Kroeber Anthropological Society Papers, Nos. 71-72, 1990
Is Sociobiology Methodologically Flawed?
P. Thomas Schoenemann
The argument between human sociobiologists and their critics is a case of competing paradigms. Two
frequent criticisms of sociobiology are: 1) it is inherently dangerous to assume that biology has an
important influence on behavior; and 2) the assumption of evolutionary adaptation is a methodological
error. These criticisms are aimed at the underlying assumptions of sociobiology with the intent of
demonstrating that sociobiology isfundamentallyflawed. It is argued that thefirst criticism disregards
examples of hwnan suffering that have occurred because of an ignorance of biology and insteadfocuses
exclusively on suffering that has occurred because of a misuse of biology, while the second criticism
represents a misunderstanding of the nature of evolutionary biological research. Individual sociobiolo-
gical argwnents may be incorrect, but the research paradigm is not intrinsicallyflawed.
INTRODUCTION
Human sociobiology means different things
to different people. The classic definition (Wil-
son 1975:2) states that sociobiology is "...the
systematic study of the biological basis of all
social behavior." Because it is recognized that
an organism's biology evolved (as opposed to
being specially created), sociobiology necessarily
involves an application of evolutionary biological
principles to the understanding of social be-
havior. There remains, however, an extreme
diversity of views on sociobiology's goals,
methods and accomplishments (or lack thereof,
depending on one's perspective). This can be
traced to the fact that sociobiology deals with
something that no one can claim to be agnostic
about. You may not, for example, have a vested
interest in the truths of ceramic tile heat conduc-
tion, or of cloud formation, but you are likely to
have some understanding of human behavior.
Because humans are highly interactive social ani-
mals, one simply cannot afford to be ignorant of
other people's behavior. To its severest critics,
human sociobiology is seen as a dangerous and
misguided attempt to speculate on the limits to
human behavior; so dangerous and misguided, in
fact, that they feel it should be rejected out of
hand as a research paradigm. How valid is this
claim?
In this article I will assess two of the most
common a priori reasons for rejecting the
sociobiological paradigm. The first criticism
emphasizes the potential negative effects of faulty
studies that attempt to detect/explain biological
influences on behavior. This viewpoint ignores
the other side of the coin, that there are serious
negative consequences to the assumption that
biology is irrelevant to behavior, even as a
working hypothesis. The second argument, ad-
vanced by some evolutionary biologists, is that
sociobiology suffers from serious methodologi-
cal flaws because it emphasizes a search for
adaptations. As we shall see, however, this cri-
ticism betrays a profound lack of insight into
how the field of evolutionary biology allows us
to understand the world around us.
THE PURSUIT OF KNOWLEDGE
My basic argument will hinge on the fact that
humans are severely limited in their ability to
perceive and understand the complexity of the
world around them. As scientists, we are con-
fronted with a bewildering array of empirical
observations which no human is able to compre-
hend as a single unit. In order to make sense of
this knowledge, we have no option but to break it
down into smaller, manageable pieces. We use
hypotheses, or theoretical constructs (which in
turn derive from our current scientific paradigm;
Kuhn 1970) to guide our explorations. Whether
the history of scientific understanding has pro-
ceeded in a tortuous and twisted route, as Kuhn
(1970) has argued, or simply by a series of suc-
cessive modifications of existing theories, it is
clear that the process requires intermediate steps.
Paradigms are constructed precisely because our
search for understanding proceeds at a snail's
pace through a series of intermediate steps. Para-
digms serve not only to organize and make sense
of the mass of empirical observations, but be-
cause they represent ways of perceiving the
world, they also serve to guide future research.
Paradigms define which questions "make sense"
to ask; that is, which questions are likely to pro-
vide meaning when answered. Even "holists"
(those who emphasize that the whole is more
than the sum of its parts) reject the "...romantic
notions that one can grasp the functional intri-
cacies of complex systems without conducting
50
scientific and technological studies of individual
components" (Soule 1985:728). We simply can-
not go from a position of complete ignorance to
one of complete knowledge in a single step.
