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SELECTION
AND NEOTENY
by
R. F. EWER
(Department of Zoology, Rhodes University, Grahamstown, South Africa)
(Received 4. V. 1959)
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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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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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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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EWER
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
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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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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
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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.
i8i
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
R.F.
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.
I83
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