NATURAL

SELECTION

AND NEOTENY

by
R. F. EWER
(Department of Zoology, Rhodes University, Grahamstown, South Africa)
(Received 4. V. 1959)

The centenary of the publication of DARWIn'S and WALLACE'S views

on evolution has, naturally, been the occasion for the appearance of numerous
articles and books .dealing with evolution in general and in particular :how
views on the subject have changed and developed since DARWIn'S .day.
Every zoologist is familiar with the fact that, although our understanding of
the causes underlying variation has advanced greatly in the last ioo years,
DARWIN'S concept of natural selection as the force conferring direction on
the evolutionary process remains basic to our comprehension of that process.
One might therefore expect that, with the advantage o.f a century's extra
knowledge, an evolutionary outlook superior in its insight and penetration
to DARWIN'S own would form the normal background of contemporary
studies in phylogeny and comparative anatomy and physiol.ogy. This, however, is far from being the case. Re-reading the "Origin" ioo years after
its publication one is impressed by the fact that DARWIN had a deeper
insight and a broader appreciation of the consequences of his theory than
many present day zoologists. "T~he reason for this seems to be quite simple:
DARWIN always saw animals as alive, making their livings in various ways
in different environments, and above all, in competition with each other.
To him the interaction of animal with animal was one of the most important
factors of all, one which was more often than not decisive for the future
course of events - - in his own words, "the relation of organism to organism
in the struggle for life being, as I have already remarked, t:he most important of all relations". By contrast, in contemporary studies an animal is
treated only too .often as a set of anatomical diagrams and an "evolutionary
approach" consists .of turning one ,diagram into another, by a series of
intermediate pictures, each of which differs only slightly from its predecessor. This might be useful training for an animator for the film cartoon
industry, but it is not zoology. The point that I wish to stress .'here is that
if we are, on the one hand to comprehend a known series of phylogenetic
changes, or, on the other, to make reasonable guesses about ones which
Acta Biotheoretica, XIII

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R.F. EWER

remain unknown, a different approach is required. It is necessary to understand what was the selective pressure behind any evolutionary c h a n g e - to
ask what advantage did it confer; what factors decided that the change
should be this particular one and no other.
Before expanding this main theme, it may be profitable to. digress slightly
and consider the .origins of the tendency to think in terms of dead .diagrams
instead of live animals. I believe that two factors have been responsible.
I. Immediately following the publication of DARWIN'S work the main
question was "has evolution really occured? If it has, then fossils should
provide us with graded series of ancestor and .descendant, and comparative
anatomy should tell the same story". So the tracing .of series became allimportant and was carried out with great enthusiasm and success: gradually
(although not without argument and disagreement on the way) the evidence
in favour of evolution became .overwhelming. And now, although we have
reached the stage where it is necessary to do more than merely trace a
series, to ask "how" instead of ",whether", the old habit dies hard.
2. The second factor is a reaction against over-simplified teleology, a
reaction which :first became apparent in the field o.f behaviour. When one
reflects that ROMANES could, in all seriousness write that the tendency o.f
a moth to fly into the flame of a candle "must be set down to mere curiosity
or ,desire to examine a new and striking object" the necessity for some
corrective reaction is obvious. Such an "explanation" is not only grossly
unwarranted anthropomorphism; it is also useless as a guide to. further
investigation: when one has guessed the animal's motives and emotions on
the unjustified assumption that they must be like our .own, the enquiry is
finished. Similarly, in the field of evolutionary theory itself came the ever
clearer discrediting of LAMARCK'S views in which both the desires and
mental efforts of the animal and its physical activities were alleged to
affect its heredity. The reaction against teleology was both natural and
necessary but the trouble was, it went too far. In rejecting simple teleology
it rejected teleology altogether; and in pointing out that no evidence was
forthcoming for the inherited effects of the animal's own activity it concluded that the latter was an irrelevancy in evolutionary studies. In this it
threw away two of the most powerful weapons in our theor.etical armoury,
and provides a classical example .of what I would like to christen "bathwaterism". The concept of selective advantage is based on a consideration
of "ends", although it does not place them inside the consciousness either
o,f a creator o.r of the evolving animal and, as will become clear later,
activities are very far from being irrelevant.
'The legitimate use of teleology, fMlowing this exaggerated avoiding re-

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action, has gradually come to be, if not exactly enthusiastically appreciated,

at least accorded toleration. Thus, although we would no longer say that
the birds evolved wings in order to fly, .only a few would object to the
formulation that in the avian lineage a positive survival value was associated
with increasing ability to fly and mutations tending to convert the fo.re
limbs to wings were therefore preserved and accumu,lated. Similarly, in
behaviour studies it is now perfectly respectable to consider the adaptive
significance .of any particular piece of behaviour.
This somewhat grudging use of the concept of survival value is, however,
not sufficient. It leaves out the live animal and concentrates too much on
what it is, too little .on what it does. It would appear to be glaringly obvious
that what an animal does, o.r tries to do, can .determine what characters are
of survival value, i.e. can decide the direction .of natural selection--and
yet even today this indirect action is frequently ignored and the only possible effect .of an animal's activity which receives consideration is the .direct
Lamarckian .one: for example CARTER (1958) writes that "the effects of
activity can only influence the evolution .of the race if they are inherited
and passed on to the next generation". If evolution results from natural
selection conferring direction on a non-directional process of genetic mutation, it s:hould be clear that factors which affect selection rather than
mutation are the ones which determine the direction of evolution: factors
affecting mutation rate may speed up, or slow down, but will not change
the direction of the process, except indirectly. Moreover, if we wish to
decide whether something has an effect on evolution we must ask the two
questions: (i) does it affect the genetic mechanism? (ii) .does it affect the
selective forces impinging on the organism? To return to the question of
the birds' wings: had the ancesto.rs of the birds used their fore limbs only
for terrestrial locomotion, what hope of preservation would there have been
for any mutations tending to. convert them into wings?
The importance of the activities and behaviour .of the animal in determining its evolutionary :fate is most obvious in cases where the animal
is in a position to make direct use of a structure in a number of different
w a y s - for example, to use its limbs for climbing, running, digging or
swimming - - but even physiological characters will also be affected. A change
of diet will a'lter the selective value .of digestive enzymes: a higher level of
activity or a tendency to explore environments poor in oxygen will alter
the selective value of changes in concentration or loading tension of blood
pigments. In fact, if FORD (I957) is justified in saying that a truly
"neutral" character is virtually an impossibility, then it becomes rather
difficult to imagine that any o.f the characterstics into. Which it is possible

