t hat t he
l ong t i me di s t r i but i on of di s pl a c e me nt s is gi ven by (V.68) i r r espect i ve
of t he val ue of k. Pr obabl y i t is qui t e r easonabl e t o expect t ha t t he l ong
32 a. O K U B O
t i me solution of p~(x, t) which woul d be obt ai ned by integrating p~(x, v, t)
of (V.56) over v will converge to (V.68).
4. Dy nami c al Model s f o r the Format i on o f Group
Model i ng t he process of group format i on is much more difficult t han
t hat of group maintenance simply because we deal essentially wi t h a
transient (time dependent ) phenomenon. The group formation process
can conveniently be classified into two categories: one is a density-in-
dependent process and the ot her is a densi t y-dependent process.
I n a densi t y-i ndependent process, animal individuals are i ndepen-
dent l y attracted to a source of attraction or core of grouping, and even-
tually a group forms. An example of this sort is t he aggregative behavior
of male insects in response to female pheromones (Bell and Card6 (84)).
Anot her example is a group formation around a confined bait; amphi pods
are at t ract ed to a bait and t hey swarmed around it (Ingrain (85)).
In a densi t y-dependent process, on t he other hand, individuals al-
ready in aggregation can influence (enhance or inhibit) t he attraction of
individuals at large. A densi t y-dependent process is usually preceded
by an initial stage which is i ndependent of density in a sense that indi-
viduals encount er at random t o form an initial group of small size.
Mut ual communi cat i on bet ween animals aggregated and t hose at
large makes t he grouping process density dependent . Some possible
machanisms by which t he animals might communi cat e are chemosensory
by means of pheromones and mechanosensory by pressure changes. A
good example of t he densi t y-dependent aggregation is seen in a massive
attack of pine beetles to host trees (for details consult Chapt er VI I . l ) .
The random encount er process is model ed by Gerri t sen and Striekler
(86) for zooplankton and by DeVita et al. (87) for arthropods. In their
model of random encount er Gerri t sen and Strickler (86) assume that 1)
t he animals are considered dimensionless poi nt s in space, 2) t he animals
are randomly distributed, 3) t he animals are moving in random direc-
tions wi t h a mean speed a, and 4) an encount er radius Re is assumed
constant in all directions.
The encount er rate Z of an animal wi t h its conspeeific individuals,
or t he total number of individuals entering t he encount er radius per uni t
t i me is given by
Z = 4/3~rR~2m2 (V. 69)
where n is t he mean density of conspecifie individuals.
DeVita et al. (87) developed an encounter model in two-dimension
space similar to that of Gerritsen and Strickler and compared their model
with field observation on arthropods.
The initial phase of group formation may also be modeled in a way
similar to Anderson (88). Namely, the change with time of the number
of individuals N( t ) in a group is determined as the difference between
the rate that individuals enter the group, which is essentially given by
(V.69) and thus considered constant independent of N without communi-
cation and, the rate that individuals leave the group, which is propor-
tional to N. We thus have
d N/ d t = a - - b N (V. 70)
where a and b are positive constants. Possible stochasticity in a and b
may be incorporated (Anderson (88)). The solution of (V.70) subject to
N=I (grouping around a single individual) or N=0 (grouping about
an center of attraction) at t =0, describes an asymptotic approach to a
stable equilibrium Ne =a / b with a rate of b -1.
This equilibrium state would not be realized, however, if a certain
density-dependent process emerges as grouping progresses. If chemosen-
sory is assumed as the mechanism of animal communication, we can
construct a mathematical model for the enhanced (or accelerated) group-
ing process that leads to a massive aggregation.
First consider diffusion of a chemical, e.g. pheromone, from a spher-
ical source of radius R( t ) . The equation of diffusion of the chemical is
given by
a C_ D ~ (r ~aC~ (V.71)
at r 2 J r \ ~-r ]
where C(r, t) is the concentration of chemical as a function of the distance
r from the source center r =0 and time t, and D is diffusivity. Eq. (V.71)
is subject to the initial and boundary conditions:
at t = to, C = Co(r) (V. 72)
at r = R( t ) , - - D3C/ ar = Q(t)/4rcRZ(t) (V. 73)
with Q( t ) being the rate of release of chemical, i.e. the source intensity.
If we regard the source as a group of animal, in which N( t ) is the total
number of individuals, q is the emission rate of chemical per individual,
and v is the mean volume occupied by one individual, then
O(t) = qN(t) (V. 74)
34 A. OKUBO
4/37:RS(t) = v N( t ) (V. 75)
We next consider the advection (chemotaxis)-diffusion equation
for the animal at large. Let S ( r , t ) , u, and K be the concentration, advec-
tion velocity and diffusivity of animal, respectively. The equation for S
reads
3 S 1 O ~ 3--~-(r2KOS~ (V. 76)
at - r 2 Or (r~uS) + Or \ Or ]
where the chemotactic flow u depends upon the magnitude and gradient
of the chemical (Keller and Segel ( 33) )
u = f ( C , 3C/Or) (V. 77)
The influx of animal through the group boundary is given by
J ~ = 4 , z R~ ( - - u S + K OS/Or)r=R (V. 78)
which is equal to the rate of change of grouping animal number, so that
d N / d t = 4 z R ~ ( - - u S + K aS/Or)r=R (V. 79)
The set of Eqs. (V.71) through (V.79) coupled with (V.70) consists of the
basic mathematical model for the grouping process.
The nonlinearity of the model equation prevents us from obtaining
an analytical solution. Instead we may deal with a simplified model under
the assumption of quasi-steady state in the concentrations of chemical
and animal density (Okubo and Anderson ( 89) ) . Thus the quasi-steady
state concentration of chemical is given by
~!~ C( r, t) = Q(t )/ 4:rDr = q N (t)/4zvDr (V. 80)
Let C* be the threshold concentration for chemosensory response. Then
the effective radius r* for chemotaxis is obtained by
r*(t) - - qN(t )/ 4~rDC* (V. 81)
For quasi-steady state the influx of animal (V.78) is determined from
the following relation
J~, = 4 ~ R~ ( - u S +KOS / Or ) ~ =R = 4 ~r * ~( - u S +KOS / Or ) r =T, (V. 82)
We further assume that
( - u S + K aS/ar)~=~. = ~So (V. 83)
where , is t he effective velocity of chemotaxis at t he t hreshol d concentra-
tion and So is t he mean background densi t y of animal.
The change wi t h t i me of t he number of animals in groupi ng is t hen
given by
d N/ d t = 4zcr*2~So = 4z:(q/4~rDC*)2~SoN 2 =_ r N 2 (V. 84)
wi t h
7 - q~So/4~rD2C*~ (V. 85)
The solution of (V.84) subj ect to N =N o at t =t o is obtained by
N( t ) / No = {1 - T N o ( t - to)} - i (V. 86)
Thi s solution describes t he avalanche of aggregation in such a man-
ner t hat t he group reaches an infinite size within a finite time interval
given by
tc ~ t - t o = (TNo) -1 = 4~D2C*2/q~vSoNo (V. 87)
The knowledge of t he pert i nent parameters woul d enable us to es-
t i mat e this critical time. The present ed model may also be applicable to
acoustic communications wi t h mi nor modifications.
I n normal environmental conditions there exists a mean flow 5 (wind
or current) so that t he substance released forms a pl ume elongating in
t he direction of mean flow rat her than a cloud of spherical symmet ry.
Hence t he effective region of t he chemical communi cat i on will somewhat
be modified by t he presence of flow. Tur bul ence in t he environmental
fluid will also affect t he region. For more details t he reader should con-
sult Okubo (26), Csanady (90), Sut t on (97), Pasquill (92), Fischer et al.
(93), and Harri s (94) among others.
As an example of modification take a case of cont i nuous emission
in a laminar flow field. The maxi mum dimensions of the pl ume charac-
terized by t he t hreshol d concent rat i on are given by L x =Q/ ( 2 z c C* D) in t he
direction of flow and Lm=( 2Q/ eTrC*Ft ) 1/2 in a plane perpendicular to t he
flow. We t hus obtain t he influx of animal by chemotaxis as
J ~ ec vLxLmSo = 71, SON 3/2
wi t h
Therefore
71 = (q/C*) a/~D~l/~
]V = ?,lvSoNa/Z
which should be compared to (V.84).
( v . 8s)
( v . 89)
( v . 90)
36 A. OKUBO
Generally speaking in t he diffusion of chemicals t he densi t y-depen-
dent attraction model may be expressed by
.N = a~SoN ~ (V. 91)
where t he parameters a and m depend upon t he model employed. The
solution of (V.91) subject to N = N o at t =t o yields
N( t ) / No = {1- ( m - 1)No' ~-l avSo(t - to)] -i/(~-1~ (V. 92)
For r n >l t he population reaches an infinite size within a finite interval
of t i me given by
t~ = [( m- 1) No~- l a~So}- 1 (V. 93)
A different approach to modeling t he process of densi t y-dependent
aggregation was used by Kawasaki (95), Mi mur a and Yamaguti (96), and
Alt (97), who i ncorporat ed long-range mut ual attraction directly into t he
advection-diffusion equation for animal concentration (V.76). Thus,
Kawasaki (95) proposed an adveetive flow of animal as
u(x) = f k ( x ' ) S ( x - x ' ) d x '
(V. 94)
wi t h t he convolution kernel k such that k ( # ) represents a weighting func-
tion vector directed t oward # = 0 . A special case where
k( x' ) = Uo~'(x') (V. 95)
reduces t he advective flow to
u(x) = uoVS (V. 96)
t hat is, individuals are attracted t oward t he region of higher density.
Models of population dispersal when t ransport of population is
long-range have been developed by Levin (98) and Levin and Segel (99)
in predat or-prey interactions.
5. Model s f o r Grouping around Mul t i pl e Centers
Thus far our dynamical models are limited to t he case of grouping around
a single attractive center. I n reality t here may be two or more grouping
centers, and t he organisms may shift t hei r locations from one cent er to
another. As far as t he dynamics is concerned, grouping is maintained by
t he balance bet ween attraction of individuals t oward t he group and dif-
fusion due to random motion of individuals. Thus an individual inside
DYNAMIC ASPECTS OF ANI MAL GROUPING 37
a group may be considered to be held by a potential barrier and to per-
form oscillations about a stable equilibrium. The random mot i on may
"act i vat e" t he individual t o such an extent that it overcomes t he barrier
and leaves the group only to j oi n a new group or ret urn to the original
group.
For a multiple cent er probl em we still use Eq. (V.9) as a basic dy-
namical equation by appropriately specifying t he force function. Thus,
in a one-dimensional case, the force has a potential:
f (x) = -- dCO (x)/dx = - ~b, (x) (V. 97)
and t he dynamical equation reads
2 +k 2 +~' ( x ) = W( t ) (V. 98)
Not e t hat the potential funct i on includes not only the effect of internal
forces of t ype (V.5) but also t he effect of external forces of t ype (V.3)
We will deal onl y with the high-friction limit (see Chapt er V.3).
Then the density funct i on px( x, t ) =p ( x , t) satisfies the Smoluchowski
equation
~p a o ~
3t - ~x ( U' ( x ) p ) +D o (V. 99)
where U( x ) - q b ( x ) / k : viscous potential (Van Kampen (100)) and D=_B/ k :
diffusivity.
We now consider t wo centers of grouping characterized by a bistable
potential funct i on U( x) as shown in Fig. 5. Ther e are t wo minima at
x =x 1 and - - x 1 and in between, a local maxi mum at x=0. For a few special
forms of t he bistable potential Eq. (V.99) has been solved explicitly. I n
general no explicit solution is known apart from the steady-state distribu-
tion. Put t i ng 3p/ 3t =O (V.99) can be integrated to find the steady-state
solution
p, ( x) = d exp {-- U( x) / D} (V. i00)
where A is det ermi ned from the normalization condition
f
~oops(x)dx = 1.
The steady-state distribution is schematically shown in Fig. 5 for the
bistable potential, ps(x) has t wo maxima at x = x 1 and - - x 1 and a mi ni mum
at x=0. The grouping animal is most likely to be found at x = x 1.
38 A, OKUBO
/ "
" " \ p~ ()
V" /
/ k_/ ' ,
\..~o
/ ' 1
/ ~ ' k k . k . /
' . ,
/ k
I
--X1 0 X1 X
Fig. 5. Bi st abl e potential function U(x) vs. distance x and steady state dis-
tribution of organisms ps(x)
T h e smal l er t he di ffusi vi t y, t he mor e c onc e nt r a t e d gr oupi ng is at x = ___x v
Pu t t i n g p ( x , t ) = f ( x ) e - ~t , we can r educe (-7.99) t o an ei genval ue
pr obl e m.
d 2 f d ,
O-dZx2+~-~x(U ( x ) f ) = - 2 f ( x ) (V. 101)
Wh e n t he mi n i mu m ei genval ue 2o is zero, t he st at i onar y s ol ut i on p s ( x )
exists and t he s ol ut i on of (V.101) is gi ven by
p ( x , t) -= p , ( x ) + ~, a~f~(x)e-~, t (V. 102)
n=l
wher e 0 < 2 1 < 2 2 < . . . and an are d e t e r mi n e d f r om t he i ni t i al condi t i ons.
Fo r t he s ymme t r i c bi st abl e pot ent i al s hown i n Fi g. 5, t he first ei gen-
f unc t i on f l ( x ) is a nt i - s ymme t r i c about x = 0 and t h e s econd ei genf unc
t i o n f 2 ( x ) is s ymmet r i c. T h e s econd ei genval ue 22 is mu c h l arger t h a n 21.
Cons e que nt l y f or t>>22-1 t he s ol ut i on (V.102) is a ppr oxi ma t e d by
p ( x , t) ---- p , ( x ) + a l f i ( x ) e - h t (V. 103)
T h e next pr obl e m of i nt er es t is h o w l ong i t t akes f or a gr oupi ng
i ndi vi dual t o s wi t ch f r om one cent er t o anot her . Cl earl y t he t i me of con-
cer n de pe nds pr i mar i l y u p o n t he he i ght of t he pot ent i al bar r i er rel at i ve
t o t he i ndi vi dual ' s r a n d o m mo t i o n char act er i zed by t h e di ffusi vi t y.