Human sociobiology is an attempt to explain
behavior within an evolutionary biological
framework, using exactly the same theoretical
foundation that evolutionists have used to explain
morphological variation. In this sense, it repre-
sents nothing more than a simple extension of a
much larger research program. The diversity of
opinions about the merits of sociobiology stem
from an extremely basic dichotomy (a conflict of
paradigms) in how people choose to study be-
havior. There are those who are fundamentally
interested in differences between human groups,
and consequently feel that cultural variability in-
dicates human transcendence over biology; and
there are others who see meaning only in com-
mon underlying behavioral patterns, and are
consequently more receptive to biological expla-
nations.
These two basic points of view define quite
nicely (but obviously only on a very general
level) the fundamental difference between the
sociocultural anthropology paradigm and the
evolutionary biology paradigm. No graduate stu-
dent in sociocultural anthropology, for instance,
could get away with claiming that the people she
studied in her fieldwork in East Africa were for
all practical purposes essentially the same as the
Balinese. It is the nature of this field to empha-
size uniqueness, and because of this, there is no
unifying theme that ties the field together, save
the omnipotence of culture. In contrast, it is the
nature of evolutionary biology to emphasize com-
monalities, links and underlying threads. A
graduate student in zoology could not get away
with studying desert pupfish without putting
them into an evolutionary framework. Evolu-
tionary biology, like the rest of the natural
sciences, is a field that attempts to explain the
greatest number of differences with the least
number of principles. Human sociobiology is
nothing more than the application of evolutionary
biological analyses to human behavior (Ruse
1985; Barash 1977), and therefore shares all the
detriments as well as all the benefits of this larger
paradigm.
THE DANGER OF BIOLOGY
Perhaps the most frequent criticism of
sociobiology stems from the historical record of
abuse of biology to support racist, ethnocentrist
and sexist ideologies. The argument is that
because there is, and has historically been, such a
potential for the abuse of biological ideas in
political arenas, then any theories purporting to
explain human behavior should operate under the
assumption that biology is irrelevant. It is impor-
tant to keep in mind that there are two basic types
of errors that can be made when discussing beha-
vior. The first error is to assume that biology is
irrelevant when it is in fact relevant. The second
error is to assume that biology is relevant when it
is in fact irrelevant. Critics of sociobiology feel
that this second kind of error is inherently more
dangerous (and likely) than the first. They feel
that it is safer to ignore biology in discussions of
human behavior than to acknowledge it.
That this fear is central to criticisms of
sociobiology is evidenced by the opening chapter
of Kitcher's (1985) critique of sociobiology. In
it he describes a policy in England which placed
children into either academic or trade schools on
the basis of psychometric tests given at age ele-
ven. His point is that children were made to feel
like failures if they did not pass these tests, and
that this quite possibly scarred them for life. In a
sense, he believes that the tests created the differ-
ences between children, rather then pinpointing
existing differences. He feels that the risks of as-
suming a biological basis of behavior are greater
than the potential benefits that such a belief might
bring, and therefore we should always operate
under the assumption that biology is irrelevant.
Furthermore, he argues that we should hold re-
search on such subjects to a higher level of
scrutiny than other fields of inquiry. As Kitcher
(1985) wrote:
If a single scientist, or even the whole
community of scientists, comes to
adopt an incorrect view of the origins of
a distant galaxy, an inadequate model of
foraging behavior in ants, or a crazy
explanation of the extinction of the
dinosaurs, then the mistake will not
prove tragic. By contrast, if we are
wrong about the bases of human social
behavior, if we abandon the goal of a
fair distribution of the benefits and
burdens of society because we accept
faulty hypotheses about ourselves and
our evolutionary history, then the con-
sequences of a scientific mistake may
be grave indeed (1985:9).
There is nothing wrong with the argument
that we should weigh the costs and benefits of
implementing policies and doing research. The
question is whether the second kind of error
(being wrong in assuming that biological
differences influence a particular behavior) is
51
inherently more dangerous than the first kind of
ewror (being wrong in assuming biology is irrele-
vant to behavior). While Kitcher (1985) and
other critics point to examples of the negative
consequences when a biological influence on
behavior was mistakenly assumed, it is equally
possible to document cases where harm has oc-
curred because biology was implicitly, if not
explicitly, assumed to be irrelevant. One specific
example is the situation with regard to lactose
intolerance. Until recently, it was assumed (by
scientists and policy makers of European ances-
try) that all humans were essentially the same in
their ability to digest the milk carbohydrate lac-
tose as adults. However, it has become clear that
Europeans, along with some populations in Af-
rica, are abnormal with regard to most human
groups (and mammals in general) in their ability
to continue producing into adulthood the neces-
sary enzyme (lactase) which allows them to
digest lactose. "Lactose intolerant" adults suffer
a variety of ailments upon ingestion of milk,
including diarrhea (often severe) and other symp-
toms of a disturbed intestinal mucosa (Leininger
1978). These facts are now common in intro-
ductory textbooks in physical anthropology, but
what is not stressed is that milk has been com-
monly sent as famine relief to needy countries
where most of the population is lactose intoler-
ant, such as Asia. Milk-induced severe diarrhea
in malnounrshed individuals can be directly or
indirectly fatal. Since it is unlikely that mal-
nourished individuals will be as cautious about
the foods they ingest as healthy individuals, the
potential for serious complications is amplified.
