Child Development, November/December 2000, Volume 71, Number 6, Pages 16871708

Child Development and Evolutionary Psychology

David F. Bjorklund and Anthony D. Pellegrini

Evolutionary developmental psychology involves the expression of evolved, epigenetic programs, as described by the developmental systems approach, over the course of ontogeny. There have been different selection pressures on organisms at different times in ontogeny, and some characteristics of infants and children
were selected in evolution to serve an adaptive function at that time in their life history rather than to prepare
individuals for later adulthood. Examples of such adaptive functions of immaturity are provided from infancy,
play, and cognitive development. Most evolved psychological mechanisms are proposed to be domain specific
in nature and have been identified for various aspects of childrens cognitive and social development, most notably for the acquisition of language and for theory of mind. Differences in the quality and quantity of parental
investment affect childrens development and influence their subsequent reproductive and childcare strategies. Some sex differences observed in childhood, particularly as expressed during play, are seen as antecedents and preparations for adult sex differences. Because evolved mechanisms were adaptive to ancestral environments, they are not always adaptive for contemporary people, and this mismatch of evolved mechanisms
with modern environments is seen in childrens maladjustment to some aspects of formal schooling. We argue
that an evolutionary perspective can be valuable for developing a better understanding of human ontogeny in
contemporary society and that a developmental perspective is important for a better understanding of evolutionary psychology.

INTRODUCTION
The new field of evolutionary psychology has captured the attention of many in academic psychology
(e.g., Buss, 1995; Daly & Wilson, 1988; Tooby &
Cosmides, 1992). Evolutionary psychologists attempt
to describe contemporary human functioning in
terms of evolved psychological mechanisms. Not surprisingly, much evolutionary research and theorizing
has focused on behaviors relating to mating (e.g.,
Buss, 1995) and social functioning among adults
(e.g., Cosmides & Tooby, 1992). Less theorizing by
people who identify themselves as evolutionary psychologists has focused on development. This is in
part because it is mature members of a species who
reproduce, the sine qua non of Darwinian explication.
Yet, individuals must survive through infancy and
childhood before reproducing, and there is every reason to believe that natural selection has acted as much
upon the early portions of the lifespan to promote
survival as it has upon adulthood. Our purposes here
are to introduce the field of evolutionary developmental psychology and to apply evolutionary thinking to the study of human development, believing
that an understanding of the whys of development
will help us acquire a better understanding of the
hows and whats of development (Geary & Bjorklund, 2000). Evolutionary developmental psychology
involves the expression of evolved, epigenetic programs in interaction with an individuals physical
and social environment over the course of ontogeny.

Central to evolutionary developmental psychology is

the idea that there are (and were in the environment
of evolutionary adaptedness) different adaptive pressures on individuals at different times in ontogeny.
We further propose that an evolutionary account
provides insight not only into developmental function, aspects of ontogeny that presumably characterize children universally and predictably, but also into
individual differences. An evolutionary account suggests that there are alternative strategies to recurrent
problems that human children faced in our evolutionary past. Such a perspective suggests that individual
differences in developmental patterns are not necessarily the result of idiosyncratic experiences but rather
are predictable, adaptive responses to environmental
pressures.
In the sections below, we first outline some of the
assumptions of the field of evolutionary psychology
as they have been developed since the mid-1980s
(e.g., Buss, 1995; Cosmides & Tooby, 1987; Daly & Wilson, 1988; Tooby & Cosmides, 1992). Our emphasis and
our examples, however, will reflect evolutionary psychology as it relates to development. Next, we introduce concepts especially pertinent to evolutionary
developmental psychology, specifically the developmental systems approach, the differential influence of
natural selection at different points in ontogeny, and
the development of evolved psychological mecha 2000 by the Society for Research in Child Development, Inc.
All rights reserved. 0009-3920/2000/7106-0018

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nisms. We then examine selective areas of research in

developmental psychology that have benefited from a
specific evolutionary perspective, including the effects
of parental investment on childrens development
and developmental antecedents of adult sex differences. We conclude by looking at the impact that an
evolutionary perspective can have for establishing a
better understanding of childrens psychological functioning in contemporary culture.

Evolutionary psychology takes these basic tenets

of Darwins theory and the advancements made to it
over the past 140 years (usually termed neoDarwinism) and applies them specifically to human psychological functioning. Although, as in any fertile area of
intellectual inquiry, there are some healthy disagreements about specifics of evolutionary theory applied to
humans, there are certain aspects of this new paradigm
that, in one form or another, most practitioners of the
field adhere to.

EVOLUTIONARY PSYCHOLOGY
Darwins (1859/1958) theory of evolution, as presented in the Origin of Species, is probably the best and
most enduring general explanation we have of the
human condition and our adaptation to the world.
The basic principles behind Darwins theory are relatively simple. First, there are many more members of
a species born in each generation than will survive,
termed superfecundity. Second, all members (at least in
sexually reproducing species) have different combinations of traits; that is, there is variation in physical and
behavioral characteristics among individuals within a
species. Third, this variation is heritable. Fourth, characteristics that result in an individual surviving and
reproducing tend to be selected as a result of an interaction between individuals and their environment
and are thus passed down (via ones genes) to future
generations, whereas the traits of nonsurvivors are not.
That is, genetically based variations in physical or psychological features of an individual interact with the
environment, and, over many generations, these features tend to change in frequency, resulting, eventually,
in species-wide traits in the population as a whole.
Thus, through the process of natural selection, adaptive
changes in individuals, and eventually species, arise.
Darwin referred to the reproductive success of individuals as reflecting their reproductive fitness, which basically refers to the likelihood that an individual will
become a parent and a grandparent. Contemporary
evolutionary theorists, taking advantage of scientific
advances that have occurred since Darwins time (particularly in genetics), use the concept of inclusive fitness
(Hamilton, 1964). Inclusive fitness includes Darwins
concept of reproductive fitness (in this case, having
many offspring) but also considers the influence that
an individual may have in getting other copies of his or
her genes into subsequent generations. For example,
by having one child, 50% of a womans genes are
passed on to the next generation. But by helping to rear
her four nieces and nephews, each of whom shares, on
average, 25% of her genes, a woman can further increase the copies of her genes in the next generation,
thereby increasing her inclusive fitness.