T h e pr obabi l i t y t ha t t he i ndi vi dual i ni t i al l y at x at t = 0 is st i l l i n
t he doma i n of pot ent i al a<x <_b is gi ven by
b !
f p ( x , t I x, O)dx' =_ G( x, t) a < x < b (V. 104)
I f T denotes t he t i me that t he individual leaves the domain (a, b), t hen we
obtain
b
Probability ( T > _ t ) = f ~ p ( x ' , t l x , O ) d x ' = G ( x , t ) (V. 105)
When t he dynamical process of grouping is homogeneous in time, we can
wri t e
p ( x ' , t I x , O) = p ( x ' , O I x , - t ) (V. 106)
so t hat p ( x t , t Ix, 0) is obeyed by t he backward Fokker-Planck equation
ap
- U ' ( X ) ~ x + D 3 2 P (V. 107)
8t 3x 2
Fr om (V.104) G ( x , t ) , too, obeys t he backward equation
3G
- U ' ( X ) ~ x + D f i ~ G (V. 108)
Ot 8x e
whi ch is subject to t he initial condition
G ( x , O) = 1 , a<_x<_b
(v. 109)
J
= 0, otherwise
Since G ( x , t ) is t he probability that T > _ t , t he mean first passage t i me
for t he individual initially at x is given by
v ( x ) = - - t 8 G / S t d t = G ( x , t i l t (V. 110)
0 0
Int egrat i ng (V.108) with respect to t from 0 to ~ and using (V.110) we
obtain
. , , , d r _ d~v
- - o ( x ) ~ f f x + O T Z x 2 = - 1 (V. 111)
For t he bistable potential shown in Fig. 5 t he mean first passage time for
t he individual initially at x = - - x a (left-side center) to cross over t he bar-
rier into t he ot her cent er is found by solving ( VAI l )
v ( - x , ) = D - l f _ x d y e x p ( U ( y ) / D ) f ~ oexp { - - U ( z ) / D } d z (V. 112)
I f t he potential barrier is large compared with D, t he funct i on
exp { U ( x ) / D } is sharply peaked at x=0, while exp { - - U ( x ) / D } is very
40 A. OKUBO
small near x=0. Therefore (V.112) is approximated by
f 0 f 0
v(--xl) ~ D -t dyexp [ - - U ( y ) / D ] dxexp { U( x ) / D] (V. 113)
- - Xl - - ~
Furt hermore if we approximate t he viscous potential near x = - - x l and
x = 0 by
U( x ) ~ U ( - x l ) - t - 1 / 2 U " ( - x l ) ( x x l ) 2, I x + x l l small
~- U(O)+ I/ 2U"(O)x~, Ix[ small
(V.113) is reduced to
v ( - x l ) ~ z c [ U~ ' ( - x l ) l Ur'(O) I } - t / ~e x p {[U(0)-U(--x~)]/D} (V. 114)
Ther e are a few special forms of the bistable potential whi ch provi de
exact closed-form expressions for the density function valid for all t i me
and also for t he mean first passage time. As an example we take
U( x ) = 1/ 2a(I x l - x ~ ) ~ , a > 0 (V. 115)
Thi s potential has two minima at x=++_xl and a maxi mum at x =0 ; t he
height of t he potential barrier, h = l / 2 a x~ 2. The eigenvalue probl em is
exactly soluble (see Banerjee e t al . ( 707) ) .
Suppose we place all the individuals at t he maxi mum point ( x=0) at
t =0 . I n t he limit of small diffusivity, the density funct i on can be expres-
sed by
p ( x , t) = c ( a / 2 K ) l / ~ e x p I--a{[ x l - - x l ( 1 - - e - ~ ) ~ / 2 K } ] (V. 116)
where
K = D(1--e -~'~t) (V. 117)
and c is a normalization factor. Thus t he animals begin to split t wo groups
of equal individuals, each group moves into its own potential domain, and
eventually settles to a Gaussian distribution around t he bot t om of t he
potential well. I n this limiting case the variance of animal positions is
given by a x 2 =- 2 Dt in the initial peri od and c r x 2=D/ a asymptotically.
The mean first passage time from x = - - x 1 to x = 0 is calculated from
(V.112) using (V.115).
v( - xl ) = (2~r)I/2a -1 f : / 2o; ' ~Xl d~e ' ~e r f c ( - ~) (V. 118)
Thus, if t he potential barrier is very large, i . e. (a/ 2D)l / ~xl >>1, (V.118) is
reduced to
~ ( - - X l ) ~-- ~ a - l e (~/2D~ :q~ (V. 119)
while if t he potential barrier is very small, i . e . ( a / 2 D ) l 1 2 x l < < l , (V.118) is
reduced to
r(--x~) --~ ( 2 z r ) l / ~ a - 1 ( a / 2 D ) l / ~ x l (V. 120)
As x I decreases for a fixed value of a, t he potential barrier becomes lower
and at t he same t i me t he two potential wells approach each ot her until
t hey collapse into a single well with t he disappearance of t he barrier.
An interesting case of escape times appears in connection with state-
dependent fluctuations or parametric noise (Horst hemke and Lefever
( 7 0 2 ) ) . The Langevin equation (V.98) contains t he st at e-i ndependent
noise W ( t ) . It is simply additive noise. Thi s kind of fluctuation does not
change t he local stability properties of t he system. Thus t he ext remum of t he
steady state density funct i on p s ( x ) always coincides with t he deterministic
steady state, i ndependent of t he intensity of t he noise. It only spreads t he
groupi ng individuals around t he bot t om of t he potential well, but does
not influence t he potential itself.
On t he ot her hand, t here may be a noise associated with t he para-
met er of t he system. For instance t he friction coefficient k in (V.98) may
contain stochasticity. The effect of this kind of noise (multiplicative noise)
depends on t he state of t he system, e . g . x or ~?. We will discuss how t he
escape process is modified in t he presence of multiplicative noise.
Consider t he following bistable potential
U ( x ) = --o~x~/2 + b x 4 / 4 , b>0 (V. 121)
and t he frequency o) contains noise as well as a deterministic element:
o) = f2o + n ( t ) (V. 122)
where 1 2 o > 0 is constant frequency and n ( t ) is a white noise such that
n ( t ) = O , n ( t ) n ( t + r ) = 2FS(r) (V. 123)
The dynami c equation in t he high-friction limit is expressed by
2 = Q o X + n ( t ) x - - b x S + W ( t ) / k (V. 124)
(see Guardia and San Miguel ( 7 0 3 ) ) . The Fokker-Planck equation as-
sociated with (V. 124) is wri t t en as
Op~t - ~xO { ( f ) o x _ b x a ) p } + ~ Z x z { ( D + F x ~ ) p } (V. 125)
42 A. OKUBO
The steady-state solution of (V.125) is
p,(x) = C( D+ Fx~) -1/~ exp {--E(x)/2F} (V. 126)
wi t h
E(x) = bx 2 - (Do + Db/F) in (x~+ D/F) (V. 127)
The steady-state distribution has t wo maxima for F<Do. For small F
t hese t wo maxima are close to t he deterministic equilibria: xl = - (~o/b) 1/~
and x2=(Do/b)l/2. As F increases t he t wo maxima approach each other
until t hey collapse into a single maxi mum for F=Do. For larger values of
F t he single maxi mum becomes more pronounced. The situation is com-
pletely different from t he previous case of t he additive noise. I n addition
t o t he disorganizing (or spreading) effect, whi ch is shared wi t h t he ad-
ditive noise, multiplicative noise can create new states of potential.
The effect of t he parametric noise is also seen in t he escape time.
Thus t he mean first passage time from xl to x 2 is calculated by
v(xl-.x~) = ;~l dx(D+ Fx2)-lp,-l(x) y~_o dyp~(y) (V. 128)
When t he intensity of t wo t ypes of fluctuations is small compared with
t he potential barrier, (V.128) is approximated by
v -~ (~/27cDo -t) exp [--[2o~/4bD{l+l/3(F/D)(Do/b)}] (V. 129)
for F/D<<I. The effect of muhiplicative fluctuations is a decrease in t he
escape time. Since a decrease of v is due to an exponential factor, t he
presence of small multiplicative fluctuations can dramatically short en the
escape time.
On t he ot her hand, t he mean first passage t i me for F/D>>I is given
by
v ~ ( ~/-ffmQo -1) {(Do/b)(F/D)}~ol 2F. exp (-Do/2F) (V. 130)
whi ch also decreases wi t h F.
The case of purel y muhiplicative noise (W(t)~-O) is analyzed by
Schenzle and Brand (104) and Graham and Schenzle (705) wi t h t he same
dynamical equation as (V.125). These investigators, too, demonst rat e t he
t hreshol d intensity of t he multiplicative noise, F t h r e s h o l d ~ - - - ~ Q o . For F<Do
t he maxi mum concentration occurs at ((Do--F)/b)l/2, while for F>Do t he
maxi mum concentration shifts to x=O. Thi s means that t he muhiplicative
noise can activate animals from grouping around two centers to grouping
around t he potential barrier.
DYNAMIC ASPECTS OFANIMAL GROUPING 4-3
VI . GR OUP S I Z E DI S T R I B UT I ON
As group-living organisms wander about in a limited space, interaction
bet ween groups can occur so that two of t hem encount er and join or
amalgamate, or a large group splits itself into two smaller groups. The
continuation of this interaction processes of amalgamation and splitting
may eventually lead to an equilibrium distribution of t he group size under
given ecological and behavioral constraints.
The equilibrium may or may not be stable within given intervals of
time. Thus, Clutton-Brock and Harvey (706) observed that groups of
social primates were usually based on stable associations of related females.
Unst abl e groups are also observed with many species of animals (Ki t chen
(707); McHugh (708); Lot t and Mi nt a (709)).
It has long been known that grouping behaviors of animal serve for
cover-seeking in whi ch each individual tries to reduce its chance being
caught by a predat or by positioning itself within a group (Galton (770);
Williams (777)). Thus a sudden stimulus causes schooling fish to cluster
very tightly (Breder (772); Springer (713)). I n this context, Hami l t on
(77d) present ed an interesting model for animal aggregation based on t he
behavior of cover-seeking ("selfish aggregation"), which may eventually
result in a very tight group or groups of animal.
Groupi ng is also considered advantageous for prey to predat or detec-
tion (Brock and Riffenburgh (775)). Pulliam (116) provided a simple
calculation whi ch shows an advantage accrued to flocking birds from a
purel y defensive standpoint of predat or detection. Thus, Pulliam' s model
predicts an inverse relationship between t he time spent in predat or sur-
veillance and t he flock size. Pulliam' s model has been tested by Elgar and
Catterall (777) with house sparrows, Passer domesticus, by Tr eher ne and
Fost er (778, 779) wi t h ocean skater, Halobates robustus, and by Har t and
Lendr em (720) wi t h ostrich, Struthio camelus.
A variety of dynamical model s for t he group size distribution has
been developed. These models range from deterministic to probabilistic
representations and vary in assumptions about t he dependence of rates
of joining and splitting groups on t he sizes and numbers of groups. Yet
an entirely different approach may be made by t he use of t he principle
of maxi mum ent ropy subject to pert i nent constraints, provided t he concept
of t he ent ropy of groupi ng be established.
4 4 A. OKUBO
1. G r o u p S i z e S t a t i s t i c s
A group of animal is characterized by the number of individuals in t he
group. Thus, we define a j - gr oup or group of s i z e j to be a group contain-
ing exactly j individuals ( j = 1, 2, 3, . . . ) . Gr oups of size 1 correspond t o
isolates. A syst em of groups is the totality of animal groups in a particular
situation.
Let g~(t) be t he number of groups of size j at t i me t. The total number
of groups G ( t ) and t he total number of individuals engaging in groupi ng
N ( t ) are given by
G ( t ) = ~ g j ( t ) (VI. 1)
j = l
c o c o
N (t) = i~=1.= j g j ( t ) = j~=l"= n~(t) (VI. 2)
where n~(t) is t he total number of individuals in j - gr oup. In reality t he
upper limits of the summat i on j are finite, but mathematically extendable
to infinity wi t h underst andi ng that g~, nj_~ 0 for j exceeds a certain value.
Inst ead of t he group size distribution g3" we often consider t he group
size frequency by normalizing g~ by t he total number G:
f ~(t) -= g~(t)/G(t) (VI. 3)
Thus,
o~
L ( t ) = 1
] L ( t ) = N ( t ) / C ( t ) =_ q ( t ) ( V I . S)
j =l
where q ( t ) is t he mean number of individuals per group.
Strictly speaking t he grouping processes of amalgamation and split-
ting are stochastic in nature. Thus t he number of groups of size ] at t i me
t is subj ect to stochastic fluctuations, and g i ( t ) should be considered as a
stochastic variable. I n ot her words we cannot predict exactly how many
groups of a particular size will be at any t i me t ~0 , given initial group
size distribution, even under identical macroscopic conditions.
I n t he purel y stochastic formul at i on an appropriate funct i on t o
describe t he instantaneous syst em of groups might be
p ( m , j ; t ) , m= 0 , 1 , 2 ..... j = 1,2,3 . . . . (VI. 6)
i . e . , t he probability that exactly m groups are found in ]-group at t i me t,
and p is subject to
p ( m , j ; t ) > 0
~ . p ( m , j ; t ) = l for a l l j and t
m=O
The kt h mome nt wi t h respect t o m o f p is defi ned by
m k p ( m , j ; t) =-- ~*~(j; t) =_ M ~ ( j ; t)
m=0
45
(VI. 7)
( v i . 8)
(VI. 9)
whi ch is t he probabi l i t y average of (number) ~ of j - gr oup at t. Not e t hat
M0(j; t) = 1 (VL 10)
M I ( j ; t) = M ( j ; t) = R/ t ):
average number of groups of size j at t (VI. 11)
Also
~ . m j p ( m , j ; t) = N ( t ) (VI. 12)
j =l m=0
t he t ot al number of i ndi vi dual s.
We are mai nl y concer ned wi t h a closed, conservat i ve syst em of
groups, wher e no i ndi vi dual s i n t he syst em can leave it, no i ndi vi dual s
out si de t he syst em can ent er it, and ani mal popul at i on i n t he syst em
cannot r epr oduce, nor die. Th e n t he t ot al number of i ndi vi dual s i n t he
syst em N remai ns i nvari ant wi t h t i me.