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1t. F. EWER

to analyse the whole that constitutes a ,live animal can be unaffected by its
activities.
This does not, of course, imply that behaviour is somehow exempt from
and superior to the rules that govern the evolution of structure. Behaviour
is the manipulation of the effector organs brought about by the activities
of the nervous system, which reacts to "information" co.nveyed to it both in
the past and in the present about changes taking place both inside the animal
itself and in the external environment. Changes in sensory equipment, or
in effector organs will, of course, affect behaviour, but progressive change
involves mutations affecting the structure and mode of action of the directing
central nervous s y s t e m - - a n d these will follow the same pattern as those
concerned in the evolution o.f any other functional organ. The relationship
between activity and anatomy will, however, be asymmetrical for two. sorts
of reasons. There is firstly the one already mentioned, which can be summarised by saying that while a genetic change causing an animal to take
to swimming wilt result in accumulation of genes making for webbing o.f
the digits, a mutation causing slight webbing in a non-swimmer will not
cause accumulation of genes making for the habit of swimming. Secondly,
it must be bo.rne in mind that an evolutionary change .does not have to. "wait
for the right mutations to turn up": the first advance wil'l always be made
on the basis of changes in frequency and recombination of genes already
present in the population 1) with new mutations bringing up the rear by
continually replenishing the pool of variability.
This evo.lutionary plasticity at the level .o:f the population applies to all
characteristics, both structural and behavioural. Behaviour, however, generally has much more plasticity at the level of the individual than has
structure. The adaptability of behaviour to varying environmental conditions
gives it a "factor of safety" allowing an immediate behavioural response to
be made at .once to a changed situation, without the necessity of waiting
for appropriate changes in the genetical structure o.f the population. Thus
behaviour will tend to be always one jump ahead of structure, and so to
play a .decisive role in the evolutionary process.
It is true that since behaviour does nol fossilise it will rarely, if ever,
be possible to .demonstrate that changes in habits did, in any particular instance, precede structural change. In certain cases, however, such an assumption does appear to provide the simplest explanation of the o.rder in which
different structural changes occurred. As an ill,ustration we may consider
I) WADDINGTON'S(1956) demonstration of the "genetic assimilation" of the bithorax
pheuotype shows how very great a departure from the norm is capable of being
produced in this manner.

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the case of the evolution of the skulls and teeth of some of the African
Suidae. The living species include the very omnivorous Bushpig, with short
low-crowned bunodont cheek-teeth, rather like those of a .domestic pig, and
the largely grass-eating Warthog, with very specialised elongated, highcrowned, multi-cusped molars. The Warthog chews with a specialised sideways grinding movement. This statement is based on personal observations,
but might 'have been inferred from the details of the skull architecture in
relation to the jaw muscles an,d from the Idirection o.f the wear marks on the
canines made as the lowers move across the uppers. In addition, a peculiar
polish on the outer surface of the lower canine results from the way in
which the Warthog moves its head as it selects out the grass tips which
form its favourite food from amongst the weeds (See EWER, I958 for
details). I have not been able to watch Bushpig chewing, but the skull structure, muscle arrangement and the wear on the canines indicate that there
is no highly developed grinding actio.n. Many of the South African fossil
pigs, belonging to a number of different lineages, show stages in the evolution of complex grinding teeth, like those of the Warthog, from an
originally unspecialised condition not unlike that occurring in the Bushpig.
On the hypothesis of "habit a jump ahead .of structure" the sequence of
events would be expected to be as follows. First a cmhange of habit, so that
grass starts to become a more important component of the diet. This will
confer selective advantage on any mutations converting the unspecialised
cheek teeth into more effective grinders, and such mutations will now be
preserved and accumulated. Once this process has advanced to a significant
degree, but not before, there will be selective advantage in a more effective
grinding with the iaws as ~/he food is chewed. Once this habit ,develops, but
no.t before, there will be selective advantage in alterations in skull architecture
facilitating a grinding jaw a c t i o n - - b u t until the animals d o .do the best
they can to grind with the equipment they have there will be no tendency
to preserve "'favourable" (actually only potentia'lly favourable) mutations.
One would therefore expect that in the fossil pigs the teeth should be
"ahead" of the skull architecture.
Unfortunately, while fossil teeth are numerous, skulls or reasonably large
fragments of skulls are rare, and in only two cases is there sufficient
material .on which to test the theory.
Potamochoero;~des shaw{ (Dale) occurs abundantly in the famous
Australopithecine deposits at Makapan. In this species changes in the teeth
away from the simple bunodont condition are clearly in progress, but the
skull arc~hitecture shows no signs of grinding adaptions, and the wear on
the canines suggests that the jaws were not used with a grinding action.

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This species would therefore appear to represent an early stage in the

development of grass-eating specialisation. A change .of diet to include a
higher proportion o,f more abrasive food would appear to have taken place
and this has conferred selective advantage on the ,development of more
high-crowned teeth with a larger surface area. Only when a critical point in
this process is r e a c h e d - - d e t e r m i n e d by a balance between increased hypsodonty and increased rate of tooth wear, and apparently not yet attained
in P. s h a w i - - w i l l the change over to a grinding action of the jaws become
advantageous and corresponding changes in skull architecture will be expected
to follow.
Potamochoerus antiquus Broom, belonging to a ,different lineage, is a
more advanced form in which the cheek teeth are very nearly as specialised
as those of the extant Warthog. Moreover, a lower tusk shows signs of the
typical Warthog type of polish, so it is a fair inference that this species
was already plucking its grass from amongst the weeds like its modern
relative. The skull shows exactly the same sort of grinding adaptations as
does that of the Warthog, but ~hey are developed to a degree which is just
about half way between the unspecialised Bushpig arrangement and the
modern Warthog. So here we have the other end of the s t o r y - - a species
in which the teeth, now almost at the most specialised stage known, are still
(as the theory demands) in advance of skull architecture.
In other cases .differences in structural details in related species can be
understood only on the same a s s u m p t i o n - - t h a t the primary factor in t~he
evolution of the structural differences was a behavioural difference. The
clearest example of this known to me is a particularly elegant analysis of the
evolution of different stridulatory mechanisms in related species of scorpions
by ALEXANDER (I960).
BOLK'S (I922) theory of the evolution of the mammalian method of tooth
replacement provides an example of how one may be led astray by concentrating on diagrams and forgetting the animals which they represent.
The discussion that follows refers only to the cheek teeth, because ~he
functional considerations which apply to these are concerned with their
chewing efficiency and are thus not relevant to the incisors and canines.
The typical reptilian repeated replacement of alternate teeth is i,deal for
an undifferentiated dentition, but is quite unsuited to complex hetero~donty,~
where the whole cheek-tooth battery forms a functional unit. In these circumstances replacement impairs the efficiency of the whole system. The
conditions necessary for an efficient masticating apparatus are that upper
and lower teeth should maintain specific occlusa'l relations, that the teeth
should be firmly anchored in the jaw and that gaps in the series should