This demonstrates that behavioral differences (the
lack of milk usage among most groups) may well
have biological bases (lactose intolerance) and the
ignorance of this possibility can and probably has
led to unnecessary suffering and death.
While much attention is given to the Nazi
atrocities carried out under the guise of "genetics"
(e.g., Stein 1988), little mention is made of the
devastation wrought in the Soviet Union because
of a rejection of genetics. From the 1930's until
the fall of Krushchev in 1964, Soviet genetic
theory and research was directed by the agron-
omist T. D. Lysenko, who rose to supreme
academic authority solely because his theories
were tailored to be desirable to the prevailing po-
ltcal environment (Medvedev 1969). Lysenko's
theories included a sort of bastardized version of
Lamarck's theory of the inheritance of acquired
characteristics along with an outight rejection of
Mendelian genetics. He insisted on the indivisi-
bility of the organism and maintained that it was
never possible to separate environmental from
hereditary influences, to the extent that the whole
field of genetics was decreed by him to be devoid
of meaning (Lemer and Libby 1976). He was
quoted as saying, "Just what is this gene? Who
has seen it? Who has felt it? Who has tasted it?"
(Medvedev 1969:257). For Lysenko, evolution-
ary change occurred in an organism when the
environment changed, because the organism was
nothing more than the sum of the environmental
conditions (Brill 1975).
The scientific basis for Lysenko's theories
was non-existent. "Scientific" arficles supporting
his claims consisted of various mixtures of
extreme polemics, guilt by association (e.g.,
Mendel was a priest and therefore his laws were
invalid), argumentwn ad hominem (arguments
that appealed to feelings and prejudices instead of
intellect), a rejection of statistical analysis, and
circular reasoning (certain results could be ob-
tained only under particular conditions, which
were defined as the conditions under which the
results were obtained) (Lerner and Libby 1976).
An incredible series of claims were made con-
cerning the miraculous transformation of one
species into another through purely environmen-
tal manipulation.
This fiasco had disastrous repercussions on
Soviet agriculture. The minimum probable loss
sustained in the Soviet Union due to its 20 year
failure to adopt American hybridization tech-
niques is estimated to have amounted to 30 to 50
billion kilograms of corn alone (Medvedev 1969:
180). Unfortunately, the consequences were not
limited to agriculture. By the end of 1936 all re-
search in human genetics was suspended because
it was considered inherently dangerous, racist,
degrading, and based on fallacious "bourgeois
genetics". Those scientists who disagreed with
Lysenko and his followers lost their jobs, were
reassigned to lower positions, or were sent to
work camps where an unknown number per-
ished. The first book in Russian on human
genetics was not published until 1964, which
meant that twenty-five successive classes of
physicians graduated from medical schools
"...without the slightest notion of the laws of
heredity" (Medvedev 1969:94). While resear-
chers in other countries became increasingly
aware that the severe behavioral effects of gene-
tically based metabolic disorders could usually be
avoided with special diets, Soviet doctors were
forced to toil under a proscribed ignorance.
This ignorance led directly to unnecessary
suffering. Research into phenylketonuria, for
example, which is known to cause early mental
retardation because of an enzymatic deficiency,
began in 1934 and a special diet had been devised
by 1953 (Plomin et al. 1990). Galactosemia,
another enzymadc deficiency that also causes
52
mental retardation, was discovered in 1956
(Cavalli-Sforza and Bodmer 1971). In all, the
specific enzyme deficiencies for some 24 differ-
ent genetic conditions (11 of which are known to
cause retardation or early death) were discovered
prior to 1964, the year of Lysenko's replacement
(Cavalli-Sforza and Bodmer 1971). If Soviet
scientists had been allowed to research human
genetics, it is likely that Soviet researchers would
also have made significant discoveries. While
we will never know the exact cost in human life
of the rejection of genetics in the Soviet Union
during this period, we can be sure it was not
trivial.