2 . D e t e r m i n i s t i c M o d e l s
We will pr esent a general gain-loss equat i on for t he group size di st ri bu-
t i on g ~ ( t ) . Le t a ( k , l ; t ) be t he rat e of amal gamat i on for groups of sizes k
and l i n ( t , t + O t ) , and b ( k , l ; t ) be t he rat e of spl i t t i ng of k-group i nt o
groups of sizes l and k - - I i n ( t , t + g t ) . Clearly,
a ( k , l ; t) = a ( l , k ; t) for all k and l (VI. 13)
b ( k , l ; t) = b ( k , k - l ; t ) , k > 2 , k > l > l (VI. 14)
We assume t hat for gt suffi ci ent l y small, t he chances for t hree or mor e
pairs of groups t o amal gamat e i nt o one gr oup and for a gr oup t o split i nt o
mor e t han t wo groups can be i gnored. Also assume t hat for t he same smal l
i nt erval of t i me t he chance t hat a gr oup is i nvol ved bot h i n amal gamat i on
and in spl i t t i ng can be i gnored.
I t is t hus obt ai ned
46
and
j - 1
~j = 1/2 2~ a(k, j - k ; t)g~g~_~-- ~. a(k, j ; t)g~gj
k=O k=l
j - I
+ ~. b ( l , j ; t ) g, - - X b ( j , k ; t)g~, j > 2
I =j +l k=l
A. OKUBO
(VI. 15)
c o o
~,~ = - - gt ~ a(k, 1; t)g~ + ~ b(k, 1 ; t)ge (VI. 16)
k=l k=2
whi ch are subject to the initial conditions
g/0) = v~, j _ l ( v I . !7)
Eqs. (VI.15)-(VI.17) are a set of deterministic phenomenologieal
equations, and wi t h specifications of the rate parameters describe t he dy-
namics of t he (mean) number of groups of various sizes. A syst em of
equations similar to (VI.15)-(VI.17) has been widely used in modeling
clustering-splitting processes. For example, in t he context of pol ymer
chemistry, Whi t t l e ( 727) , Van Dongen and Ernst ( 722) , DeLi si and Perel-
son ( 723) , and Hendri ks e t al. ( 724) have analyzed such a quadratic model
for t he kinetics of polymerization and fragmentation, and provi ded an
equilibrium solution for the model.
In general little is known about t he solution of t he syst em of quad-
ratic equations. Oft en an assumption is made for t he principle of detailed
balance, i.e. t he forward rate of amalgamation of i- and j - gr oups is equal
to t he backward rate of splitting of ( i +j ) - gr oup;
a(i, j)g~gj = bq + j , i)g~+j
The detailed balance condition guarantees t he existence of a stationary
(equilibrium) group size distribution g~*.
I n t he context of grouping, James (725), Coleman and James ( 726) ,
Whi t e ( 727) , Goodman ( 128) , and Cohen ( 729) devel oped model s for a
syst em of "casual groups" or "freely-forming groups" in whi ch indi-
viduals are relatively free to maintain or break off contact wi t h one an-
other, that is, informal controls on behavior are at work and spontaneity
is at a maximum. I n these model s t he relevant processes are t he depart ure
of individuals from groups and arrival of individuals to groups. Thus,
t he possibility of a group of size j_>2 joining or leaving a group size of
i >2 is ignored.
Thi s model specifies t he rates of individuals joining and leaving
groups in the syst em as follows:
DYNAMI C ASPECTS OF ANI MAL GROUPI NG 47
a( k, l ; t ) = a for k = 1, l >2 or for k >l , l = 1 (VI. 18)
0 otherwise
b( k, l ; t) = bk for l = 1 or for k - l = 1 (VI. 19)
0 otherwise
Eqs. (VI.15) and (VI.16) are t hen wr i t t en as
9g = agl gy _l - agl g~+b( j +l ) gj 1- bj gj , j>_2 (VI. 20)
: b o
gl --agl ~ gk-k ~, kg~: (VI. 21)
k=l k=2
Summi ng up for all j = 1, 2, . . . , we obt ai n
, 4
c o
-~- Y~ gy(t) = 0 for all t (VI. 22)
tit j=l
i f g l = b ~, kgk/ a ~, gk, so t hat t he t ot al number of groups is conserved;
k=3 k=2
gj = G = constant (VI. 23)
j =l
For an equi l i br i um di st r i but i on of groups, gs*, t he l ef t - hand sides of
(VI.20) and (VI.21) are set zero and solve for gj t o fi nd
KJ
g j* = G (e/~_ 1)j! ' j = 1, 2, 3 . . . . (VI. 24)
wher e
K= In (-q~-+ 1) (VI. 25)
(VI.24) is t he t r uncat ed Poi sson di st ri but i on. The equi l i br i um relative
f r equency of groups g~*=f ~*/ G is gi ven by
gJ
f~* - ( e K, 1) j ! , j = 1, 2, 3 . . . . (VI. 26)
whi ch is charact eri zed by a single combi ned par amet er K.
Gi ven t ot al numbe r of groups, K increases as t he t endency for i ndi -
vi dual s t o j oi n a group and decreases as t he t endency for i ndi vi dual s t o
leave a group, For a part i cul ar val ue of K t he f r equency di st r i but i on of
groups may exhi bi t a single mode at j = [ K] : t he great est i nt eger not
great er t han K. I n part i cul ar, when K< 2 or a G/ b < e ~ - - l , isolates f or m
t he most numer ous gr oup and t he number of groups decreases mono-
48 A. OKUBO
t onousl y wi t h t he size. Since t he rat e for i ndi vi dual s t o leave a group in-
creases l i nearl y wi t h t he gr oup size whereas t he rat e t o j oi n a group is
i ndependent of t he group size, larger sized groups t end t o be unf avor abl y
"sel ect ed" against smal l er sized groups, t hus l eadi ng to an equi l i br i um
size di st r i but i on t hat st rongl y decays t owar d groups of large size.
Cohen' s model for casual groups ( Cohen ( 729) ) assumes sl i ght l y dif-
ferent rat e paramet ers. Bot h t he rat e of j oi ni ng and t hat of leaving a gr oup
consist of t he s um of t wo rates, one i ndependent of t he group size and
ot her proport i onal t o t he size;
a(1, l; t) = a o +a l l (VI. 27)
b(k, k- - I ; t) = bo + b~k (VI. 28)
wher e a 0, a~, b0, bl are all non-negat i ve const ant s. Hence
~j = { a o +a l ( j - - 1 ) } g j _ l - - ( a o + a j j ) g j + {(bo+ b l ( j + l )} g~+l
- - ( bo+bl j ) g~, j > 2 (VI. 29)
oo oo ~o b
gl = --ao 52 g ~- - a o g l - - a l ~ i g~- - al gl +bo ~.] g i + og2
1 1 2
c o
+b l 2~ ig~ +2btg2 (VI. 30)
2
Thi s model conserves t he t ot al number of i ndi vi dual s such t hat
or
d
c,o .
i~__13g~ = 0
~7
N = constant
(VI. 31)
but does not conserve t he t ot al number of groups G unl ess
d
c o
- ~- ~ g~ = - a o G - a l N + b o ( G - n l ) + b l ( N - n l ) = 0
a t . / ' = 1
(VI. 32)
I t is i nt er pr et ed t hat a 0 specifies t he at t ract i veness of groups t hem-
selves, a 1 t he at t ract i veness of j oi ni ng i ndi vi dual s i n groups, b 0 a group
depar t ur e rate, and b I an i ndi vi dual depar t ur e rate. Not e t hat i n t hi s
model t he arrival rates for groups j > 2 are i ndependent of t he number of
isolates i n t he syst em.
Cohen ( 7 2 9 ) f ound t hat his model generat es ei t her t r uncat ed Poi sson
or t r uncat ed negat i ve bi nomi al di st ri but i ons. Specifically t he case ( a l =
D Y N A M I C ASPECTS O F A N I M A L G R O U P I N G 4 9
bo=O ) leads to t runcat ed Poisson, and t he case (bo=O, a l <bl ) leads to
t runcat ed negative binomial, i.e.
g~ = G F ( j + p - - 1 ) ( 1 _~)o~-1 (j = 1, 2, ...) (VI. 33)
r ( p ) r ( j )
where p =a o / a l, f l = - a l / b l < l , and F is a gamma function. Shorrock (130)
and Bart hol omew (131) deal wi t h this particular case.
Especially for p = l ( a o = a l ) , t he negative binomial distribution is re-
duced to t he geometric distribution
g , = G( ~ ) y ( j = l , 2 ) .... (VI. 34)
wi t h f l <l . The geomet ri c distribution is also obtained when a l = b l = 0
as a special case of t he Yul e-Si mon model (Bartholomew (131)).
The present ed model s for group size di st ri but i on are classified as
deterministic simple because neither the rate parameters are t reat ed as
random nor t he model pertains to systems of groups. I n ot her words,
gj ( t ) ( j = 1, 2, . . . ) represent a state of groups of various sizes rather t han
an ensemble of groups of various sizes at t i me t.
Goodman (128) first clarified this issue of a syst em vs. systems of
groups, and also brought out certain probl ems inherent to mathematical
procedures involving averages over quadratic terms. Suppose Eqs. (VI.20)
and (VI.21) are valid for an instantaneous, particular state of groups, and
g(t) may be decomposed into an ensemble average g~(t) and fluctuations
g/ ( t ) ;
gj(t) = o~(t) +g/ ( t ) (VI. 35)
Averaging (VI.20) and (VI.21) over an ensemble after making use of
(VI.35), we obt ai n
d _
~-g~ = @lgj-1 +agl'g~_l--aglg~--agl'g~_l
+( j +l ) g j +~ - b j ~ j , j>_2 (VI. 36)
d - co oo
d{g~ = --ag~k~---lg~--ak~==igt'gzt+bk=2 ~' k~ (VI. 37)
Since gl and gj are random variables, g~g~ cannot be replaced by gl gj unless
t he fluctuations in gl and g3" are mut ual l y uncorrelated. The covariance
funct i ons appeared in (VI.36) and (VI.37) are unknown, and t hus t he set
of equations is not closed. In ot her words, if g~(t), j = l , 2, . . . , represent
50 A. OKUBO
st ochast i c variables, t he ensembl e averages g~(t), j = l , 2, . . . , cannot be
satisfied by a set of equat i ons (VI.20) and (VI.21) si mpl y by repl aci ng g~
by g~ and wi t hout knowl edge of t he covariance of t he fl uct uat i ons we can-
not obt ai n t he sol ut i on for t he ensembl e averages.
3 . C o n t i n u o u s M o d e l s : Di f f u s i o n A p p r o x i m a t i o n
Whe n t he number of ani mal s f or mi ng groups is sufficiently large, we
may develop a si mpl e cont i nuous st ochast i c model , whi ch leads t o a di f-
f usi on equat i on known as a Fokker - Pl anck equat i on.
Le t n ( t ) be t he number of i ndi vi dual s i n a group of size n at t i me t.
The rat e of change of n wi t h t i me is det er mi ned as t he di fference bet ween
t he rat e ani mal s j oi n t he group and t he rat e ani mal s leave t he group. We
assume t hat t he bot h rates of j oi ni ng and leaving consist of a component
whi ch is i ndependent of n and a component whi ch is pr opor t i onal t o n
( Ander son ( 88) ) .
Th e n t he basic dynami c equat i on for t he number of i ndi vi dual s reads
or
where
dn
d-T = ( A o + A ~ n ) - ( Bo + B t n ) (VI. 38)
d n
d t = c~-- fin (VI. 39)
a ~ A0--B0, f = B 1 - A 1 (VI. 40)
We also consi der t hese net rates a and/ ~ as bot h det er mi ni st i c and sto-
chastic, and t he st ochast i c part s represent whi t e noi se fl uct uat i ons of
zero mean whi ch var y rapi dl y compar ed wi t h vari at i ons i n n ( Ander son
( 88) ) .
He nc e
= ~ +~1(t) (vI. 41)
f = ~ +rCKt) (vI. 42)
where ~,, ~, 2, and y are const ant s i n t i me, and
~ = 0 (i = 1, 2)
O~(t),(t+v) = a(v) ( i = l , 2) (vI. 43)
~(t ' )~)/ t ") = 0 for i : / : j
Usi ng (VIA1) and (VI.42) we rewri t e (VI.39) as
dn
- ~ - = 5 - - f i n + 2l(t)--Tn2(t) (VI. 44-)
L e t p ( n , t ) b e t h e p r o b a b i l i t y d e n s i t y f u n c t i o n t h a t t h e n u mb e r of i n-
di vi dua l s i n a g r o u p l i es b e t we e n n a nd n + d n at t i me t. F r o m ( VI . 43)
a nd ( VI . 44) a F o k k e r - P l a n c k e q u a t i o n f or p ( n , t ) is gi ve n b y
31) 3 1 3 2
3t - - 3n {(cT--/3n)p} + ~ - ~ffn2 {(22+TZnS)p } (VI. 45)
wh e r e I t o - c a l c u l u s is a s s u me d ( Ar n o l d ( 79) ; I t o ( 7 5 2 - 7 3 4 ) ) .
We wi l l d i s c u s s a s t e a d y - s t a t e d i s t r i b u t i o n b y s e t t i ng 3p/ Ot =O. I n -
t e g r a t i o n o f t h e r i g h t - h a n d s i de o f ( VI . 45) yi e l ds
- l d 2 ~
(ff--fln)p*(n) + 2 dffn {(2 + r n )p*(n)} = 0 ( v I . 46)
wh e r e p * ( n ) is t h e s t e a d y - s t a t e ( e q u i l i b r i u m) p r o b a b i l i t y d e n s i t y f u n c -
t i on, a nd t h e c o n s t a n t o f i n t e g r a t i o n i s z e r o s i nc e b o t h p * ( n ) a nd d p * / d n
mu s t a p p r o a c h z e r o f a s t at v e r y l ar ge v a l u e s o f n. ( VI . 46) c a n b e i nt e -
g r a t e d t o r e s u l t
p*(n) = C(2~+rZn2)~/r 2-1 exp ~2 ~ - t an -1 (-Lr n~l (VI. 47)
wh e r e C is t h e n o r ma l i z a t i o n c o n s t a n t t o b e d e t e r mi n e d f r o m t h e c o n d i -
t i o n
f : p * ( n ) d n : 1 (VI. 48)
A f e w s peci al cas es o f i n t e r e s t a r e c o n s i d e r e d .
1) ~: <0, / ~ = 7 = 0 : a ni ma l l e a ve s a g r o u p wi t h a c o n s t a n t r a t e a nd
t h e j o i n i n g r a t e i s s t oc ha s t i c .
p*(n) = 22/25 exp {-- (2c7/2~)n} (VI. 49)
He n c e t h e si ze d i s t r i b u t i o n is e xpone nt i a l or g e o me t r i c .