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not occur. BOLI<, quite correctly, assumed that with the .development of
mastication there would be selection for reducing to a minimum fhe number
of replacements, bo.th because they create gaps when a tooth is actually
shed and because the anchoring of the tooth is weakened for some time
before the actuM shedding. In his theory, reduction in ~ehe number o.f
replacements is thought to have proceeded until each germ produced one
tooth only. I n the final stage the members of the exo- and endo-stichous
series, instead of erupting side by side come to replace each other, forming
the milk and permanent series respectively. This theory lo.oks so well in a
series of diagrams that it still graces the pages .of a very recent text-book
(GlaASS~, I954) with no more critical comment than that it is "speculative".
It will, however, not bear a closer examination. In the first place, this
theory requires that, at the last step, the animal's jaw must be filled by
half the number of teeth that had occupied it previously. Even if we assume
that this need not have .'happened all at once, but could have occurred progressively from front to rear (or vice versa), this helps very little, for the
theory still requires that a complex .dentition, evolved and .selected as a
functional unit, must continue to work effectively when one after another
alternate teeth are removed from the original series, and they in their turn
must constitute a functional permanent dentition. This, however, is not the
only difficulty. What BOLK failed to appreciate is that distichial replacement precludes the development of an efficient masticatory system. True,
it keeps a set of teeth always functional, but it does so only by continually
bringing up reserves between each pair of teeth actually in use. The presence
of these not-yet-functional reserves is incompatible with the conversion of
the first set into an efficient chewing unit, which demands that the surfaces
of the who.le battery of cheek teeth shall be on a single level, without gaps.
T~he more effective the individual teeth become as grinding tools, the more
necessary it is for the inequalities of level resulting from distiohial replacement to be abolished. In other words, to assume that distichial replaeement was preserved during the evolution of complex chewing teeth is to
assume that the latter were evolved in circumstances in which it was impossible for t~hem to w~ork . There must therefore have been selection not
only for fewer replacements and for a smaller number of more complex
teeth but also for .obliteration of the distinction between the members of
the two series, and distichial replacement can have been preserved only in
species which did not chew very effectively.
Since it i's obviously .desirable to keep to a minimum the number of teeth
non-functional at any one time, one would expect that there would be a
gradual transition from alternate replacement to replacement pro,ceeding

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EWER

gradually from front to rear .of the jaw. The following changes would therefore be expected to .occur, more or less simultaneously:-I. Increase in size and complexity of individual teeth and reduction in
number. This last would most simply be achieved by suppression of the
most posterior germs.
2. Exaggeration of the antero-posterior gradient thus established, so that
teeth .differentiate in order from front to rear and the distinction between
exo- and endostichous series is lost.
3. Reduction in number of replacements, so that after a certain time in
the individual ontogeny no further teeth are formed. This, coupled with
the antero-posterior gradient, will finally result in the mammalian condition where anterior germs produce two. teeth but posteri.or ones, starting
late, produce only one.
Since the distinction between exo- and endostichous families has been
lost early on in this complex process, the question as to which reptilian
tooth families the mammalian milk and permanent series represent
ceases to have any meaning. Palaeontological evidence whioh is not in
agreement with BOLK's theory is now steadily accumu'lating (CRo~PTO>r,
1955, KER~aACtr 1956), and there is little doubt that his .diagrams are on
the way to extinction: the surprising thing is that they should have survived so long, when the Iehe.oretical .deficiencies of he assumptions on
which they are based should have been obvious to anyone able to see the
animal behind the diagram.
Another instance of failure to think in terms of live evolving animals is
shown in the too facile invocation of the concept of neoteny which appears
to be at 'present in vogue. This tendency is clearly shown in an article by
DE B E E R and SWI~TOaT (I958), publis.hed in the set of essays presented
to D.M.S. Watson under the title of "Studies on fossil vertebrates". Here
almost any .structural resemblance between an adult of one fo.rm and a
development stage of another is forthwith attributed to "neoteny", without
either a close scrutiny of the facts, a di'scussion as to how or v~hy the change
is thought to have occurred o.r a consideration .of exactly what the term
"neoteny" implies. It may therefore be profitable firstly to consider this last
point, and then proceed to examine in a more critical manner some cases
of alleged neoteny.
It is not sufficient to ,define neoteny as the attainment of sexual maturity
by a larval form, with the result that a .descendant resembles in its adult
form a developmental stage of its ancestor: it is necessary also to consider
how this neotenic condition could have been evolved. Modern genetical
work has tended to emphasise the importance in evolution of cumulative

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small changes, and the improbability of large mutations being wholly

favourable in their effects. This point is very ably treated by FORD (1957),
to w~om the reader is referred for details. If this is correct, then it is
unlikely that neoteny could be achieved at a single mutational step, and a
gradual change as the result of cumulative mutations of small effect is
more probable. There are two slightly .different ways in which such a
gradual change might occur. The first is what might be described not as
the larva becoming sexually mature but as readaptation of the adult to. the
larval mode of life, with consequent gradual loss of any adult characteristics
which make it less suited to the larval mode of life, or have simply become
functionless. The result will be a gradual elimination of everything which
distinguished adult from larva, with the exception of reproductive organs
and behaviour and possibly larger size.
This appears to have been the way in which neoteny has been attained
in the Amphibia.
Although amphibian metamorphosis is "thyroid triggered", neoteny does
not, in general, seem to have been achieved by the single step of abolishing
the thyroid stimulus (see NOBLE, 1931 for details). Amongst the Perennibranchiata, Necturus and Proteus, Cryptobrcmch'us and Siren all l~ave functional adult thyroids, capable o,f inducing metamorphosis in anuran tadpoles.
The two former have no metamorphosis and are unresponsive to thyroxine
injection: in tthe two latter, metamorphosis involves only the skin and
injection of thyroxine into young larvae causes precocious skin metamorphosis
only. In other words, neoteny has involved loss of tissue sensitivity, and
there is evidence that this was not achieved at one step.
A consideration of the secondarily aquatic anuran, Xenopus, tells the
same story. Many features of this animal have been interpreted as neotenous,
for example the retention of the lateral line canals. Similarly, on the
physiological side, ~he adaptive responses to posterior pituitary hormones,
characteristic of adult Anura but absent in tadpole's, are also reduced or
absent in adult Xenopus (EwER, 1951). Nevertheless, Xenopus remains
a frog, not a tadpole, tailless and with hind limbs even more developed than
those .of most Anura. On our present theory the reasons for this are clear.
The adult has returned to the larval habitat, but unlike the urodeles, it has
no tail and swims perforce with its legs. Selective pressure for better
swimming can therefore result only in bigger and better-webbed hind legs
and Xenopus will never regain its lost tail.
T~he case of /tmbystorna is more difficult. Within the genus, .//. tigrinum
metamorphoses in warm conditions but does not do so when it lives in the
cold at high altitudes, whereas other species are regularly neotenous.