Is it possible, then, to justify the position
that it is always more humane to operate from a
non-biological paradigm in the absence of more
specific knowledge? Unfortunately, if we con-
tinue to operate from a non-biological mindset,
the only way to obtain specific knowledge of
biologically important influences on behavior will
be to stumble blindly onto them. The Lysenko
affair, along with the situation with regard to lac-
tose intolerance, serves to remind us that severe
costs have been paid as a direct result of igno-
rance. While neither situation involved a
conscious conspiracy to cause harm, this did not
make the costs any more palatable. It would
have been easy to avoid such costs if the right
questions had been asked. It will always be
more prudent to investigate every possibility,
including the possibility that biological differ-
ences influence behavior. This is not the same as
stating that we should assume all behavioral
variation is genetic. It is an appeal for more
knowledge. If differences exist, it is essential
that we understand them. Tmly humane policies
can only be derived from an understanding of
boti biological and environmental influences.
JUST-SO STORIES
In the post-Darwinian era, a reaction
against uncritical acceptance of the se-
lection theory set in, which reached its
climax in the great days of Comparative
Anatomy, but which still affects many
physiologically inclined biologists. It
was a reaction against maling uncritical
guesses about the survival value, the
function, of life processes and struc-
tures. This reaction, of course healthy
in itself, did not (as one might expect)
result in an attempt to improve methods
of studying survival value; rather it
deteriorated into lack of interest in the
problem -- one of the most deplorable
things that can happen to a science
(Tinbergen, quoted in Clutton-Brock &
Harvey 1979).
The second a priori objection to sociobio-
logy arose from within the ranks of evolutionary
biologists. Sociobiologists believe that behavior
itself should be considered an adaptation, just as
morphological characters, like the wings of
birds, have been considered adaptations (Barash
1977). As a working hypothesis, an animal's
behavior is not assumed to be the result simply of
learning, in a sense "independently created" in
each generation, but rather it is assumed to have a
heritable basis and therefore to have been shaped
by natural selection. This idea has been chal-
lenged primarily because of this emphasis on
behavior as an adaptation (Lewontin 1979).
Gould and Lewontin (1979) have been the pri-
mary critics of the way many biologists place
exclusive emphasis on the delineation of adapta-
tions in species. They argue that not enough
attention is given to such questions as "to what
extent are we justified in assuming a character (or
behavior) is an adaptation?", "How should we
define adaptation?" and "What other evolutionary
forces besides natural selection are operating to
produce the phenotypic and genotypic variation
we see today?" In defense, adaptationists ac-
knowledge these problems but argue that some
starting assumptions are necessary for any evolu-
tionary investigation (we do not start with perfect
knowledge), and that there is no reason to as-
sume that adaptationism as a worldng hypothesis
(i.e., as a guide to future research) is invalid. In
fact, as I will argue, the "adaptationist pro-
gramme" (Gould and Lewontin 1979) is the only
useful paradigm for evolutionary research. As a
result, sociobiology cannot be rejected out of
hand simply because it is "adaptatonist".
How do evolutionary biologists define adap-
tation? Lewontin (1978) has defined it as a
"process of evolutionary change by which the
organism provides a better and better 'solution'
to the 'problem.' The 'problem' is set by the
external world, and the agent of change is natural
selection" (1978:56). Function is therefore im-
plicit in the concept of adaptation. Futuyma
(1986) states that "the analysis of adaptations
entails showing that the trait has been developed
by natural selection, and specifying the nature of
the selective agent or agents that have favored the
trait" (1986:251). Clutton-Brock and Harvey
(1979) have argued that the concept of adaptation
should not be restricted to traits of known genetic
origin, since ontogeny always involves the
interaction of environment and genes. Any con-
ceivable trait would therefore have to be divided
53
into adaptive (genetic) and nonadaptive (envi-
ronmental) segments. By not restricting the
definition in this way Clutton-Brock and Harvey
(1979) acknowledge, however, that adaptations
could arise from forces other than natural selec-
tion. Most workers reserve the term adaptation
for those features specifically built for some
function by natural selection (Williams 1966;
Gould & Vrba 1982).