2) ~ > 0 , ~ > 0 , 7 = 0 : a ni ma l j o i n s a g r o u p wi t h a c o n s t a n t r at e,
l e a ve s wi t h a r a t e p r o p o r t i o n a l t o t h e n u mb e r , a n d t h e j o i n i n g r a t e is al so
pa r t i a l l y s t o c h a s t i c
p* (n) = 2(/~/:r2~)e -Z~/~ ~' exp (25/22n -- fi/22n 2) (VI. 50)
He n c e t h e si ze d i s t r i b u t i o n i s t r u n c a t e d Ga u s s i a n wi t h t h e mo d e at n =
2~//~ a nd t h e va r i a nc e of 22/4/3.
52 A. OKUBO
Ander son and Okubo (/35) proposed a st ochast i c model for zoo-
pl ankt on swar m number di st ri but i on. Thei r approach is based on a si mpl e
concept of fission and encount er processes i nvol ved i n swarmi ng. Le t
g ( t ) be t he number of swarms in an area. The rat e of change of g is det er-
mi ned by t he rat e of fission whi ch is pr opor t i onal t o g and by t he rat e of
encount er whi ch is pr opor t i onal t o g2 as a quadr at i c encount er process
suggests. Th e fission rat e also cont ai ns a r andom factor.
Thei r basic equat i on for t he dynami cs of swar m number is given by
d g
- d t - = a g - - b g ~ + c g f ( t ) (VI. 51)
where a and b are spl i t t i ng rat e and encount er rate, respectively, c is t he
ampl i t ude of t he r andom spl i t t i ng rate, and f ( t ) is a whi t e noi se f unct i on
of zero mean and uni t vari ance;
f ( t ) = 0 I
f ( t ) f ( t +r ) = ~(r) / (VI. 52)
The probabi l i t y densi t y f unct i on of fi ndi ng t he number of swarms
bet ween g and g + d g at t i me t , p ( g , t ) is obeyed by a Fokker - Pl anck equa-
t i on
~ p _ ~ c 2 ~ 2
a t a g { ( a g - - b g ~ ) P } + ~ - T ~ ( g 2 P) (VI. 53)
o g
A st eady-st at e di st r i but i on p * ( g ) is obt ai ned f r om (VI.53) i n a way similar
t o t he previ ous case. I t resul t s
p * ( g ) ( g / m) ~ - l e - g / ~ (VI. 54)
mF(v)
where F(u) is a gamma funct i on, and
m = c~/2b )
v = ( 2a/ c 2) --1 >0 / (VI. 55)
The probabi l i t y densi t y f unct i on for p * ( g ) is t he gamma probabi l i t y f unc-
t i on. The charact eri st i c propert i es of t he swar m number di st r i but i on are
mean ~ ~ = ( a / b ) ( 1 - c 2 / 2 a ) (VI. 56)
mode _= gmoa = ( a / b ) ( 1 - c Z / a ) (VI. 57)
variance _-__ a ~ = ( c Z / 2 b ) 2 ( 2 a / c ~ - l ) (VI. 58)
A part i cul ar case wher e a = c 2 or u = l reduces (VI.54) t o an exponent i al
DYNAMI C ASPECTS OF ANI MAL GROUPI NG 53
di st ri but i on
p*(g) = m-ie-g/'~ (VI. 59)
4. Ent r opy o f Grouping
Gi ven a total number of groups G the total number of ways in which G
may be formed into groups of different sizes, gl , g2, . . . , is given by
W( g) = G!/ ~I g; ! (VI. 60)
j =l
When gj is large, we may use t he Stirling approximation for t he factorials,
and (VI.60) can be wri t t en
co
In W = - G )Z. (g,/G) In (&/G) (VI. 61)
i =1
Since g i / G = f i , frequency of / - gr oup, (VI.61) is expressed as
We define
ln W
e o
- - Y2~ f , I n f ~ ( V I . 6 2 )
e i =1
H --- -- ~. f , in fi (VI. 63)
f =l
as t he ent ropy of t he frequency di st ri but i on of group size or simply t he
"ent r opy of groupi ng. " The form of H is in fact analogous to t hat of t he
ent ropy of t he set of probabilities p, as defined in statistical mechanics.
It is seen that H has the following properties that substantiate it as
reasonable measure of disorder in grouping (Shannon and Weaver (136));
i) I f all t he animals gather in one group (i----3, say): corresponding to a
state of maxi mum order, fa----1, ~----0 ( i ~3) . Hence, H= 0 , that is, t he
disorder is zero. ii) I f each group size contains exactly one group, cor-
responding to a state of maxi mum disorder wi t hout any constraints ot her
than Z f l = l , f l = l / G . Thus, for given G, H has a maxi mum value; H=
- - 1/ G( l n 1 / G) G= l n G. As G--~ co, H- - * co. iii) Any change t oward equal-
ization of t he frequencies j~ increases H. I n view of these properties we
consider (VI.63) as a proper definitian of t he ent ropy of grouping wi t hout
requiring g;- to be large.
For our ent ropy of grouping t here are t wo "nat ural " constraints to
t he frequency distribution:
f~ = 1 (VI. 64)
54 A. OKUSO
i f , = N / G = q (VI. 65)
where N is t he total number of animals in grouping, and N / G is t he mean
number of animals per group, whi ch is fixed constant.
The principle of maxi mum ent ropy states t hat t he most probable
distribution f i corresponds to t he maxi mum of (VI.63) subject to given
information or constraints, (VI.64) and (VI.65) (Jaynes (737); Dias and
Shimony (738); Tikochinsky e t al . ( 1 3 9 ) ) . To maximize t he ent ropy under
t he constraints, t he met hod of Lagrange multipliers can be used. The
distribution ~ of maxi mum ent ropy is t hen given by
]~ = exp {--(1+20+21i)} (VI. 66)
where 2 o and 21 are Lagrangian multipliers selected to fit t he given data
(VI.64) and (VI.65). Substituting (VI.66) into (VI.64) and (VI.65) we find
e- I1*~0 = e c - - 1 ( V I . 67)
e - ~ , = e - ( v I . 68)
where
c = In (q/ (q--1)) (VI. 69)
Fr om (VI.66), (VI.67), and (VI.68) we obtain
A
f , = (e c - 1)e -c* (i = 1, 2,...) (VI. 70)
OI
f ~= 1 [_q_q ~-~ ( i =1, 2, . . . ) (VI.71)
q - - l \ q - - l J
Hence, t he frequency distribution is exponential or geometric.
VII. EXAMPLES OF ANI MAL GROUPI NG AND COMPARI SONS WI TH
MATHEMATI CAL MODELS
1. I n s e c t S w a r m s
Al t hough insect swarming is one of t he most extensively studied areas of
animal grouping, only a few works have focused on t he mot i on of individ-
uals in a swarm. Among these few, studies of t he migrating swarm of t he
Desert Locust ( S c h i s t o c e r c a g r e g a r i a , Forsk) using consecutive photo-
graphy have done much to reveal t he orientation and flight behavior of
swarming individuals (Betts, (TdO); Rainey ( l d 7 - 7 4 3 ) ; Sayer ( 7 4 4 ) ; Waloff
( 7 4 5 , 7d6)). These investigators demonstrate that t he orientation of in-
dividual insect is somewhat irregular despite the fact that t he swarm as a
whol e drifts downwind. The interior of t he swarm consists of a number
of subswarms each moving in a more or less random direction. On the
ot her hand, insects at the leading, peripheral, and rear edges show a con-
sistent orientation t oward t he swarm interior. Thi s behavior at t he edges
apparently provides a mechanism by which t he swarm is kept cohesive
as it is travelling long distance for many hours. Were t he swarming in-
dividuals carried passively by winds, t hey woul d spread owing to atmos-
pheric t urbul ence (Rainey ( 7 5 ) ) .
Recently Baker e t a l . ( 1 4 7 ) have analyzed high-speed films of swarms
of L o c u s t a m i g r a t o r i a to measure locust' s body orientation, flight track,
height and speed of flight. I n all swarms st udi ed mean course angle and
mean track angle relative to wi nd direction were significantly different
from zero as a whol e swarm drifted predomi nant l y downwind.
Radar has proved to be a useful t echni que for the st udy of insect
flight ( Har dy e t a l . ( 1 4 8 ) ; Atlas e t a l . ( 1 4 9 ) ; Schaefer ( 1 5 0 ) ; Greenbark
e t a l . ( 1 5 7 ) ) . Radar observat i on enables us to st udy such factors as day
and night airborne insect populations, species and sex identification, air
speed and orientation, and wing-beat rates. Even a single insect echo
is able to observe, and t hen we can track the individual and st udy its move-
ment pattern. The potential of radar in t he st udy of insect swarms is
clearly demonst rat ed by Schaefer' s work (Schaefer ( 7 5 0 ) ) .
Many species of di pt era are know to form swarms (Downes (4);
Sullivan (5); Syrfiimiiki ( 7 5 2 ) ) . Characteristically swarming is performed
by a single sex, usually male; females are attracted t o a swarm and mating
may result (Chiang e t a l . ( 7 5 3 , 7 5 4 ) ) . The swarming activity is diurnal
wi t h peaks at dusk and dawn, although there are some exceptions. Con-
spi cuous objects, animals, and special environmental conditions serve as
swarm markers and i nduce individual insects perform t he so-called
"swarmi ng dance" in grouping. The pat t ern of ttight of swarming indi-
viduals varies wi del y from very regular movement , hovering and circling
around a definite position in t he air, to very irregular, almost random
movement (Dowries (d); Koyama (5, 1 5 5 ) ) .
Ther e is some cont roversy regarding t he interaction of individuals
wi t hi n a swarm. Downes (d) claims that the swarm is essentially a group
of individuals responding i ndependent l y to a swarm marker; it does not
owe its existence to responses among t he insects themselves. On t he ot her
hand, Koyama ( 6 , 1 5 5 ) suggests t hat t he number of individuals of Y a n n i a
s c a l a r i s in a swarm is limited by interaction among swarming flies. He
56 A. OKUBO
observed that swarming males frequent l y chased and became entangled
wi t h each other. Details of chasing behavior of houseflies ( Fanni a cani-
cularis) have been studied by Land and Collett (77) by means of high-
speed filming.
Int erference bet ween individuals is also det ect ed in midge swarming
(Okubo and Chiang (72); Okubo et al. (73); Chiang et al. (753, 754));
these investigators analyzed high-speed movies of Anar e t e pr i t char di
swarming and showed t he presence of interactions and an inward force
exerted on individuals at t he peri phery of a swarm. Gol dsmi t h et al.
(57) analyzed t he same movies to show that t he speed of female midges
was reduced in t he presence of more males in a swarm, while t he speed
of males was not affected by a large number of their own kind. Shi nn and
Long (756) claim that insect orientation to a swarm marker probabl y acts
to keep t he swarm in t he maker area, while orientation t o ot her individuals
is important to cohesion of the overall swarm body.
Verification of t he mathematical models for grouping t hus far pres-
ented requires comparisons wi t h appropriate data. Fort unat el y such data
exist in insect swarming../1, pr i t char di Ki m is a midge wi t h a length of
2 mm and a wingspan of 3 ram. The peak of swarming activity occurs
mi dday and t he midges are most active in bright, hot, calm environ-
ments. The animals can be i nduced to swarm by placing a whi t e marker
on t he ground. These characteristics facilitate photographic studies of
swarming behavior (Chiang (7, 3, 757, 158)). The insects swarm about
5 cm above t he ground and cast their shadows on t he marker. The shadow
is used to det ermi ne t he three-dimensional position of swarming midges
wi t hout stereoscopic phot ography (Okubo et al. (759)). A more general
t echni que for three-dimensional phot ography has been developed by
Shinn and Long (754) by t he use of t wo cameras.
The film analysis wi t h Anar et e midges shows t he following details.
1) The swarm as a group moves to and fro above t he swarm marker wi t h
t he swarm dimension pulsating around a mean size. 2) The trajectories
of individuals may be classified into two disinct patterns, one being a
"l oose" pat t ern and t he other a "t i ght " pattern. I n the former pattern,
t he insect exhibits a relatively straight to- and fro-movement that might
resemble a pendul um motion. I n t he latter, on t he other hand, t he insect
exhibits a relatively short, zigzag mot i on that might resemble a random
flight. I n practice most individuals display a pat t ern which combines t hese
t wo extremes. 3) The swarming midges move wi t h a mean speed of 30
cm/sec and maxi mum speed of 90 cm/sec. They also achieve very high
(Veloclty ~ ) (Acceleration ~ )
Fig. 6. Velocity and acceleration field of individual mi dges in an instance of
swarming.
accelerations, as high as more t han t en times t he acceleration of gravity,
G=980 cm/sec 2, and 4) t he kinetic energy expenditure of swarming
midges is esitmated to be 3.5 kcal per kg of body weight per hour (Okubo
et al . (73)).
The high-speed film of An a r e t e mi dge swarming provides us with
basic kinematic data for evaluating swarm maintenance characteristics
such as velocity autocorrelation coefficient, velocity frequency distribu-
tion, acceleration filed, swarm dimensions, and insect number density
distribution (Okubo and Chiang (12); Okubo et al . ( 73) ) .
Figure 6 shows t he velocity- and acceleration-vector field of indi-
vidual midges in an instance of swarming. The mi dge moves with high
accelerations, occasionally twice or more t he acceleration of gravity. The
mot i on inside t he swarm looks more or less random in both velocity and
acceleration, but t he mi dge is subject to an inward acceleration t he magni-
t ude of whi ch increases with distance from t he swarm center and is partic-
ularly high at t he swarm edge. The frictional force on t he flying mi dge
is "essentially" negligible near t he edge, where midge' s speed diminishes,
and t he acceleration may be i nt erpret ed as t he "dri vi ng force" per unit
mass of t he midge. As we present ed modelling for grouping in Chapter
58 A. OKUBO
2,000!
1, 000
m
E
o
l I I I
(No. of individuals)
, )
I I I l n ~
- - , , wv - - 3 - - 2 - - I
I I i
0 ~1 2 3
X rr
Fig. 7. Mean acceleration of swarmi ng mi dges v s . normalized distance from
t he swarm center.
V, t he presence of this inward-oriented force plays a crucial role in main-
taining t he swarm against a general t endency of spread by t he randomness
in t he motion.
The acceleration field in swarming can be seen more clearly in Fig.