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Thyroxine can induce metamorphosis and one might therefore conclude

that in this case neoteny has been achieved per sc~ltum, by a single genetic
change causing failure of the normal thyroid metamorphic trigger. Some
caution about accepting this explanation is, however, necessary ; the case may
rather be comparable with the polymorphism shown by many species of
butterflies. Here, although the difference between one form and another may
be controlled by a single "switch gene", FISHER (I930) has pointed out that
this is not likely to be a primitive condition, but .has more probably resulted
from the accumulati.on of modifiers which have "built up" the effect of
the major gene in the course .of an evo,lu.tion which has been by successive
small steps. A similar explanation may hold for Ambystoma. Moreover,
physiological adjustments enabling the animal to. function at a reduced
thyroxine level would be required in any case where neoteny is achieved
by suppression of the thyroid trigger. It seems very improbable that a
single geneticM change could have been responsible for the whole process.
This path to neoteny can be followed only by an adult which is capable
of living after the larval manner in the larval habitat. T~here are, however,
many cases in which this is not true: for example, the origin of the
Ctenophora as neo.tenic Turbellaria requires the transition from creeping
adult to pelagic larva. It is not, however, necessary to assume that the only
alternative to "larval readaptation" is the attainment of ne.oteny at one jump
as the result of a macromutation. 'The condition's in which neoteny is
advantageous are those in ~hich the adult is a less successful form than
~he larva. In such circumstances there will be selection for shortening the
adult stage as much as possible, so that its main function is to attain sexual
maturity and reproduce as rapidly as possible. There will thus be a tendency for a shift of emphasis in ontogeny, with the onus of feeding coming
to lie more and more with the larva. Final,ly a stage may be reached in which
the germ ce'lls are laid down in ~he larva, together with sufficient food
reserves to last through the brief adult stage. From this point to the actual
breeding o,f the larva is not a major jump. Moreover, provided adult and
larva are both aquatic and fertilisati.on is external, there is no reason why
gametes shed by a larva should not be fertile with those shed by a metamorpho,sed individual. This avoids anot, her difficulty inherent in neoteny
attained per s a l t u m - that numerous neotenic individuals would have to.
be produced at once if they were to have any chance .of breeding and so
passing .on their new genetic constitutions. T~he alternative of self-fertilising
hermaphroditism, with its reduction of evolutionary plasticity through recombination, is not likely to lay the foundations of a new evolutionary
advance.

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The proponents of neoteny have, very justifiably, been highly critical of

recapitulation theory, DE BEER (1948), for example, denouncing the biogenetic law as "an unfortunate theory, unsound in its premises, illogical in
its .deduction's and disastrous in its results." One may certainly agree that
the idea that phylogeny consists only of terminal addition of .developmental
stages is incorrect and has, in the past, greatly impeded the development
of evolutionary theory. It is also true that a life-history may 'be complicated
by intercalation of a larval .detour arid that subsequent neoteny may result
in the adult assuming the form of the ertswhi~e larva. It would, however,
be rank bathwaterism to conclude that modification of terminal developmental stages can .never occur, to .cry "HaeckeI !" and let sli:p the .do~s of
war without further ado on any theory which assumes such terminal modification. The facts are explicable only if we assume that modification at
any stage is pos'sible, w~hether in the form of an embryonic adaptation such
as amniote embryonic membranes, of intercalated larval stages, or of modification of terminal s t a g e s - - w h i c h must be invoked to explain such
structures as the plates and stalks of adult barnacles or the baleen plates of
whales.
With these points in mind, it may now be profitable to examine critically
one of the cases which is often considered to be an example of neoteny. It
has been suggested by vario.us authors, from GARSTANG (1928) to IBERRILL
(1955) that the chordates are secondarily glorified neotenous sea-squirts.
T,he structural similarity o.f the Ascidian larva to the basic chordate plan
certainly indicates r e l a t i o n s h i p - - b u t is the relationship one of neoteny?
The adult squirt is not able to. live in the larval' manner at all : neoteny by
simple readaptation t.o the larval mode of life is therefore .out of the question.
The second mode .of achieving neoteny implies an ancestral form in which
the adult is relative unsuccessful an,d the larva a successful actively feeding
stage. The condition's in the asci.dians are t~he exact oppisite. Moreover
BE~RILL'S attempt to convince us that the ascidian tadpole ,larva was evolved
within the group and that its structural 'features are necessary adaptations to
achieve the correct site for the sessile adult is not convincing. H e shows us,
indeed, that the larva is well fitted to do what it does do - - but his plea that
the complex tadpole structure is necessary for this end is refutable by the
simple expedient of turning over a few stones or looking underneath overhanging rocks .on any of our local beaches. There you will find not only
ascidians but also sponges, not to mention other sessile forms with ciliated
larvae of one sort or another; and the sponge, with a much less complex
larva appears to be perfectly successfuI in achieving the same situation as
the tadpoled ascidian. It would seem that some variant .of the older view

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expressed by WILLY (I894) i's more likely to be correct; to wit, that the
common ancestor of squirts and chordates was a swimming filter-feeder,
whose locomotion was still far from perfect and whose young stage was
simply a young stage, not justifiably ,described as a larva. Two evolutionary
possibilities are open to such a creature and both appear to have been
realised. The first is the path of increased swimming efficiency, which
resulted in the evo.lution of the chordates. The second is the assumption of
a sedentary habit by the adult. This line gives rise to the ascidians, with
subsequent reduction of the motile young 'stage and elaboration of adaptati.ons of the adult to its sessile mode of life.
The next question that arises is whether neoteny can occur in an animal
which does not possess a true larval stage, with a mode of life ,differing from
that of the adult. Clearly, in such a case what can occur is 'simplification of
structure when altered conditions of existence result in loss of functional
importance of some adult structure or confer selective advantage on a
juvenile characteristic. The altered conditions may be the result of changes
either w'ifhin the animal itself .or external to it. For instance, armour plating
might be lost either becau'se increasing locomotory .efficiency made speed
the most effective means of escaping predators or because colonisati.on of
an island uninhabited by predators rendered armour redundant. Such
changes need not necessarily reflect readaptation to the mode .of life of an
earlier developmental stage and it might therefore be argued that they should
not be called neoteny. On the other hand, neoteny senstr stric~o does include
loss of structures and functions which, owing to changed circumstances, have
become useless to the adult. One might compromise by restricting the term
neoteny to cases where a true larval form is involved and refer to the other
type of change as paedomorphosis. The two differ slightly, not only in the
selective ,directional force postulated to account for the change, but also in
possible end results. Since a larva is, by .definition, an active self-supporting
feeding stage, not necessarily any less structurally complex than its adult,
it follows that neoteny .on the one :hand need not result in any structural
simplification and, on the other, that it can proceed to completion, i.e. to
the production of ~hat contradiction in terms, the sexually mature larva.
An embryo, in contrast, is not self-supporting and is structurally 'less complex than its adu'it: pae.domorphosis will therefore commonly result in
structural simplification, but will never (unless a parasitic habit is acquired)
proceed to completion: certain structures may become paedomorpho,sed, but
the whole animal cannot revert to a pre-functional condition. Moreover, if
paedomorphosis is not necessarily readaptation to the exact mode of ,life of
an earlier developmental stage, it follows that although the stages of reduc-