Gould and Lewontin (1979) argue that the
common strategy among evolutionists is to: 1)
atomize the organism into traits; 2) argue that the
traits are optimally designed for some function by
natural selection; and 3) argue that traits not
perfectly adapted to some function are so because
of the inability to optimize more than one trait
without imposing expenses on others, such that a
balance between opposing traits is reached. They
find fault in this strategy for two main reasons.
First, not enough care is given to the problem of
how to atomize traits, or how to find what
Lewontin (1979) calls "the 'natural' suture lines
for evolutionary dynamics" (1979:7). Both
Gould (1977) and Lewontin use the example of
the human chin: it is not an adaptation but simply
a result of two growth fields (alveolar and
mandibular) that, in humans, have regressed at
different rates (the former faster than the latter).
Second, they argue that the exclusive focus
on the adaptive function of traits, the "adapta-
tionist programme," obscures the fact that some
traits may not be adaptations. They feel that one
should not assume a trait is an adaptation even if
it is clearly advantageous for the organism: "The
mere existence of a good fit between organism
and environment is insufficient for inferring the
action of natural selection" (Gould & Lewontin
1979:592). Some evolutionists try one "adaptive
story" after another until they find one they can
live with. These explanations are dubbed "just-
so stories" after Kipling's children's tales. Other
evolutionists simply assume an adaptive expla-
nation exists without considering alternative,
nonadaptive explanations. Possible alternatives
include: 1) pleiotropy, where the trait is simply
one nonadaptive expression of a gene with many
effects (these other effects may or may not be
adaptive themselves); 2) multiple selective peaks
of equal adaptiveness, such that chance deter-
mines which peak an organism happens upon
first (e.g., one-horned Indian rhinoceros versus
two-horned African varieties); 3) allometry, in
which traits are unequally affected by develop-
mental processes, so that an increase in the size
of, for instance, cervine deer antlers is more than
proportionately greater than the corresponding
increase in body size; and 4) random gene fre-
quency changes due to finite population size, in
which combinations conferring lower genetic
fitness can even become fixed (if population and!
or fitness differences are small enough) (Gould
& Lewontin 1979).
Given these confounding variables, under
what circumstances are biologists ever justified in
advancing adaptationist arguments? Gould and
Lewontin (1979) give no explicit guidance, but
remark that they "would not object so strenuous-
ly to the adaptationist program if its invocation,
in any particular case, could lead in principle to
its rejection for want of evidence" (1979:587).
What they object to is the use of adaptationism as
a working hypothesis. They emphasize that bio-
logists need to treat the organism as a whole
entity, not a collection of discrete adaptations: "If
selection can break any correlation and optimize
parts separately, then an organism's integration
counts for little" (1979:596). Lewontin (1979)
states outright that "If sociobiological theory is to
make a lasting contribution to our understanding
of evolution, it must abandon the naive adapta-
tionist program..." (1979:14).
This is a fundamental methodological issue,
and to properly analyze it we must not forget
how science proceeds. Adaptationism is simply
one way of ordering the mass of empirical obser-
vations.  It does not follow  that we are
unjustified in using it to guide our investigations
simply because it might yield "incorrect" an-
swers. In fact, nonadaptative arguments suffer
from exactly the same theoretical problem.
Adaptationalism and nonadaptationalism do not
represent opposite ends of a spectrum (such as
"high metabolism" versus "low metabolism") but
rather the presence or absence of a phenomena.
The opposite of "adaptive" is not "less adaptive",
but "not adaptive". We just cannot know, a pri-
ori, which hypothesis is operational. But if we
start with the assumption of nonadaptation, the
quest for understanding is ended. Mayr (1983)
makes this explicit: "As a consequence of the
adaptationist dilemma, when one selectionist ex-
planation of a feature has been discredited, the
evolutionist must test other possible adaptationist
solutions before he can resign and say: This phe-
nomenon must be a product of chance" (1983:
326). We have no choice but to start from an as-
sumption, as a working hypothesis, that a given
trait is an adaptation if it confers some advantage
to the organism. There can never be direct
"proof' of nonadaptation, only a lack of adapta-
tionist explanations that fit our observations.