7, where we plot t he mean acceleration of swarming insects against nor-
malized distance from t he swarm center. Also shown are the 90% con-
fidence intervals of t he mean values. The normalized coordinate x" is
calculated as ( x - ~ ) / a x for each frame of film, where x, ~, ax are t he posi-
tion, mean position, and standard deviation of x respectively. The effects
of fluctuations in t he swarm center as well as in t he swarm dimension
are most l y eliminated by using the normalized coordinate system (Okubo
et al. (13)).
Fi gure 8 shows t he velocity autocorrelation coefficient against t i me
lag. The autocorrelogram oscillates up and down about t he zero value
as is expected for swarming (consult Chapt er I I I ) . To see it more pre-
cisely we comput e the t wo integrals in Eq. (III.10). Let
Y
I~(t) = t R(r)dr (VII. 1)
0
+,ol
o
- 1 . o 0
Fi g. 8.
- - @- - , dat a; - -
z f f ' x
/ ' ; . o :
I I I |
0.2 0.4 0.6 0. 8
Z" ( s e c }
Ve l oc i t y a ut oc or r e l a t i on coef f i ci ent of s wa r mi n g mi d g e s v s . t i me l ag.
, t he or y.
I 2 ( t ) - - - f t o V R ( r ) d r (VII. 2)
Fi gure 9 shows t he values of I i and 12 against t with R(r) obtained in t he
same series (see Fig. 8). It is seen that R(r) nearly fulfills t he require-
ment s for swarming t ha t / 1 approaches zero and 12 approaches a positive
t
constant value or . r 2 r R ( r J v approaches a negative constant value. At
about t =0. 3 sec, I s begins to take over 11 as t he latter remains nearly zero.
The convergence of 12 with increasing t does not look evident, however.
The non-stationary nat ure of t he data is partly responsible for masking
t he convergence behavior (Okubo and Chiang (72)). Similar data obtained
by Jansen ( T d ) with C h i r o n o m u s midges show t he behavior of t he velocity
autocorrelation more clearly. Figure 9 suggests that, should t he swarm-
ing insects be squeezed into an infinitesimally small spatial domain and
t hen released, t hey woul d spread out for a while, 0.2 sec say, as if t hey
were processed by diffusion and t hen would settle into swarming. Fi gure
10 shows a simulation of this hypothetical experiment. We comput ed a
60 A. OKUBO
A
e 4
e 4
I
O
x
-16
1 " 1 I I J I
t
1~ =- f t :R~(v)d:
o
/
I
g
1 I I I i I
0.1 0.2 0.3 0.4 0 5 0.6 0.7
Time (sec)
Fig. 9. Time behaviors of two integrals related to the velocity autocorrela-
tion coefficient. C), I1; A, 12.
"rel at i ve" variance ~xl ~, defined by t he variance of insect coordinates wi t h
respect to their initial positions; thus, at t =0, axl e- 0. The relative var-
iance starts to increase monotonically wi t h t i me as a diffusion process
does, and t hen ceases to increase, ending wi t h fluctuations. The theoretical
curve shown in Fig. 8 is const ruct ed on t he basis of Eq. (V.40) wi t h h =
6.8/sec and O~e=31.6/sec. The observed behavior of t he autocorrelation
coefficient is consistent not only wi t h the kinematic interpretation of swarm-
ing but also wi t h t he dynamical model for swarming.
As ment i oned in Chapt er V.1, t he frictional coefficient k may be
i ndependent l y estimated by equating h]~ I to t he quadratic frictional drag,
CD/2 pA ]vl~/m wi t h C~, p, A, ~, and m being respectively t he drag coef-
ficient, density of air, area of projection of t he animal body, mean speed,
and t he insect' s mass. A numerical choice that CD=3 (midge' s Reynol ds
number is of t he order of 10), p =1 0 -a g/ cm 3, A= 2 X1 0 -z em ~, and m=
10- 4g gives k=9. 0/ see for ~=30 cm/sec. As this evaluation should be
considered valid only for an order of magnitude, t he agreement bet ween
t he t wo values of k (6.8/sec vs. 9.0/sec) is satisfactory.
D Y N A M I C ASPECTS OF A N I M A L G R O U P I N G 6 1
I I I I 1 }
15-
g-
E
>
U u . I u . z u . o u . 4 u . o u . o 0 . 8
T i me (sec)
Fig. 10. Ti me behavior of relative variance (simulated case).
o
E
J
Fi gur e 11 depi ct s t he f r equency di s t r i but i on of i nsect speed i n a
swar m, whi c h can be wel l a ppr oxi ma t e d by t he Maxwel l i an s peed di s-
t r i but i on. T h e t heor et i cal cur ve is based on Eq. (V.61) wi t h k = 6 . 8 / s e c
and B = I . 1 X 104 cm2/secS; t he val ue of B is es t i mat ed f r om t he obs er ved
me a n s quar ed vel oci t y of i ndi vi dual i nsect s: Eq. (V.38), ~ = B / k = I . 8 X
l 0 s cm2/sec 2.
T h e as ympt ot i cal val ue of t he s t a nda r d devi at i on of di s pl acement s
is es t i mat ed f r om Eq. (V.33) gi ven t he val ues of t he par amet er s :
(2z) 1/2 = ( B / k c o e z ) l / z - 2 cm (VII. 3)
whi c h r easonabl y agrees t o t he val ues di r ect l y obs er ved wi t h mi dges
( Okubo and Chi ang ( 1 2 ) ) .
T h e n u mb e r dens i t y di s t r i but i on of s war mi ng i ndi vi dual s is i n fact
pl at ykur t i c as our ma t he ma t i c a l model , Eq. (V.68), pr edi ct s. Ta bl e I
pr ovi des t he val ues f or skewness (fl11/~) and kur t osi s (/92) c o mp u t e d f r om
data. Smal l val ues of skewness i ndi cat e t he spat i al di s t r i but i on t o be
near l y s ymme t r i c about t he cent r oi d, and st at i st i cal t est s for kur t osi s
shows si gni fi cant devi at i on f r om nor ma l i t y (fl 2=3) wi t h t he onl y excep-
62 A. OKUBO
150
0
" 0
._~ 100
6
z
0 "
50
I,.I_
I I I I ' I ' ' I I ' ' I I
I
10 20 30 40 50 60 70 80
Speed (crn/sec)
Fig. 11. Speed frequency di st ri but i on of swarmi ng midges.
- - , t heory (2D Maxwell).
I
90 100
F ( ~ , data;
TABLE I
Paramet er of Spatial Densi t y Di st ri but i on of Swar mi ng Mi dges: Skewness and Kurt osi s
Series Skewness Kurt osi s
2/3 ( Not availab!e) 2.38
4 --0. 10 3.10
5 0.18 2.61
100-1 0.13 2.07
100-2 0.11 2.78
100-3 0.13 2.68
tion of series 4. Thi s deviation from normality arises from anharmonie
attractive forces that correspond to densi t y-dependent advective velocity
t oward t he swarm cent er (see Chapter V). To demonst rat e this point we
shall take
b(x) = 2 2 x 8 ( V I I . 4 )
D Y N A M I C ASPECTS OF ANIMAL GROUPING 63
in t he dynamical model Eq. (V.13), and use t he steady-state density
di st ri but i on given by Eq. (V.68), i . e.
p*( x ) = Po exp { -- (oJ2k/2B)x 2 - - (2~k/4B)x 4} (VII. 5)
The kurtosis/~s is calculated from ( v i i . 5 ) :
/gs = 3 (1 -- 2B22/ko) 4) (VII. 6)
whi ch guarantees that fl2<3 and t he di st ri but i on is platykurtic. Usi ng
t he observed values of/5~ we can estimate 22 given B, k, and o); for example
~s=2. 61 (series 5) gi ves 22=37/ cm ~ sec 2. Fr om Eq. ( v i i . s ) a critical dis-
tance from t he center, X c , is defined in such a way that
J2k ~X2 2~k
2B -- -4B -X~4
or
Xc = ~/2 ~/2 (vii. 7)
For distances less t han Xc t he harmonic force dominates t he anharmonic
force and vi ce ver s a for the distance larger than X c . For t he above series
X c is estimated as
Xc = 1.2 cm (VII. 8)
Since (X2)1/2=2 cm from (VII.3), t he anharmonic force dominates at
distances of the order of one standard deviation of mi dge positions.
The intensity of acceleration of swarming insects may be estimated
from Eq. (V.48). Thus for A n a r e t e midges, B- - - 1. 110 ~ cm2/sec 8, k =
6.8/see, and we=31. 6/sec, whi ch give
(52) 1/~ = 1,340 cm/sec 2
The mean intensity of mi dge acceleration exceeds the value of t he ac-
celeration of gravity ( G=980 cm/sec2). The result of this calculation is
consistent wi t h t he observed accelerations of midges as shown in Fig. 6.
The maxi mum acceleration observed with A . p r i t c h a r d i is more t han
10 G. According to Alexander ( 760) t he maxi mum forces exerted by
animals in a wi de range of activities is proportional to t he 2/3 power of
body weight; t hus the maxi mum acceleration due to the maxi mum force
is proportional to (body weight)-l/~G. For a body mass of 0.1 mg for the
A n a r e t e midge t he maxi mum acceleration should range from 100 to 1,000 G
according to Alexander' s figure.
64 A. OKUBO
A massive aggregation of pine beetles to host trees has been paid
much attention as t he beetle causes a severe damage to t he forests (Gara
(161); Cost er and Gara (162); Gara and Cost er (163); Pi t man and Vit~
(164); Renwick and Vitfi (165); Geiszler et al. (166, 167)). The attack
behavior of the sout hern pine beet l e (Dendroctonus f r ont aI i s Zi mm) can
be represent ed as a series of events such as directed flight, landing on a
pine t ree and entering t he bark, and it is conditioned primarily by ol-
factory stimuli and communication.
The attack process may be considered in t hree phases. The initial
phase is due to a few "pi oneer " females who select their host in response
to unknown host volatiles in addition to visual and ot her cues. Upon
landing on t he host tree, these females release their pheromones, which
induce aggregation of large number of beetles; at first predominantly
males are at t ract ed and t he responding males t hen release their phero-
mones whi ch reduce t he response of other males, while are attracting
ot her females, and t he process repeats itself. Thus an accelerating mass
attack is a feature of t he second phase. Eventually t he concentration of
animals reaches a carrying capacity of t he host, t he attack is complete,
and t he popul at i on shifts t he focus to another host trees.
Let us consider a simple model for this entire process. N ( t ) is the
number of beetles who have aggregated to a host tree at t i me t. We assume
t he sex ratio to be equal. I n any t i me interval At , t he number of aggregat-
ing individuals increases by A N , which is assumed to be proportional
to t he product of t he number of beetles available for attack:
A N = A ( M - - N ) A t (VII. 9)
where A represents t he rate coefficient and M is t he total number of bee-
tles engaged in attack.
The rate coefficient consists of t wo parts; a part due to the percep-
t i on of a susceptible host t ree by pioneering females who respond to host
volatiles and a part due to t he attraction of other individuals by means
of pheromones released by t he aggregated beetles. The former process is
considered to be density i ndependent and the latter to be density depen-
dent. Hence we have
A = a + b N (VII. 10)
where a and b represent, respectively, the rate constant of a susceptible
t ree and that of a uni t number of insect. The density dependent attrac-
t i on is assumed t o be l i near l y pr opor t i onal t o t he numbe r of i nsect ag-
gr egat ed, i . e . , m= l i n Eq. (V.91).
Subs t i t ut i ng ( VI I . 10) i nt o ( VI I . 9) and t aki ng t he l i mi t At - - ~0, we
obt ai n
2V = ( a + b N ) ( M - N ) (VII. 11)
E
Z
180-
140-
100-
60-
20:
0"
180-
140"
100-
60~
40-
0
a
c
S
100
e
- A
0 2 4 6 8
7 b
80
60
40
20-
0
2 4 6 8 10 12
Days
14
Fig. 12. a-e. Comparison of mathematical model for the aggregation of pine
beetles with field data. O, data; - - - , theory.
6 6 A. OKUBO
The initial condition simply states t hat
N- 0 at t = 0 (VII. 12)
Eq. ( I I . 11) subj ect to (VII.12) has t he solution
N(t ) = M[1 - exp {-at (1 +c)} ]/[1 +c exp { - at ( 1 +c)} ]
where for simplicity M is taken constant and
(VII. 13)
c - bM/ a (VII. 14)
The behavior of t he aggregation curve, i.e., N( t ) vs. t may be classified
into t wo categories according as c_<l or c >1. When c is less t han or equal
to 1, t he aggregation curve possesses no inflection point, while when c
is greater t han 1, t he curve has an inflection point, As c increases indef-
initely, t he inflection point approaches t he mi dpoi nt of aggregation, i.e.
N/ M=1 ~2 . For values of c much larger t han 1, t he curve resembles the
logistic equation.
Thi s simple model can be t est ed against field data on t he daily cumu-
lative number of pine beetles attacking t est trees (Gara and Cost er (763);
Gara et al. (768)). Fi gure 12a-e show t he result of comparison. Tabl e I I
summarizes t he result of evaluation of t he relevant parameters in t he
model. Ther e appears to be an inverse relation bet ween t he t wo rate
constants, a and b.
2. Zooplankton Swarms
The pat chy distribution of marine zooplankton over a large range of
scales is wi del y recognized (Dam-Guerrero et al. (769); Fasham et al.
TABLE II
Estimation of Parameters Used in Modelling the Aggregation of Two Southern Bark
Beetle Species
) I a b
Species (number) (day -1) (day_ ~ no._l) c
D. frontalis 146 0.0004 0.0082 3,000
D. frontalis 142 0.44 0.00094 0.3
D. frontalis 170 0.20 0.0014 1.15
D. frontalis 40 0.013 0.028 89
Ips avulsus 80 0.0061 0.010 135
M, total number of beetles engaged in attack; a, rate constant of host tree; b, per capita
rate constant of settled insect; c=--bM/a
(170); Mackas and Boyd (171); Pugh (172); Wiebe (173)). The degree of
zooplankton patchiness is more intense t han t he patchiness of co-occurr-
ing phytoplankton distribution (Dam-Guerrero et al. (769); Mackas and
Boyd (171)) and t he difference becomes mor e noticeable at smaller scales.
These observations suggest t hat zooplankton should have a special be-
havioral mechani sm for formation of small scale aggregates or swarms
(Clutter (10)).