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tion of an organ may closely resemble ancestral stages, they need not necessarily ,do so. This presents something of a p r o b l e m - - i f the stages of
reduction do not resemble previous developmental stages, does such reduction count as paedomorphosis ?
T,his is not the .only difficulty; there is also the converse problem.
Neoteny has been hailed as "escape from specialisation" (HARDY, I954 ).
It is true that, in species which possess a larva, neoteny provides an escape
from a situation in which the adult is not an evolutionary success. This may
be because it .had become over-.specialised (i.e. so well adapted to .do. one
thing that any change in circumstances leaves it unable to make a living
effectively) but it may equally well be because it was insufficiently specialised and in danger .of being ousted by more efficiently adapted forms, as
is almost certainly what occurred in the case of the urodeles. Nevertheless,
in either case neoteny opens up new evolutionary possibilities (which, of
course, may or may not be realised) by "giving the animal another chance".
Similarly with paedomorphosis, reduction of those adult structures Whose
development restricts the animal to a narrow specialisation constitutes an
escape from specialisation: but reduction proceeding to the ~stage of complete loss of an organ or structure usually spells specialisation, rather than
its opposite. For instance, it is not usual to refer to. the loss of teeth in an
ant-eater, .or of eyes in a cave-d.weller as paedomorphosis; and yet, both
may reproduce earlier developmental stages.
The resolution ,of these difficulties comes from considerations not .of
comparative anatomy, but of function and ecology. The important question
is not "has loss of an adult character occurred and does this result in a
condition resembling an ancestral developmental stage ?" but '"what are the
changes which have permitted or encouraged the loss .of adult structures
and are these changes such as to restrict the animal to a narrowed range of
activities or such as to remove restrictions ?" The primary link in the chain
of events is not the occurrence of paedomorphic changes, but the situation,
whether internal or external to the organism, which creates the .selective
forces resulting in ~he change. Paedomorphosis is not a sort of magic password, automatically opening up new evolutionary possibilities, but only the
recognition, by giving it an official name, of the fact that evolution by
elaboration and development of structures is not necessarily irreversible"
provided the selective forces alter their direction, the elaboration may b,
eliminated or the new structure lost again.
Whether the stages of loss ,do .or do not parallel the stage.s of ontogenetic
.development .does not seem to be particularly important, and we may thus
expect to find cases in Which "escape" from an adult specialisation occurs

I74

R. ~'. EWE~

along a path which does not exactly repeat ontogeny. To converse is also
true: if adult specialisation involves reduction in relative importance of one
structure or organ .during ontogeny, t~hen a new development of that p a r t - actually the reverse .of paedo.morpho.sis- may result in proportions which
resemble those of a juvenile stage. In .other words, resemblance o.r lack o.f
it to a juvenile stage is not necessarily by itself a reliable guide to. what i.s
going on.
With these considerations in mind we may now examine those characteristics of 'human evo.lutio.n which are commonly cited as cases of paedomorphosis. Of these we x~,ill take first the relatively large .size of the braincase.
Although frequently quoted as a paedomo,whic character, no reasons for
regarding it in this light are given, beyond the fact that young mammals in
general and young primates in particular have relatively large heads. But a
zoologist does not say that a dolphin is a fish because it is fi.s'h-sihaped, and
he should be equally cautious and aware .of the :dangers of trusting to
purely superficial resemblance when he is dealing with the problem of
paedomorphosis.
In the case of relative head size, two processes are involved.
(i) Phylogenet~c: in the Primates as a whole there is an evolutionary trend
towards increased size of brain. Later members of a phylogenetic series may
therefore be expected to ~have larger brains than earlier ones. (It is, of course,
not legitimate to compare directly the brains of animals of different sizes;
brain weight/square root of body weight gives the best comparative measure.)
(ii) Ontogen'etic: in ontogeny the brain does not grow at the same rate as
the skeleton; it reaches a large size early in development and increases very
little during the greater part of po.st-natal skeletal growth. Brain size therefore .decreases relative to body size as the individual grows. In man the
relatively large size .of the brain does not result from a general curtailing
of growth, so that juvenile propo.rtions are preserved in the adult. 'The adult
human .skeleton, with its extremely long limbs, in fact shows proportions
very different from ~hose of the youthful stages where the hind limbs are
relatively short. The large head of the adult i.s a result .of a greater manifestation of process (i) than is shown by any other primate. The fact that this
results in brain/body proportions resembling those of a young ape i,s only
an incidental consequence, as discuss.ed above, of the negative allometric
growth of the brain in post-natal development and has nothing to do with
paedomorphosis. A less superficial approach wilt bear out the correctness of
this interpretation. %he human adult brain, were it paedomo.rphic, should
in its structure show resemblances to that of a young ape, but, of course,
does no such thing. The development .of the special co.rtical areas concerned