Nonadaptive arguments therefore suffer
from an intractable problem. Lewontin (1978)
points out that "biologists are forced to the
extreme adaptationist program because the alter-
natives, although they are undoubtedly operative
54
in many cases, are untestable in particular cases"
(1978:169). This is not true of most adaptive
arguments (Mayr 1983). As Clutton-Brock and
Harvey (1979) point out:
Though many functional arguments are
initially post-hoc, relatively few are ad-
hoc: almost all claim generality of some
kind and are consequently refutable by
more detailed observation, by experi-
ment or by examination of their validity
in other groups of animals... The fact
that they originated in post-hoc expla-
nations is not an objection: this is an
inevitable stage in any observational
science. In fact we rather doubt that
post-hoc explanations are more mis-
leading than many cases where
predictions are formulated and subse-
quently tested: it is seldom difficult to
produce facts that are generally con-
gruent with a theory, and the process
can appear falsely conclusive (1979:
550).
It is instructive to note that in Gould's (1984)
attempt to demonstrate nonadaptive variation in
the shells of land snails in the genus Cerion, he
empirically invalidates the adaptive arguments he
had previously advanced (Gould 1969). He
acknowledges that "some unexamined selective
agent might be clinally distributed throughout the
islands" (1984:235) that would account for the
observed variation. In order to argue for non-
adaptation, Gould himself implicitly recognizes
that he must demonstrate that the adaptive hy-
potheses are inoperable. Thus, whether one
operates from the adaptationist program or es-
pouses nonadaptationist explanations, one is still
obligated to test possible adaptive arguments.
THE CASE FOR ADAPTATIONISM
Aside from methodological issues, however,
there are other arguments that support the
adoption of adaptationism as an initial working
hypothesis. Mayr (1978) points out that our
world view is dominated by the knowledge that
both cosmic evolution and biological evolution
have proceeded under "more or less directional
natural processes consistent with the laws of
physics" (1978:47). Yet he goes on to state that
biological evolution is fundamentally more
complicated than cosmic evolution; that living
systems are "far more complex than any non-
living system" (1978:47). The tremendous
complexity of living organisms points to the im-
portance of natural selection in evolution. While
there have obviously been random, or nonselec-
tive, forces helping to shape organisms, such
forces could not have been dominant. No one
has been able to demonstrate how such complex-
ity could arise without selection. In fact, Wright
(1980) has argued that pure random drift (with
no accompanying selection) must lead to degen-
eration and extinction. While Wright's adaptive
landscape metaphor incorporates random factors,
he is in fact arguing that adaptations superior to
those produced solely by selection could be ob-
tained from a mixture of selection and drift.
Experiments have demonstrated that tremen-
dous changes can be induced in a population by
artificial selection, given some genetic variation
(e.g., Yoo 1980; and see Falconer 1981). Daw-
kins (1982) warns us that human subjective
judgement can be quite unreliable in assessing
just how unimportant weak selection may be to
have any effect. Mathematical calculations by
Haldane (1932) demonstrate selection pressure as
weak as 1 in 1000 on an advantageous allele
would lead to fixation in only a few thousand
generations, given reasonable assumptions about
population size. Lande (1976) has shown that,
assuming the heritability we see in extant species
has been constant throughout horse evolution,
only about two selective deaths per million indi-
viduals would account for the observed changes
in the paleontological record. Selection, there-
fore, can at least theoretically exert a tremendous
influence on biological evolution.
The utility of these calculations on extant
organisms to help us understand the past is
limited because we do not fully understand the
constraints imposed by developmental processes
on traits. Allometry, of course, is one such
developmental constraint. But allometrmc con-
stants are not constant for all time. Dawkins
(1982) points out that "the allometric constant is a
parameter of embryonic development. Like any
other such parameter it may be subject to genetic
variation and therefore it may change over evo-
lutionary time" (1982:33). Clutton-Brock and
Harvey (1979) have noted that allometric con-
stants vary between phylogenetic groups. It is
certainly not inconceivable that these differences
are due to selection. But even this can be tested:
if no other major ecological or behavioral factor
is correlated with increased antler size in relation
to body size, for example, then we may confi-
dently assert that the allometric constant, as far as
we are able to judge, is nonadaptive. Clutton-
Brock and Harvey (1979) found that large deer
species tend to be strongly polygynous. Since
males of polygynous species are likely to com-
pete for access to females, they may well invest
55
more heavily in large antlers than the males of
small, non-polygynous deer species. The acid
test would be to see how much of the variation in
antler size can be explained by these other vari-
ables.