Certain zooplankters t end to aggregate in regions of high concentra-
tions of phytoplankton (Bainbridge (174)) or amino acids (Poullct and
Quellet (175)), but no data are available for tracking t he swi mmi ng pat-
t erns of individuals within patches. It seems yet unknown if t he indi-
viduals can detect concentration gradients and orient t hei r swimming
paths accordingly.
Ther e are a number of observations on swarming of zooplankton:
copepods (Hamner and Carleton (7); Byron et al. (8); Ueda et al. (9);
Kawamura (176); Omori and Hamner (177)), euphausiids (Kawamura
(176); Omori and Hamner (777); Hamner et al. (178); Nemot o (779);
Sameoto (180); Weber et al. (181); Witek et al. (782)), planktonic shrimps
(Hamner and Hamner (183)), Cladoceran (Brandl and Fernando (784)).
These studies all indicate that swarming behavior is part of t he life history
of many zooplankton populations. Field observations by Hamner and
Carleton (7) show t hat copepods' vision contributes to swarming behavior,
although ot her senses may also be involved. The dispersion or loosening
of swarms at night (Hamner and Carleton (7); Cl ut t er (10); Shaw (185))
furt her supports t he role of vision in t he mai nt enance of swarm integrity.
I n comparison with insect swarming, however, no kinematic data
appropriate for verification of our mathematical models really exist in
t he mari ne field. The mai n difficulty in such a quantitative st udy lies in
t he process of obtaining adequate photographic recordings of swarming
individuals in aquatic environments, although recent progress in high-
speed oceanic mi croci nemat ography has proved very encouraging (Price
and Paffenh6fer (786)).
Hebert et al. (187) report ed that adults of Heterocope septentrionalis,
a large calanoid copepod common t hroughout arctic Nort h America,
swarmed over areas of pale substrate in pond and lake habitats. Swarm
formation was i nduced by placing panels of yellow or white plastic and
t he Heteroscope aggregations were most dense when light intensity was
high and wat er was calm. Hebert et al. suggest that t he swarming behavior
may enhance t he frequency of mat i ng encounters. Thi s finding alone
68 A. OKUBO
indicates that copepod swarming in the wat er should be t he count erpart
of insect swarming in t he air. As report ed by Cl ut t er (70), Hamner et al.
(778), Brand and Fernando (784), and Ratzlaff (788), zooplankton swarms
are usually composed of similar sized individuals, t he feature of which are
seen also in insect swarms.
All t hose studies on zooplankton swarms suggest that the kinematical
aspects of zooplankton swarming shoul d be very much similar to t hose
of insect swarming, and hence t he mathematical model s that are favorably
t est ed wi t h available data on insect swarming shoul d also be applicable
to some zooplankton swarms. To proceed into marine grouping, how-
ever, kinematic data on zooplankton swarming are urgent l y needed for
evaluation of t he mathematical models. For example, information on t he
basic swarm propert i es: t he density of swarms in t he sea and t he distribu-
tion of zooplankton in a swarm will enable us t o estimate at least t he ratio
of t he diffusivity and t he attractive forces in t erms of t he number density
of organisms (Okubo and Chiang (72)).
Wi t ek et al. (782) st udi ed t he aggregation of krill (Euphausia superba).
They describe t he distribution and characteristics of krill swarms in
Antarctic Peninsula region. An effort has been made to obtain the fre-
quency distribution of krill swarm dimensions; t hey range from about
6 0
A 4O
E
O"
20
An t a r c t i c k r i l l
Fi g. 13.
a n e x p o n e n t i a l d i s t r i b u t i o n ) . @, d a t a ; - -
Q
, ~ 0 ~ T
2 0 0 4 0 0 6 0 0
Swar m l engt h (m)
L e n g t h f r e q u e n c y d i s t r i b u t i o n o f kr i l l s wa r ms ( Da t a ar e f i t t ed t o
-, t h e o r y .
D Y N A M I C A S P E C T S O F A N I M A L G R O U P I N G 69
10 m (lowest measurable value) to over 500 m (Fig. 13). A semi-log linear
fit suggests t he following relationship
f ( l ) = f o e - ~ (VII. 15)
where f(/ ) is t he frequency of swarm length l , andf o and 2 are constant.
Organisms are usually distributed heterogeneously in both space
and t i me (Taylor e t a l . ( 1 8 9 ) ) . Thi s uneven distribution of organisms,
oft en referred to as patchiness, is a particularly conspicuous feature of
aquatic populations (Cassie ( 7 9 0 ) ; Tonol l i and Tonolli ( 7 9 1 ) ; Steele
( 1 9 2 - 7 9 4 ) ; Haur y e t a l . (795)). The patchiness occurs in scales ranging
from centimeters (Cassie (796)) to met ers (Fasham e t a l . ( 1 7 0 ) ; Wiebe
( 7 7 3 ) ; Smi t h e t a l . ( 7 9 7 ) ; Greenbl at t (198)) to kilometers (Cushing and
Tungat e ( 7 9 9 ) ) . For mari ne bacteria Mitchell e t a l . ( 2 0 0 ) suggest a pat ch
size of less t han 1 mm.
To what extent zooplankton swarming influences t he scale of patchi-
ness is not clear, but Fasham ( 2 0 1 ) stresses t he i mport ance of swarming
in underst andi ng zooplankton spatial spectra and patchiness in com-
parison to phyt opl ankt on spectra and patch size that have been much
more extensively studied,
Dam- Guer r er o e t a l . (769) have investigated zooplankton patchiness
in Long Island Sound and developed a mathematical model for zoo-
plankton patch size distribution. Thei r model depends upon t he follow-
ing assumptions. 1) Ther e exists a statistical equilibrium in t he kinetic
energy spect r um of t he wat er in Long Island Sound. The equilibrium
spect rum is established in t he range of scales less t han t hose of predo-
mi nant tidal components, and t he energy spect rum is characterized by a
definite group of eddies that contain most of t he energy. 2) The energy-
containing eddies possess a field of velocity gradients. The scales of t he
velocity-gradient field is of t he order of those of t he energy-containing
eddies. 3) A patch of zooplankton is embedded in this velocity field of
t he environmental water.
Thei r model equation for t he length of zooplankton aggregation l ( t )
reads
1 d l ~ I I \ ~ , .
l d t = e~ef~x~x~f[~o)q-c~ (t) (VII. 16)
where e ( t ) is t he uni t vector specifying t he direction of t he velocity gradi-
ents at t i me t , a ~ i / ~ x y is t he velocity-gradient t ensor in t he water, and
d ( t ) is a random variable. I f l ( t ) is much smaller t han lo (a characteristic
7 0 A. OKUBO
l e ngt h of t h e ener gy- cont ai ni ng eddi es), t h e n a gr oup of z oopl a nkt on is
de f or me d and s t r e t c he d by t he vel oci t y- gr adi ent field associ at ed wi t h t he
ener gy- cont ai ni ng eddi es and f ( l / l o ) is uni t y ( Bat chel or a nd To wn s e n d
( 2 0 2 ) ; Saf f man ( 2 0 3 ) ) . As t i me pr ogr esses, t he gr oup l e ngt h l t e nds t o
i ncrease, and t he vel oci t y gr adi ent effect i ve t o t he zoopl ankt on pa t c h
t e n d s t o decr ease and ul t i mat el y vani shes as l is equal t o l o. Da m- Gu e r r e r o
e t a l . pr opos e d s i mpl e f or ms
f ( l / l o ) = 1 - l / l o
e~esOffffOx s = ~7 (constant)
~ , ( t ) = 4 2 ~ ( t )
(VII. 17)
(VII. 18)
(VII. 19)
whe r e 2 K is t h e i nt ens i t y of r a ndomne s s i n t h e gr owt h rat e, a t and ~b(t) is
whi t e noi se wi t h zero me a n and u n i t vari ance, i . e . ,
~ ( t ) = o
(t)~b(t + v) = 3(v) J (VIE 2O)
Combi na t i on of ( VI I . 16) wi t h (VII. 17), (VII. 18), and ( VI I . 19) yi el ds t he
dynami cal equat i on for l
d ~ = - - + ~ / 2 K l ~ ( t ) (VII. 21)
whi c h is essent i al l y t h e same as Eq. (VI. 51), and hence t he pr obabi l i t y
dens i t y f unc t i on f or t he pat ch size, p ( l , t ) is obeyed by a Fokker - Pl anck
e qua t i on same as Eq. (VI. 53).
Un d e r t he a s s umpt i on t ha t t he e nvi r onme nt a l eddi es are i n st at i s-
t i cal equi l i br i um wi t hi n t he t i dal per i od, t he pr obabi l i t y dens i t y f unc-
t i on p ma y also achi eve st at i st i cal equi l i br i um. T h e st eady- st at e p. d. f , is
gi ven by (see Eq. VI. 54))
( l / a ) b-1
p , ( l ) - - a F ( b ) exp ( - - 1 / a ) (VII. 22)
whe r e F ( b ) is a g a mma f unc t i on and
a = lo K / ~ ]
( v i i . 23)
/
b = (s/K)- 1
Not e t ha t K / ~ r epr es ent s a coefficient of r a ndomne s s i n t he vel oci t y
gr adi ent of t he e nvi r onme nt a l wat er. I n a speci al case of ~ / K = 1, ( VI I . 22)
is r e duc e d t o an exponent i al di s t r i but i on (VII. 15).
A
H = 3 m
B
4 "
H=3m
31 o o
g
H = 5 m
5-"
4 -
3-
2-
1-
H = 5 m
H = 1 0 m 5 - H = 1 0 m
4 -
2 o o 2 -
1 o 1 -
1 / I r
200 400 6 0 0 800 2 0 0 4 0 0 600 800
Patch l engt h (m)
Fig. 14. Length frequency distribution of zooplankton patches (Data are
fitted to a gamma distribution).
Th e char act er i st i c pr oper t i es of t he size di st r i but i on are eval uat ed
i n t er ms of t he par amet er s :
i (mean) = l o ( 1 - K / ~ ) I
l,~ (mode) = l o ( 1 - 2 K / ~ ) I (VII. 24)
(standard deviation) = l o ( 5 / K - 1 ) l / 2 ( K / ~ )
Fr e que nc y di st r i but i ons of obser ved and model pr edi ct ed pat ch size are
s hown i n Fig. 14. Th e non- par amet r i c Kol mogor ov- Smi r nov t est s f or
goodness of fit reveal t hat i n five out of six cases of size di st r i but i on t her e
is not si gni fi cant devi at i on bet ween t he model pr edi ct i ons and t he ob-
ser vat i ons at t he l evel of 5%. Th e val ues of t he par amet er s lo and K / ~
est i mat ed f r om t he fit are gi ven i n Ta bl e I I I . Al so s hown are i nde pe nde nt
eval uat i ons of t hese par amet er s ; t he char act er i st i c eddy size l engt hs (lo)
wer e est i mat ed f r om t he power s pect r um of t he cur r ent vel oci t y and t he
coeffi ci ent of r andomnes s ( K / ~ ) was est i mat ed f r om t he rat i o of t he mean
72 A. o~:uBo
TABLE III
Comparison of Mathematical Model and Observation for the Values of 1 o and K/~
Dat e De pt h
(m)
Pat ch size (m) l o (m) g / ~
Me a n S, D. A B A B
9/27/83 3 187 139 291 484 0.36
0.36
9/27/83 5 318 204 499 659 0.29
9/27/83 10 483 257 585 705 0.26 0.31
6/11/84 ; 3 289 148 364 668 0.21
0.16
6111184 5 161 71 192 731 0. 16
6/11/84 10 153 97 214 643 0.20 0.17
A, model est i mat es; B, di r ect est i mat es f r om cur r ent met er r ecor ds; lo, char act er i st i c
l engt h of t he ener gy- cont ai ni ng eddi es; K[~, coefficient of r andomnes s i n t he vel oci t y
gr adi ent of t he envi r onment al wat er.
absol ut e r esi dual val ue of t he shear t o t he mean cur r ent shear. Est i mat es
of K / ~ are r oughl y si mi l ar t o t hose of t he model , whi l e est i mat es of lo are
consi st ent l y l arger t han t hos e of t he model .
Th e l at t er di sagr eement may be par t l y at t r i but ed t o behavi or al
f eat ur es o f zoopl ankt on, such as swar mi ng. I t is i nt ui t i vel y cl ear t hat
swar mi ng woul d make t he val ue of lo effect i vel y smal l er t han t he l efi gt h
of t he ener gy- cont ai ni ng eddi es i n t he envi r onment al wat er . Anot her r ol e
of swar mi ng woul d be t he r educt i on of t he shear effect. Thus , i f t he ani-
mal s t e nd t o concent r at e at a cer t ai n dept h, t he hor i zont al di sper si on by
shear effect will be suppr essed ( Okubo (20d, 205)). Th e quant i f i cat i on of
t he effect of swar mi ng is not possi bl e at pr esent , however .
I t has been known f or s omet i me t hat t he Ant ar ct i c krill, E. superba
f or m swar ms ( Ha mne r et al. (178); Ne mot o (179); Wi t ek et al. (182);
Ever s on (205); Par f enovi ch (207)). Recent st udi es by t he use of acoust i c
t echni ques show t hat t he size of krill swar ms can r ange hor i zont al l y f r om
a f ew met er s t o several ki l omet er s and vert i cal l y f r om less t han a me t e r
t o a few hundr e d met er s. Th e l argest krill swar m ( " s uper s war m" ) was
obser ved i n Ma r c h 1981 near El ephant I sl and (61S, 55W), and t he t ot al
bi omass of t hi s swar m was est i mat ed t o be 2.1 mi l l i on met r i c t ons dis-
t r i but ed over 450 km ~ ( Macaul ey et al. (208)).
Th e f or mat i on of such a s uper s war m i nevi t abl y r equi r es a densi t y-
dependent communi cat i on syst em as model ed i n Chapt er V.4. Chemi cal
or acoust i c communi cat i on mi ght be t hought of a possi bl e mechani sm.
Th e kri l l ' s concent r at i on i n l arge swar ms i t sel f pr esent s an i nt er est i ng
fl ui d dynami cal pr obl em. Si nce kri l l ' s densi t y is about 2% heavi er t han
DYNAMI C ASPECTS OF ANI MAL GROUP I NG 7 3
the surrounding sea water, a krill swarm would appear as a volume of high
density. Hofmann e t a l . ( 2 0 9 ) speculate that the density difference be-
tween the krill swarm and the surrounding water should be sufficient to
produce a water circulation within the swarm. The proposed mechanism
of circulation is as follows. The higher density associated with the krill
swarm produces a downward deflection of constant pressure surfaces.