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with speech, for instance, and the enormous size of the areas representing
hand and eye and reduced importance of those concerned with smell are not
characteristics .of the young ape, lost in ~he adult, but are something peculiar
to Homo. The human brain is a further .development of the evolutionary
trends shown in other primates and not an example of return to a juvenile
condition as a result .of decreased functional importance in the adult.
A point which does reflect decreased functional importance and which
may legitimately be regarded as a case of paedomorphosis, is the smallness
of the jaws relative to any .other bodily dimension. This diminished importance of Vhe jaws is also reflected in the general lightness of the skull
and reduction of heavy boney strutting and ridges for muscle attachments.
Thus, as far as the head goes, it is these characteristics rather than the
large brain of man, that are paedomorphic.
We may next consider the question of human dental morphology. This
is cited by DE BEER & SWINTON (I958) aS an example of paedomorpho,sis,
the only supporting evidence being a statement that "the milk teeth .of the
Australopithecines resemble Vhose of modern man, whereas their permanent
dentition resembles that of apes". This statement is taken from a brief
report, in a semi-popular journal, of a paper .delivered by SPUULER (I954 a)
before the American Association of Physical Anthropologists. One cannot
but express surprise that two zoologist,s of such distinction as Sir GAVIN
DE BEE~ and Dr. SWI?CTON should accept as their sole authority a statement
of this sort, without any attempt to .discover the basis on v~hiCh it rests,
especially w[hen detailed descriptions of Au.stralopithecine dental anatomy
are readily available in the numerous papers published by BRoo51, DART
and ROmNSOhT 1).
SPUHLEX'S paper may have been published in full, but I have not succeeded in tracing it. An abstract (SPuLER, 1954 b) has, however, appeared.
It is clear from this that a series of measurements on teeth of various
primates, including two types of modern man, "Australian" and "European",
was used to arrive at an index known as the "'coefficient of .divergence".
Unfortunately, fhe meaning of this index does not appear from the abstract
and its value seems highly questionable since it results in a number of very
peculiar series .of which one sample, based on milk teeth, is as follows: - "Australopithecus afrfc,cmus, Australian, Orang-utah European, Gorilla,
Chimpanzee", while another, based on the permanent dentition, reads:

I) All the information about the teeth of the Australopithecinae has now been collected together and discussed in detail by Rom~rsoN (I956) to which paper the reader
is referred for details.

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~. F. EWER

"Paranth'ropus robustus, Orang-utan, Gorilta, PIesSanthropus transvacdensis,

Australian, Chimpanzee, European."
These series are difficult to understand but it seems possible that SPUHLER
may have fallen into the error of attaching too much importance to size, in
contrast to morphological pattern, in devising his method of comparison.
It is true that when we study closely related species in which the morphological pattern i,s well-nigh identical, the sizes of the individual teeth, bo{h
absolute and relative to each other, may provide reliable systematic characters. Size, however, is a characteristic in which parallel evolution has
occurred in Mmost every mammalian lineage: there is hardly a genus that
does not include larger and smaller species and the larger the taxonomic
unit we con,sider the greater the size range covered. It therefore follows that,
wt.hen comparing forms which are not extremely closely related, size becomes
useless as a guide to relationships. An example should make this clear enough.
The dentitions of the lion and domestic cat are very similar morphologically,
although the difference in the size of the teeth is considerable. The teeth
of the spotted hyaena (Crocuta crocuta (]~rxl.)) resemble those of the lion
in size, but differ considerably in morphological pattern. The systematist
who would suggest that these facts indicate that the lion is mere closely
related to the hyaena than to the cat would be laughed to scorn, but an
exactly comparable argument has in fact been seriously advanced in the
case of the ~igher primates (AsHTON & ZUCI~EI~ANN, 1950 ).
This ,di~scussion has led us away from our main point, the possible
paedomorphosis of the human dentition. In studying this problem we must
consider the relationships of the permanent and .deciduous dentitions. In
the case of the cheek teeth, the two sets are differentiated under the influence
of the same mMarisation field. However, unless that field extends forwards
pari passu vCith the elongation of the jaw as the animal grows, it is to be
expected that milk molars and permanent molars will resemble each other
since they are affected by the full strength of the field but that the anterior
premolars will differ in being less molarised. If there i.s an evolutionary
trend towal~ds more efficient grinding cheek teeth this will involve both
elaboration of the mMarisation field and its extension forwards during
ontogeny, so as to bring not only the molars, but the premolars too, under
its influence. The result will be the process of "molarisation of the preremarks", shown most clearly in the Per~ssodactyla (see BUTLER, I952 ).
Milk teeth are not juvenile stages of permanent teeth and the resemblance
between the tv~o thus finally achieved is not an example of paedomorphosis,
but rather its exact opposite.
If it were true that Australopithecine permanent teeth resemble those of

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pongids, but their milk teeth those of man, it would be a most remarkable
state of affairs, with the molarisation field altering its characteri,stics, not
merely quantitatively, but qualitatively during its ontogeny. It is not very
clear why this should be interpreted as showing that the human dentition
is paedomorphic: the simplest way in which such a peculiar condition coul,d
be explained w~uld be to suggest that the Austral.opithecine dentition was
evolving towards a pongid type from an originally more hominid condition,
and that change,s which affect the molarisation field only in its 'later stages
of ontogeny had started. Fortunately it is not necessary to ,draw such a
revolutionary conclusion, for the facts of Australopithecine dental morphology are not as stated by SPUHLEP~. The position may be summarised as
follows: the differentiation fields of the cheek teei~h of Australopithecines
and hominids are very ,similar but .differ considerably from the p ongid
pattern, particularly as affecting the specialised seetorial first 'lower premolar
of the latter, which acts as a sharpener for the large upper canine. In the
Australopithecinae the molarisation field keeps pace with the growth of the
jaw, so that the premolars, differentiated under its influence, are also highly
molarised. In man, however, the anterior premolars are les,s molarised, reflecting a molarisation field which does not extend forwards so far during
the time at which the premolars are being Jdifferentiate.d. That this should
be regarded as a secondary reduction, consequent upon ,diminished importance of the grinding function of the cheek teeth is suggested by the
following facts. In Homo sapiens, P1 is highly variable, from bicu.sped to
almost unicusped. High variability is normally associated with the relaxed
selection which accompanies decreasing functional importance. In Pithecanthropus, Pl i.s regularly more molarised than in Homo. The human
.dentition, like the jaw size, therefore show's characteristics of dental pattern
which reflect .decreased functional importance. There i,s no evidence that
these changes are such as to make ~he teeth resemb'le growth stages of the
permanent teeth of an ancestral form, and they are such as to make them
different from the milk teeth both of their own species and of the Austral-.
opi~hecinae. We thus have here an example of "paedomorphosis" without
detailed resemblance to an earlier ontogenetic stage. The condition of the
pelage in man i,s another character frequently regarded as pae.domorphic,
but the condition is by no means simple. There is not merely a r~duction
of the hair covering to a pre-functional juvenile stage, but a highly specialised
condition in which, although body hair is scanty and arrector pill muscles
poorly developed, the ~air on the head is retained and in some races is longer
than in any other mammal. Moreover, the development of hair in certain
Acta Biotheoretica, X I I I

~3

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1~. F. EWEI~

areas as a secondary sexual character can hardly be attributed to paedomorphosis.