There are evolutionary biologists that have
no faith in their ability to make decisions about
what constitutes an adaptation and what consti-
tutes a trivial character before actually studying
an organism. Cain (1964) remarks that, before
studies by Sheppard and himself demonstrating
selective forces operating on the snail species
Cepaea nemoralis and Cepaea hortensis, "it had
been confidently asserted that it could not matter
to a snail whether it had one band on its shell or
two" (1964:48). This remark comes frighten-
ingly close to Gould's (1984) remark about vari-
ation in Cerion: "Are such small differences,
each involving a simple variation in basic onto-
genetic pattern, necessarily adaptive? Must they
make a difference to a snail?" (1984:235).
Dawkdns (1982) notes that history seems to be on
the side of the adaptationists, in that time after
tdme aspects of organisms thought to be "trivial"
have been found to be adaptive (1982:31). This
does not mean that every trait must be an adap-
tation, but simply that "merely to fail on a casual
inspection to see any selective significance in a
particular variation does not license the observer
to proclaim that there is none" (Cain 1964:48).
Every question must be tackled afresh, on its
own merits (Dawkins 1982).
Cain (1964) also calls into question the
concept of neutral adaptation. What pattern of
variation should we expect to find for truly
neutral traits? He argues that we would find
enormous variation. The example he cites is of
fingerprints. Every human being has dermal
ridges. This lack of variation in morphology
suggests dermal ridges are functional (i.e., are
adaptive). However, provided there are enough
ridges to produce an adequate friction pad, and
assuming they run in all directions so that the
finger is not likely to slip in one direction more
tan another, "the exact pattern is imaterial and
can be allowed to vary. The resulting variation is
certainly tremendous, and this, and not relative
constancy, is what we would expect of neutral
characters" (1964:48). With this logic he main-
tains that ancestral traits that characterize major
clades (like the aquatic phase of amphibians) are
not merely neutral holdovers (as per Gould
1986), but represent adaptations to less special-
izedways of life. If these characters were truly
aeutral and not soundly functional, they would
varywildly, especially given their antiquity (Cain
1964:37). This must be true, since mutations are
continually occurring at all loci and would accu-
mulate unchecked in neutrl genes.
While it is true that adaptationists maintain
that natural selection is inherently an optimizing
process, this does not mean that they ignore con-
straints. Dawkins (1982), Clutton-Brock and
Harvey (1979), Cain (1964) and Mayr (1983) all
list essentially the same possible constraints on
perfection. The difference is that adaptationists
insist that we test for adaptations. This requires
that we construct plausible "just-so stories".
Adaptationists operate from the assumption that
natural selection, given enough time, closely
approaches the optimal solutions given the possi-
ble constraints. In sum, there is no reason to
assume: 1) that nonadaptationism as a working
hypothesis is most appropriate a priori; and 2)
that adaptation and nonadaptation are mutually
exclusive explanations of a trait: in order to test
for nonadaptation one is forced to exclude (test
for) adaptation.
CONCLUSION
Sociobiology is a research program aimed at
understanding behavior as an evolutionary adap-
tation. As such it carries all the underlying
assumptions of the field of evolutionary biology.
But because every paradigm has underlying as-
sumptions, this fact cannot be a criticism of
sociobiology, but instead is a criticism of human
limitations. The main reasons that have been
proposed for why sociobiology should not be
used as a research program simply do not hold
water. Nonbiological explanations of behavior
are not inherently less dangerous than biological
ones. Human suffering has resulted from the
uncritical acceptance of nonbiological expla-
nations. Critcisms of adaptationalism reflect a
misunderstanding of evolutionary biological rea-
soning. Adaptationist hypotheses are an essential
step in the process of gaining knowledge. Indi-
vidual sociobiological explanations of particular
behaviors may well be incorrect, but the socio-
biological world view is not "wrong" a priori.
ACKNOWLEDGMENTS
This paper owes much of its genesis to dis-
cussions with Vincent Sarich. David Wake and
Reina Wong provided useful comments and
criticisms on earlier drafts.
56
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