Assuming the pressure surfaces below the swarm are level, such a distor-
tion of the pressure filed by the swarm will produce an inward-directed
horizontal pressure gradient field,
If no other forces are present, the pressure gradient force must pro-
duce a water circulation in such a manner that the centrifugal accelera-
tion balances the pressurc gradient. This force balance, for a radially-
symmetric, steady swarm, is expressed as
v 2 / r = p l - l Op / a r (VII. 25)
where v is the velocity, r the radial distance from the center of the swarm,
p the pressure, and Pl is the density of the volume containing water and
krill.
Using the hydrostatic approximation we can express the radial pres-
sure gradient in terms of the density gradient, and the circular velocity
within the swarm is given by
v 2 = - - g r p 1 - 1 Op l / ~ r d z (VII. 26)
- - H
where g is the acceleration of gravity, z is the vertical coordinate (positive
upwards), and z =- - H is the bottom of the swarm. Thus the knowledge
of the radial distribution of krill concentration within the swarm will pro-
vide us with the evaluation of the velocity. Hofmann e t al . ( 2 0 9 ) estimate
that for a swarm thickness of 50 m and densities of 0.1 kg/m a and greater,
velocities of 3 to 4 cm/sec can be produced. Thicker and higher densities
swarms could produce velocities approaching 30 cm/sec.
The biologically-induced circulation would have some interesting
implications. At swarm densities of 2 kg/m 8, oxygen depletion in the
water would limit the duration of swarming to a few hours. The induced
swirl circulation acting together with a vertical plane circulation could
provide a mechanism to transport oxygenated water from the environment
into the swarm.
74 A. OKtmO
3. Fi s h School s
Schools are groups of individuals engaging in cohesive movement s with
parallel orientation. The presence or absence of parallel orientation dis-
tinguishes schools from swarms. In a fish school neighboring fish orient
themselves similarly such that t he distance bet ween individuals is fairly
uni form and t he mot i on of individuals is synchronized ( Hunt er ( 270) ) .
Van Olst and Hunt er ( 211) st udi ed t he organization of schools of
Pacific mackerel ( Sc ombe r j aponi c us ) , nort hern anchovy ( Engr aul i s mor -
dax ) , and others, with t he aid of photography. Fi l m analysis demonst rat ed
t hat fish were closer to adjacent neighbors in file t han in rank and also
t hat adjacent fish in file were more likely to have similar directions or
headings t han adjacent fish in rank.
Symons ( 272) , t oo used phot ography to st udy spacing and density in
schools of sticklebacks ( Gas t er os t ens acul eat us ) and mummi chog ( Fu n d u l u s
het erocl i t us) . Fi sh were usually spaced regularly when t he average distance
bet ween nearest neighbors was less t han one fish length, When, on t he
ot her hand, t he distance was greater, i . e. , at lower density, spacing be-
t ween fish was found to be in a state of loose aggregations. Thi s finding
suggests that some critical density is necessary for t he occurrence of
schooling.
Parr ( 213) pioneered in modelling fish schools and discussed t he main-
tenance of fish schooling in t erms of t he balance bet ween two counteract-
ing forces, i . e. , attraction and repulsion of individuals. Lat er Breder ( 274) ,
using an analogy to intermolecular forces, formul at ed these attractive
and repulsive forces as
F = - a / r ~ + bi t ~ (VII. 27)
where r is t he distance bet ween individual fish, a, b, m, and n are constants,
and m < n since attraction can be expected to be more effective at long
distances t han repulsion. The equilibrium distance r e is given at whi ch F
vanishes
r , = (b/a) 1/{~-~) (VII. 28)
Breder ( 214) furt her assume that attraction might be nearly i ndependent
of distance ( m=0) as it is accomplished by vision and also that repulsion
varies inversely wi t h t he square of t he distance (n----2) as in Coul omb' s
repulsion law of electromagnetism. Hence (Vl i . 27) simplifies to
F = - a + b/r z ( v i i . 29)
I n this case t he equilibrium distance becomes
by
75
re = (b/a) 1/~ (VII. 30)
Anot her interesting expression for t he schooling force may be given
F = - - a +B e -b~ (VII. 31)
where a, b, and B are positive constants and it requires B>>a. I n this case
t he equilibrium distance becomes
r~ = b -~ in (B/a) (VII. 32)
and t he force is repulsive for rKKre and attractive for r>)re. I n fact t he
force given by (VII.31) is derivable from t he potential of t he Toda lattice
(Toda (275, 276)). A little more detail discussion on this special force
will be present ed at t he end of this section in t he context of t he propaga-
t i on of a wave of agitation in a school.
Observations of various fish schools reveal that re is 16N25% of the
mean body length of fish, and that repulsion nearly vanishes at distances
beyond one body length. Under normal stable conditions of schooling,
individual fish rarely allow themselves to be separated from others by
more t han 40% of their body length.
Interfish distances estimated from two-dimensional measurement s
t end to be underestimated, and t he nearest-neighbor may even be incor-
rectly identified (Symons (272)). Several techniques for t hree-di men-
sional measurement of school st ruct ure have been developed (Cullen et
al. (217); Dill et al. (278); Kol t es (279); Pitcher (220, 227)).
Okubo et al. (222) present ed a dynamical model for fish schooling.
They consider a two-fish school for simplicity. The equations of mot i on
for a pair of fish in t he direction of school movement , x, are given by
21 = - k21- oJ2xl +A1( t ) }
22 = -k22-~o2x2 +A2(t) (VII. 33)
where x l ( t ) and x2(t ) are, respectively, t he deviations of the position of t he
first fish and t he second fish from its equilibrium. The equilibrium posi-
tion is defined to be t he position of each fish at time t as t hey swim to-
gether at which t he attractive and repulsive forces balance each other.
The first t erms on t he right-hand side represent the frictional force per
unit mass, assumed to be proportional to the velocity, t he second t erms
76 A. OKUBO
represent t he harmoni c force t oward t he equilibrium position, and t he
t hi rd t erms represent t he "swi mmi ng force" per uni t mass.
I n this model of schooling, t he xl-fish is assumed to follow t he x 2-
fish; t hus as t he x~-fish accelerates, t he xl-fish responds to t he leader wi t h
a small but finite t i me lag (latency) (Shaw ( 785) ; Hunt er (223)). We t hus
express t he swimming forces for two fish as
A / t) = A(t)
(VII. 34)
f
As(t) = A~(t+~) = A ( t + e ) , s>0
The smallness of e permits us to expand A ( t + s ) in a Tayl or series in e;
A ( t +e) = A( t ) +e A( t ) +e2/2A'(t) + . . .
Fr om (VII.33) we can obtain t he equation of t he cent roi d velocity, V(t)=_
(&1+~2)/2 and t he equation of t he relative separation bet ween two fish,
r ( t ) - x 2 - - x r They are
V + k I T + w 2 V = S( t ) +~/ 2A' ( t ) +2/ 4 Ai t ) }
(i:/~) + k(~/e) +~o~(r/s) = A( t ) +,/2 A(t) +e2/6 A'(t) (VII. 35)
Hence V and r/e obey t he same dynamical equation to first order in ~.
Thi s is consistent with experimental results due to Inagaki e t al. (224).
The frequency spect rum for t he fluctuations of V or r/~ can be cal-
culated from (VII.35) by assuming t hat A(t) has a white noise spect rum,
i.e~ A varies extremely rapidly compared with t he variations of V or r / z .
We t hen obtain t he spectral density funct i on ( f ) of V or r / z as (see Eq.
(v.20))
J(f) = 22{f2-~oZ)U+ k~ f u } - * (VII. 36)
where f is t he frequency and ~ is t he (constant) spectral density of t he
white noise variable A. For small values of f , (VII.36) t ends to
( f ) ~ 2 / w 2 = constant (vi i . 37)
while for large va!ues of f , (VII.36) t ends to
q~(f) ~ 22f-* (VII. 38)
Inagaki et al. ( 224) observed these two limiting relations in experiments
using Gnat hopogon elongatus eIongatus. The theoretical spect rum fits very
well to t he observation by taking o=l . 3/ s e c , k=2. 0/ see, and , =0 . 2 sec
(Okubo (26); Okubo et al. ( 222) ) .
DYNAMIC. ASPECTS OF ANIMAL GROUPING 77
It may be i mport ant to poi nt out that the dynamical model for fish
schooling is essentially t he same as t he linear dynamical model applied
t o insect swarming, Eq. (V.15). The only difference appears in the forcing
funct i on A vs. W; in fish schooling t he force bet ween fish is closely related
wi t h a small t i me lag, while in insect swarming each insect is subject pri-
marily to an i ndependent random force.
An extension of this model for two-fish to many-fish schooling may
require significant modification as Williams (225) and Partridge (226)
observed that internal organization and st ruct ure of two-fish schools were
quantitatively different from those for large schools. Partridge ( 226) made
t i me series analyses wi t h European mi nnows ( Ph o x i n u s p h o x i n u s ) and
showed that corrleation bet ween fishes' instantaneous velocities increased
wi t h school size and as interfish distances decreased. Also leader-follower
relationships are common in two-fish schools, but t hey are not seen for
schools wi t h a number of fish (Shaw ( 785) ) . The number dependence in
swarming behavior is also observed in insect swarms (Gol dsmi t h e t al .
(51)).
Breder ( 112) described that a distinct wave of movement passed
t hrough a school of fish when only fish on the peri phery were agitated
by a sound. The speed of this wave was slower than that of sound in water.
Radakov ( 227) st udi ed t he behavior of fish school to frightening stimuli.
It was observed that a wave of agitation propagated t hrough a school of
At h e r i n o mo r u s . Ther e was a rapidly shifting zone in which the fish reacted
to t he actions of their neighbors by changing their positions. The speed
of propagation of this wave of agitation reached 11.8-15.1 m/see in t he
experiments. Thi s speed was much higher than the maxi mum speed of
forward movement of individual fish (about 1 m/see). These observa-
tions suggest a soliton-like wave woul d be propagating t hrough a school
in agitation.
We will briefly discuss a simple model for t he wave of agitation in
fish schooling. Consider a one-dimensional school of an infinite number
of fish. Let y j ( t ) be t he position of t he j - t h fish in t he school. Ignoring
t he frictional and random forces t he equation of mot i on of the j - t h fish
reads
m~j = F ( y ~ - - y j - l ) - F ( y j . 1 - y j) (VII. 39)
when y j - - y ] _ l = y j + l - - y ~ = r e (equilibrium distance), t he fish school is
either at rest or moving wi t h a constant velocity as each fish is separated
from its nearest neighbors exactly by t he distance re. I f we define
7 8 A. OKUBO
r j -- Y~+i-- Yj (VII. 40)
as t he rel at i ve di st ance bet ween j and j + 1 fish, ( VI I . 39) is expr essed by
mrs, ---- --F(rj+i) +2F( r j ) --F(rj _i ) (VII, 41)
Th e us e of t he f or ce (VII. 31) r ewr i t es (VII, 41) as
m r j = - - B ( e - ~ r J+l-- 2 e - b r J ~ - e - b r J -1) (VII, 42)
To d a ( 2 J 5 , 2 1 6 ) f ound i ngeni ous exact sol ut i ons of ( VI I . 4 2 ) i n t e r ms
of t he el l i pt i c f unct i ons. A par t i cul ar sol ut i on of ( VI I . 4 2 ) gives a sol i t ary
wave wi t hout changi ng its shape, i . e . " s ol i t on" :
m
e-Or1 -- 1 = -~b-V 2 sech 2 ( k j + v t ) ( v i i . 43)
wi t h a speed of pr opagat i on c t hat
c = v / k = ( B b / m ) i / ~ ( s i n h k ) / k (VII. 44)
and k and v are wavenumber and f r equency of t he sol i t on, r espect i vel y.
Th u s a s ol kon of l ong wave pr opagat es muc h f ast er t han t hat of s hor t
wave.
60
4O
~Y
u_ 20
Fig. 15.
exponential distribution.
Spot t ai l shiner

@
2 4 6 8 10 11 14
School size
Comparison Of the school size distribution of the shiners with the
@, data; - - , theory.
Al t hough a group of fish exhibits a characteristic spatial orientation
in schooling, some groups are "pol ari zed" (Radakov (227)) constantly
and ot hers only occasionally. Breder (228) distinguishes facultative, or
occasional, from obligate, or constant, schoolers. Seghers (229) st udi ed
t he behavior of facultative schooling in t he spottail shiner (Notropis hucl-
sonius) in Veronica Lake, Ontario, Canada. The juvenile shiners (mean
size is 1.5 cm) are facultative schoolers. Schools consist of two or more
fish of similar size swimming t oget her usually in a polarized fashion.
Single fish are not affiliated wi t h a school and are usually farther than
50 cm from t he nearest shiner. They may, however, eventually form or
j oi n a school. Few schools maintain their integrity for more t han a minute.
Thus the composition of schools is highly variable. Fi gure 15 shows t he
frequency (or probability) of occurrence of a particular school size. The
group (school) size varies from 2 to 49 individuals, but in 39. 8% of t he
observations t he shiners were solitary. Al most 50% of the shiners were
di st ri but ed as solitary fish and schools of less than five fish. The school
size distribution of t he shiners is well represent ed by t he geometric dis-
t ri but i on or exponential di st ri but i on
f l = (ec-1)e -c~ ( i = 1,2 .... ), c>0 (VII. 45)
As discussed in Chapt er VI.4, t he geometric distribution is a consequency
of maximizing group ent ropy under t he constraints that bot h t he total
number of animals in grouping and t he total number of groups are held
constant.
Usi ng sonar Smith (230) observed fish schools in waters off sout hern
California, and he obtained the frequency distribution of school diameter.
The di st ri but i on showed a well-defined peak frequency at a diameter of
about 15 m and decreased in an exponential like manner t owards larger
and smaller diameters. Thi s simple, and relatively stable, distribution
stimulated Anderson (88) to const ruct a mathematical model for t he size
of fish schools.