One 'last point may be considered, to wit, t~he prolonged period .of Childhood
characteristic of man. It is difficult to .decide whether this should be regarded
as paedomorphosis, since man has not only a longer childhood but also a
longer total life-span than other primates. The Chimpanzee is sexually mature
at 7 years old, man at about 15: Chimpanzees in zo.os live to about 3o year,s
of age (FLowER, 1931), man in similar protected conditions to about 7~ .
It is unlikely, however, that .during the early stages of evolution man's potential longevity was very impo.rtant; even today it i.s only in the more advanced civilisations that the average expectation .of life is more ~han 5o years.
It would seem therefore that in the early stages selection for long Childhood
mu~st have been more important than selection for long life. In general, the
more quickly the young can grow to the adult stage the more chance they
have of reproducing before they get killed. Prolongation of youth can be
selected only if it provides some advantage which outweighs the increased
chance of pre-mature death. In man, this of course was provided by the
learning that takes place during childhood. Clearly therefore the prolongation of childhood could ,have begun only after a certain .degree of ability to
learn from parental "instruction" had been established. T,his ability to. learn
involves two components: not only the characteristics of the nervous .system
of the learner, but als.o the parental behaviour and social organisation of the
"instructors". We thus have a series: - (I) Development of social behaviour favouring learning by the young
which creates selective pressure for greater learning abi,lity.
(2) Development of learning ability, particularly w~en young. This creates
selective pressure for a quantitative increase in the amount learned.
(3) Development of a longer learning period.
This series will now become a circle with (3) making (I) more "worth
w:hile" and (2) making (I) more easily changed, so that evolution along these
lines will be expected to proceed very rapidly until the point is reached
where the increased chance o.f pre-mature .death, acting as negative feedback, brings it to a ~halt. Step (2) involves qualitative changes in the functioning of the ,central nervous system, for a diminution of the importance
of endogenous behaviour patterns and highly specific releasers is a necessary correlate of the growing emphasis on ability to learn. "Phe es.sential
rMe of parental protection in this process has been mentioned elsewhere
(Ewer, 1957). These changes in behaviour, like the increase in brain size
which presumably reflects them, are not just a retention of juvenile characteristics, but a further evolution along the lines ,shown to a lesser extent

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in other primates. Although an element of paedomorphosis may be involved

in the lengthening of the youthful phase, to designate the whole complex
process as paedomorphosi,s serves only to conceal the qualitative e'hanges
that have been taking place.
We thus seem to have reached a state of comp{ete obfuscation: certain
characters, suctl as relative head size, which mimic juvenile stages, are not
paedomorphic; others, such as the c~haracteristi.cs .of l~he teeth, reflect
reduction due to loss of importance of erstwhile adult specialisations, but
do not result in a juvenile appearance; still others, such as the condition of
the pelage and the behavioural characteristics, are extremely complex. Once
again, 'however, the situation beconles comprehensible w~en we put it into
its context in the live animal and ask the right question: "what was it that
permitted man to lose certain adult specialisations?" Although we may
normally make our ontogenetic entry into the world head first, we undoubtedly made our evolutionary debut the other way r o u n d - - f e e t first.
The "key character" in the evolution of man was bipedal locomotion, with its
anatomical correlates in changes in limbs, girdles, shape of thorax and head
orientation. This was what opened the way for the coordinated evolution of
hand, eye and brain which finally put man in the position to become the toolmaking animal. This iclea is not new: it was first put forward on purely
logical and fheoretical grounds by ENGELS about 8o years ago. Since then
the .discovery of lehe bipedal Austral.opithecinae has provided concrete supporting evidence and a number of autho.rs have expressed similar views,
(see, for example WASHBURN, I950, RoBrNsotr I956 ) but the relation to
human paedomorphosis has not always been appreciated. It was the use of
tools--"extra-corporeal organs", as they have been c a l l e d - - t o perform
functions v/hich previously had to be carried .out by bodily organs, xghich
permitted man to reduce or lose certain of the specialisations characteristic
of other adult primates. Some of these reduced characteristics signify
nothing more than toss of functional importance: but others may have
secondarily acquired new positive values. For instance, the wearing of
clot,hes associated with reduction in body hair not only provides a form of
t'hermoregulation rapidly adjustable to extremes of climatic variation, but
production and exchange of clothing forms .one component of ,social' organisatien and the role of clothing, as distinct from bodily characteristics
alone, in influencing the relations between the sexes or between social subgroups is by no means negligible. Simi'larly the large sensitive lips were very
likely elaborated first as a juvenile adaptation, but retained because they
became of importance in sexual behaviour, as suggested by BE BEER (I948).

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R. I~'. E W E R

It is, however, more likely that the lips were originally not simply, as
DE BEER puts it, "an adaptation to prolonged suckling" but that their
primary function was concerned with finding the maternal teat, which must
have begun t.o present difficulties in an animal .devoid of vibrissae and with
the .sen,se of smell decreasing in importance and the distinction between
naked teat and hairy surroundings becoming less marked.
In dealing with human evolution there has been a tendency to imply that
it is to his paedomorpho.sis that man owes his evolutionary success. This,
'however, is really (to use a mixed metap~ho.r) patting ourselves on the back
with the wrong end of the 'stick. The characters which have made man
".different from all other animals" are not those in which he is paedomorphic
but those in which he shows himself to be an advanced primate with an
aberrant type of locomotion: the paedomo.rphic characters although undoubtedly a n.ecessary element in the "ascent of man" are secondary consequences of the non-paedomorphic key characters. Man may have escaped
from primate arboreal specialisation and in so ,doing found the whole world
literally at hi.s feet, but this was done not merely by"becoming as little
children" which, while it may qualify for entry into the Kingdom of Heaven,
would have resulted in earthly extinction, had it not been preceded by the
acquisition of new characters not present in either the adult or the baby of
any other primate.
Although many other examples might be considered, what ~has been said
should suffice to make the main point clear: that evolution proceeding with
natural ,selection as its direction-conferring component is a process remarkable, like Cleopatra, for its infinite variety. In attempting to understand it,
we will fare best if we are able, as DARWIN was, to see it as a set of interactions between organisms, always capable of change, always affecting and
being affected both by each .other and by the physical environment; and
althoug~h definitions and .descriptive labels are useful and neces,sary, we
must beware of the danger of too glibly attaching a name and assuming
that this constitutes an explanation.

NATURAL SELECTION AND NEOTENY

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SUMMARY
Even today, a century after the publication of the 'Origin of Species', current
~oological literature often reveals an insufficient grasp of the implications of the now
generally accepted view that it is natural selection that confers direction on the evolutionary process.
This is, in part, due to a reaction against oversimplified teleology and against
Lamarckism. In rejecting Lamarck's thesis that the activities of an animal directly
affect its hereditary characters it is frequently assumed that this implies that such
activities are irrelevant to the study of evolution. This is a non-sequitur, for activities
may affect evolution not directly, through heredity, but indirectly, by influencing the
direction of the selective forces impinging on the organism. Reasons are given for
concluding that changes in habits and behaviour frequently precede structural change
and, in fact, determine the direction of the latter.
As an example of a concept which deserves a more evolutionary treatment than it
commonly receives, the subject of neoteny is considered. It appears that failure to think
in terms of the selective forces involved has frequently led to error. A n analysis in
functional terms is made of the theory that the chordates represent seconda.rily
glorified sea-squirts and it is concluded that this idea is unlikely to be correct. The
characteristics of man that are commonly considered to be neotenous are also discussed.
It is concluded that a facile and unanalytical application of the label "neoteny" to many
of the evolutionary changes involved has tended to obscure rather than to clarify their
significance: the neotenous features of man are not those that have been of primary
importance in human evolution, but are secondary to the acquisition of new characters
which are not to be found in either the adult or the young stages of any other primate.

RgSUMP~
Aujourd'hui, un si~cle apr~s l'apparition de l'Origine des Esp~ces, la comriction est
presque g~nfirale que la sglection naturelle dirige le proc~s de l'~volution. N~anmoins,
les publications zoologiques actuelles rfiv~lent souve~t une comprehension insuffisante
de ce qui en r~sulte.
Ceci doit en partie ~tre consid~r~ comme r~action contre une t~l~ologie simpliste et
contre Ie Lamarckisme. En rejetant la th~se de Lamarck, que les activit~s d'un animal
ont une influence directe sur ses caract~ristiques h~r~ditaires, on admet fr~quemment
que de pareilles activitgs n'ont rien 5. voir a~cec l'~tude le l'~volution. Cependant, une
telle conclusion n'est pas ~ustifi~e, car les activit~s peuvent influer sur l'~rolution non
pas directement, par la voie de l'h~r~ditg, mais indirectement er~ agissant sur la direction des forces s~lectives que l'organisme rencontre. Des raisons ont ~tg donnfies qui
m~nent ~ conclure que des changements dans les habitudes et dans le comportement
precedent maintes lois le changement structural et mfime d~terminent la direction de
celui-ci.
Comme exemple d'un concept qui m~rite un traitement plus ~volutionniste qu'il ne
re~oive en g~n~ral, la n6ot6nie a ~t6 examin6e. I1 parait que l'on a fait beaucoup
d'erreurs en ne tenant pas compte des forces s61ectives engag6es. Une analyse au pointde-rue de la fonction a 6t6 appliqu6e ~ la th6orie que les chord6es ne sont que des
ascidies ndot6niques transfigur6es par apr~s, et il s'ensuit que cette id6e n'est probablement pas correcte. Aussi les earact6ristiques de l'homme g6n6ralement regard6es comme
neot6niques ont 6t6 discut6es. I1 en r6sulte que, en d6signant 16gfirement et sans analyse
plusieurs changemenls 6volutionnaires comme n6ot6nie, on a plut6t obsurci qu'6clairci
leur signification. Ce ne sont pas les traits ndoteniques de l'homme qui ont jou6 le
r~61e principal dans l'dvolution humaine; au contraire, ils n'apparaissent qu'apr~s I'acquisition de qnelques nouvelles caract6ristiques qui ne se trouvent ni dans l'adulte ni dans
les phases plus jeunes des autres primates.

Ig2

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EWER

ZUSAMMENFASSUNG
Sogar heutzutage, ein Jahrhundert nach dem Erscheinen der ,,Origin of Species",
zeigt die jetzige zoologische Literatnr oft eine unzuI~ingliche Einsicht in die Folgerungen, zu denen die jetzt allgemein akzeptierte Annahme leitet, dass die natiirliche
Selektion die Riehtung des Evolutionsprozesses bedingt.
Dies ist teilweise zu sehen als eine Reaktion gegen eine einseitige Tele~logie und
gegen das Lamarckismus. Bei der Ablehnung der These LAMAI~CK':,class die Aktivitiiten
eines Tieres einen direkten Einfluss ausfiben auf seine erbliche K6~;stitution, nimmt
man oft an, dass dies bedeutet, dass derartige Aktivit~iten den:,!~.ch nic~~ zur Sache
tun bei dem Studinm der Evolution. Zu einer derartigen Konklusion ~" J s tman jedoch
nicht berechtigt, denn, obwohl die Aktivitiiten die Evolution nicht direkt~beeinflussen
kgnnen mittels der Erblichkeit, so k6nnen sie dies jedoch indirekt, weil sie die Richtung
der selektiven I~riifte, welche auf das Organismus einwirken, bestimmen. Es werden
Argumente gegeben, welche die Konklusion rechtfertigten, dass Anderungen in Gewohnheit und Verhalten oft StrukturS.nderungen vorhergehen und tats~ichlich die Richtung dieser letztgenannten bestimmen. Die Neotenie wirdt als ein Beispiel eines Begriff
betrachtet, welcher eine mehr evolutioniire Behandlung verdient als bisher geschehen ist.
Es ergibt sich dass man viele Irrtiimer hat gemacht, indem man den betreffenden selektiven Kr~iften keine Rechnung getragen hat. Eine Analyse der Theorie, welche die
Chordateaa als neotenische Ascidien betrachtet, wird yon einem funktionellen Gesichtspunkt aus durehgefiihrt. Daraus scheint der Schluss berechtigt, dass diese Theorie falsch
ist. Auch werden die menschlichen Kennzeiehen, welche gewghnlich als neotenische %etrachtet werden, besproehen. Die Konklusion lautet, dass, wenn man leichtfertig und
ohne geniigende Analyse den Zettel ,,Neotenie" fiir mancherlei evolutive Nnderungen
anwendet, man die richtige Bedeutung dieser Erscheinung vielmehr verdunkelt als aufkUirt: nicht die neotenische Kennzeichen des Menschen sind am wichtigsten in seiner
Evolution; in Gegenteil sie erscheinen nur sekund~ir, nachdem die neuen Kennzeicheaa,
welche sich weder in den Erwaehsenen noch in den jiingeren Stadien yon den anderen
Primaten linden tassen, erschienen sind.

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REFERENCES
ALEXANDER,A. J. (1960). A note on the evolution of stridulation in the family Scorpionidae. - - Proc. zool. Soc. Lond. C X X X I t I , p. 391-399.
ASHTON, E. H. & S. ZUCKERMAN (1950). Some quantitative dental characters of fossil
anthropoids. - - Phil. Trans., ]B, C C X X X I V , p. 485-52o.
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