Anderson' s model starts wi t h (VI.44) for t he stochastic equation of
t he number of fish in schooling. To express t he frequency di st ri but i on
in t erms of school diameters Anderson uses t he relation
n = 4 PXZ( 1 h( t ) ) (VII. 46)
where n is t he number of fish in schooling, p is t he average number den-
sity of fish per uni t horizontal area, X is t he diameter of a school wi t h a
8 0 : A . O K U B O
disklike shape, and h ( t ) is a vari abl e representing stochastic variations on
t he density. Combining (VI.44) wi t h ( v i i . 4 6 ) and assuming [hi is much
smaller t han unity, Anderson obtains t he equation for X
d~-t--= ~ p X 2 + m + X f ( t ) (VII. 47)
where ~:, ~ , m S, and r are constants and f ( t ) is white noise of zero mean
and uni t variance,
The probabi l i t y density function p ( x , t ) for t he syst em having a
diameter X at t i me t is obeyed by t he following Fokker-Pl anck equation
Op 1 0 I { 4 5 _ O / 4 r X ' , q ~
8 t 2 a X ~ - f l X - m ~ (VII. 48)
- + ) / q
whereas ( 8 8 ) used t he Stratonovich calculus (Goel and Ri cht er-Dyn ( 2 3 7 ) ) .
The steady-state probability density function is given by
P , ( x ) = k X 2 ( X Z + c ) -2(1+b) exp - { 2 ( a + b c ) / ( X Z + c ) } (VII. 49)
where a _ ~ 8 / ~ r p m 2, b==_~/4m 2, c=_4r/~rp, and k is a constant det ermi ned by
t he normalization condition O f p s ( x ) .
Eq. (VII.49) can be compared favorably wi t h Smith' s ( 2 3 0 ) data by
appropriately choosing t he parameters a, b, and c. Anderson also in-
vestigated t he sensitivity of P s ( x ) to variations in t he model parameters.
For small values of t he entrance rate into schools, ~, small diameters are
favored, and as t he entrance rate increases a wide range of large schools
is favored. The entrance rate t ends to increase wi t h increase in t he stock
popul at i on of an area. Thi s suggests that large stocks woul d contain a
wi de range of school sizes. The coefficient for t he average exit rate of fish
from a school, fi, has a si gni fi cant effect on P s , wi t h large values favoring
a narrow range of small diameters and small values favoring a wide range
of large diameters. The randomness in t he schooling process is parame-
terized by m S. For small levels of randomness, i . e . small m S, t he pr ob-
ability distribution converges on t he deterministic steady-state diameter
Xo whi ch is given by
Xo = 2 ( ~ / ~ p i ) - ~ ( v i i . 50)
For large values of m S t he di st ri but i on spreads away from t he deter-
ministic value. The density of fish, p, has a strong effect on P s . Thus a
dense grouping of fish, correspondi ng to large values of p, favors a narrow
range of small school diameters, while a low density favors a wi de range
of large diameters. The density of fish in schools is inversely related to
t he fish length, so that large fish (low value of p) should have a wide prob-
ability distribution of large diameters and small fish (high value of p)
should develop a nar r ow distribution of small diameters.
4 . B i r d F l o c k s
Ecological and behavioral aspects of bird flocks have been investigated
rat her extensively. They include t he problems of foraging success and
predat or detection for bi rd flocks (Pulliam ( 7 1 6 ) ; Elgar and Catterall ( 117) ;
Har t and Lendr ew ( 7 2 0 ) ; Herrera ( 2 3 2 ) ; Pulliam e t al . (233); Bertram
( 234) ) . I n t he context of animal dynamics, however, studies on bird flock-
ing have been limited to aerodynamics of bird flocks (Higdon and Corrsin
( 2 3 5 ) ; Major and Dill (236)) and energetics of locomotion (Alexander
( 56, 5 7 ) ) .
Recently Potts ( 237) has analyzed slow-motion films of dunlin ( C a l i -
dr i s alpina) flocks to st udy how bi rd flocks can execute abrupt maneuvres
with very precise coordination. Potts shows t hat a single bird may initiate
a maneuvre whi ch spreads t hrough t he flock in a wave.
Major and Di l l (236) developed a stereoscopic camera t echni que to
st udy t he t hree-di mensi onal st ruct ure of flying flocks of small birds (dun-
lin and starling) in t he field. Winds played an i mport ant role in t he flock-
ing behavior of bot h species of birds. When wi nd velocity was greater
t han about 20 km/hr, bot h species t ended to fly close to t he ground or
water, and to spread out horizontally more t han vertically, and flock
st ruct ure appeared to become looser or less dense.
When predators such as perigrine falcons ( Fal c o peregrinus) attacked
a dunl i n or starling flock, it quickly coalesced into a nearly spherical
"ball" of birds and appeared to increase t hei r flight speed. At times t he
flock "pul sat ed", e . g . expanding and contracting in spatial dimension,
presumably as i nt erbi rd distance varied.
Ther e appear to be striking similarities in t he st ruct ure and behavior
of bot h fish schools and bi rd flocks. Thus analyses of t he t hree-di men-
sional st ruct ure of schools of mi nnows (Pitcher (220)) found that nearest
neighbors in schools t ended to be behi nd and possibly below t he reference
fish. Thi s t endency is similar to t he distribution of nearest neighbors in
dunl i n flocks (Major and Dill (236)). The changes t hat occur in flock
st ruct ure when predators appear are also very similar to those observed
in schools of t hree species of prey fishes when attacked by predat or fishes
(Major and Dill (236)). The general characteristics of polarization and
8 2 A. OKUBO
synchronization of bi rd flocks are in fact not unlike those of fish schools.
Ther e may be a fundament al difference bet ween fish schooling and
bird flocking, however. I n fish schools individuals are reacting more or
less to t hei r immediate neighbors owing to t he limitation in vision, while,
in bird flocks, individuals are able to view a wi der part of flock motions.
Schnei der (238) studied t he location and st ruct ure of pre- and post-
foraging seabird aggregations in relation to oceanic fronts. Seabird aggre-
gations oft en coincided with chlorophyll maxima in t he sea water, A
direct linkage t hrough zooplankton and fish is suggested.
5. Mammal i an Herds
The great herds of wildebeest, buffalo, and zebra among others in t he
Serengeti are a phenomenal sight (Sinclair (239)). Herdi ng for these
animals is oft en i nt erpret ed as a defense mechani sm against predators.
Group living increases predat or detection, cooperative defense, and
feeding efficiency, but at t he same t i me increases intraspecific compet e-
tion such as for food, water, and resting sites (Bertram (23d)). Group
living species t end to form stable social groups that are oft en composed of
relatives; groups of social primates are usually based on stable associa-
tions of related females.
I n ot her times even group living species can form "r andom associa-
t i on" (Lot t and Mi nt a (709)), in particular adult females are known to
be solitary once t he period of nearly continuous association bet ween
female and her offspring ended.
Wirtz and Lorscher (2dO) presented data on t he group size frequencies
of t he eight most common antelope species in Lake Nakuru National
Park, Kenya. These species are steenbok (Raphicerus campestris), Kirk' s
dikdik (Madoqua Ki r k i ) , bushbuck (Tragelaphus scriptus), Bohor reedbuck
(Redunca redunca), mount ai n reedbuck (Redunca f u l v o mf u l a ) , Thomps on' s
gazelle (Gazella t homsoni ), impala (Aepyceros melampus), and wat erbuck
(Kobus ellipsiprymnus). Among t hese antelopes Bohor reedbuck, moun-
tain reedbuck, and wat erbuck are grazers, steenbok, bushbuck, and dikdik
are browsers, and Thompson' s gazelle and impala are mi xed feeders.
The t hree browsers have significantly lower mean group sizes t han t he
ot her species.
Wirtz and Lorscher' s data showed that singletons (single animals)
were prevailing and t he group size distribution was generally a decreasing
funct i on of t he group size, although t he proportion of singletons appeared
to decrease and t he maxi mum group size to increase with increasing
40
20
0
80
60
60
5 10 15
4O
"~ 20
0 2 4 6 8 ' 10
3O
2O
L
G
/
Group size
Fi g. 16. Co mp a r i s o n o f g r o u p si ze f r equenc i es o f T h o ms o n ' s (a), b o h o r
r eedbuck (b), a nd wat er buck (c) i n an East Af r i can nat i onal par k wi t h t he
geomet r i c (or exponent i al ) di s t r i but i on. (~), dat a; t heor y.
84 A. OKUBO
population density.
Somewhat surprisingly the group size distribution of the eight ante-
lopes is represented generally by a negative binomial distribution and even
better by a geometric distribution. Figures 16a-c show some examples
of fitting Wirtz and Lorscher' s data to a geometric distribution.
Another example of the geometric distribution is seen in American
bison (Bison bison) herds. Lott and Minta (709) recorded individual as-
sociation in determining group stability in a population of 400 freely
roaming American bison on Santa Catalina Island, California over a 44
month period excluding cow-calf bonds and rut periods. Figure 17 demon-
strates a close fit of a geometric distribution to the observed frequency
distribution of the size of groups of mature cows during the non-rut
period. Although the authors (Lott and Minta (709)) represented their
data by a negative binomial, the geometric distribution seems providing
a better fit.
The size of herds of the African buffalo (Syncerus caffer) in May
1968 in the Serengeti varied from 50 to more than 1,500 animals (Sinclair
(239)). Many of the smaller groups were temporary aggregations with
splitting and joining, while the large herds appeared to be stable units.
300
200
o
e -
6
Z
10C
American bison
Fig. 17.
bison herd sizes with the exponential (or geometric) distribution.
- - , theory.
O O
5 10 15 210
Size
Comparison of the observed frequency distribution of the American
(~), data;
60 African buffal o
4O
~d
6
z
2 0 o o e e
0 2 0 0 4 0 0 6 0 0 8 0 0 1, 000 1, 200
Size
Fig. 18. Comparison of the frequency distribution of herd sizes of the
African buffalo in the Serengeti population with the exponential (or geometric)
distribution. @, data; - - - , theory.
Small herds up to about 10 individuals are family groups. However, with-
in the large herds of more than 50 individuals, there were no obvious
divisions into subunits or families except the basic one of a family with a
calf and possibly with an older offspring. Thus the dynamics of grouping
of African buffalo is expected to obey the models described in Chapter
VI. In fact the observed distribution can be fitted to a geometric distribu-
tion (Fig. 18). Except for the large herds (size>800) which appear to
form stable units, the observed frequency is closely described by the
geometric distribution with a mean of 145 animals per group.
Another example of the geometric distribution is seen in the case of
desert bighorn (Ovis canadensis) herds. Deming (1953: cited in Hansen
(241)) recorded group sizes of desert bighorn in Desert National Wildlife
Range, Nevada between 1947 and 1952. These data show that bighorn
are encountered most frequently solitary, or in groups of two or three.
Groups of less than nine are most common. Again the group size fre-
quency is well represented by a geometric distribution (Fig. 19). The
universality of the geometric frequency distribution for group sizes among
freely roaming herbivorous mammals may look strange, but it might be
a result of maximizing group entropy under the conservation of the hum-
86 A. OKUBO
2 O0
D e s e r t b i g h o r n
o
"= 1 0 0
"8
6
7
o

o Q
Q Q 0
O( 10 2 0 3'0
Size
Fig. 19. Comparison of the frequency distribution of herd sizes of the De-
sert bighorn with geometric (or exponential) distribution.
ber of gr oups. I n ot her wor ds t he ani mal s t e nd t o f or m r a ndom associa-
t i on except f or f emal e- of f spr i ng bonds, st r ong def ense behavi or and so
f or t h. Ra ndom associ at i on may faci l i t at e f or each ani mal t o mat ch its
physi ol ogi cal needs at any mome nt wi t h t he needs of ot her pot ent i al co-
gr oup me mbe r s at t he same t i me ( Lot t and Mi nt a ( 1 0 9 ) ) .
Of cour se non- r a ndom associ at i ons do occur i n mammal i an gr oups.
Th u s humpback whal es off Ne wf oundl a nd i n s umme r f or me d gr oupi ng
cont ai ni ng f r om 1 t o over 10 i ndi vi dual s ( Whi t ehead ( 2 4 2 ) ) . Th e size of
f eedi ng gr oups was cl osel y r el at ed t o t he hor i zont al scale of t he pr ey
school , wher eas nonf eedi ng whal es wer e general l y f ound i n pai rs. Smal l
gr oups wer e consi der abl y mor e st abl e t han l arger ones; t hi s is also a
cont r ast t o t he ant el ope her ds. Tya c k and Whi t ehead ( 2 4 3 ) r epor t ed fast
movi ng gr oups cont ai ni ng t hr ee or mor e adul t wi nt er i ng humpback
whal es off Hawai i and on Si l ver Bank i n t he West I ndi es. Ma ny of t hese
gr oups had a defi ni t e s t r uct ur e wi t h a cent r al nucl ear ani mal (female),
with or wi t hout a calf, surrounded by escorts (males) who compete for
proximity to t he nuclear animal.
S UMMARY
Groupi ng of animals is a natural phenomenon in whi ch a number of
animal individuals are involved in movement as formi ng a group. Ex-
amples are insect swarms and fish schools. I n this article an at t empt is
made to describe t he mot i on of groupi ng individuals kinematically as
distinct from simple diffusion or random walk, to model t he grouping
on t he basis of dynamics of animal motion, and to i nt erpret t he grouping
from t he standpoint of advection-diffusion processes. Also present ed
is dynamical modeling for t he group size distribution as a result of amal-
gamation and splitting processes of groups.
Examples of animal grouping are described in detail. They are insect
swarms, zooplankton swarms, fish schools, bi rd flocks, and mammal
herds. The present ed mathematical models are compared with data of
t hese animal groupings.
Acknowledgments
I n t he preparation of this article many friends and colleagues have helped
me by providing valuable suggestions, encouragement, and stimulation;
particular thanks are due to Joe Ackerman, Ji m Anderson, Huai Chiang,
Cay Craig, Hans Dam- Guer r er o, Eileen Hofmann, Pet er Kareiva, Mari-
anne Legier-Visser, Mike Levandowsky, Si Levin, Ji m Mitchell, Ci ndy
Monaco, Tom Powell, Jerry Schubel, Nanako Shigesada, Sharon Smith,
and Ei Teramot o (alphabetically last but most i mport ant to convince t he
editorial board of Advances in Biophysics of consideration for publication
of my work). Last but not least I wish to t hank Eileen Goldsmith for
her skillful and patient t ypi ng of t he manuscript. Contribution No. 528
of t he Mari ne Sciences Research Center, State University of New York
at Stony Brook and Cont ri but i on No. 093 of t he Ecosystems Research
Center, Cornell University.
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Received for publication November 9, 1985.

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