Evolved Psychological Mechanisms

Evolutionary psychologists have proposed that
psychological mechanisms are the missing link in
the evolution of human behavior. This is a position
presented by Cosmides and Tooby (1987, p. 277),
who proposed that cognitive processes in interaction with environmental input, generate manifest
behavior. The causal link between evolution and behavior is made through psychological mechanisms.
According to Cosmides and Tooby, at least in humans, adaptive behavior is predicated on adaptive
thought. Natural selection operates on the cognitive
level information-processing programs evolved to
solve real-world problems. Moreover, mechanisms
evolved to solve specific adaptive problems faced by
our ancestors in the environment of evolutionary adaptedness. These are domain-specific mechanisms, what
Cosmides and Tooby (1987) referred to as Darwinian
algorithms. That is, rather than influencing general
intelligence, for instance, Darwinian algorithms affect very specific cognitive operations, such as face
recognition, language acquisition, or the processing
of certain types of social interactions. Pinker (1997,
p. 21) captured this perspective succinctly: The
mind is organized into modules or mental organs,
each with a specialized design that makes it an expert in one area of interaction with the world. The
modules basic logic is specified by our genetic program. Their operation was shaped by natural selection to solve problems of the hunting and gathering
life led by our ancestors in most of our evolutionary
history.
If we possess domain-specific mechanisms for
solving specific problems, the implication is that our
mind is not a general-purpose problem solver and
that some things will be very difficult or impossible to
learn. Stated differently, this perspective proposes
that there are constraints on learning (Gelman & Williams, 1998). Constraints imply restrictions, and restrictions are usually thought of negatively. The human mind is notable for its flexibility. We, more than
any other species, live by our wits and have been able

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to adapt to the most varied range of environments of

any large animal. But constraints, from this perspective, enable learning, rather than hamper it.
Children enter a world of sights, sounds, objects,
language, and other people. If all types of learning
were truly equiprobable, they would be overwhelmed by stimulation that bombards them from
every direction. Instead, infants and young children are constrained to process certain information
in core domains (such as the nature of objects,
language) in certain ways. They come into the
world with some idea of how the world is structured, and this leads to faster and more efficient
processing of information within specific domains.
According to Gelman and Williams (1998, p. 600):
From an evolutionary perspective, learning cannot be a process of arbitrary and completely flexible
knowledge acquisition. In core domains, learning
processes are the means to functionally defined
ends: acquiring and storing the particular sorts of
relevant information which are necessary for solving particular problems.
The evolved psychological mechanisms proposed
by evolutionary psychologists have some things in
common with the innate-releasing mechanisms, or
fixed-action patterns, identified by ethologists to explain the often complex behaviors of animals in response to specific environmental conditions. There
are also some differences, however. For example, Tinbergen (1951) described the aggressive behavior of
male stickleback fish in response to the presence of
the red belly of another male stickleback fish (the red
belly being an indication of a readiness to mate). Stereotypic aggressive behavior was displayed to any
red stimulus that closely resembled the underbelly of
another male fish, and this is adaptive, in that it limits
access of other males to a prospective mate. Such responses could be thought of as evolved psychological
mechanisms; but most (if not all) such mechanisms
possessed by humans are more flexible in nature, reflecting general propensities to respond in certain
ways depending on the environmental conditions. It
is not the case, for example, that human males act aggressively toward any male stranger who enters their
territory (red belly or not). This pattern may be found
in some cultures, however, and depending on the
social organization of the group and the developmental
history of the individual, how a person (male or female)
responds to a stranger will vary. Nonetheless, according to evolutionary psychological theory, what underlies such responding are evolved psychological
mechanisms, which find their expression as a result of
interaction with the environment over the course of
development.

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Functional Analysis
Evolutionary psychological explanations focus on
adaptationist thinking stressing the function of a behavior or trait. For example, pregnancy sickness is
quite common during the early months of pregnancy,
occurring in the majority of women around the world
(e.g., Tierson, Olson, & Hook, 1986). Symptoms include
nausea, vomiting, and food aversions. Given these
symptoms, pregnancy sickness is understandably considered an illness. Profet (1992), in an elegant review
of the literature, however, showed that pregnancy
sickness can be better understood as an adaptation to
protect the health of the developing fetus. For example: (1) modern women acquire aversions to food that
are highest in toxins and tend not to develop aversions to foods that are more apt to be toxin-free; (2)
pregnancy sickness, including food aversions, corresponds to the time when an unborn child is most susceptible to the effects of teratogens; (3) pregnancy
sickness appears to be universal; and (4) women who
experience pregnancy sickness have lower levels of
spontaneous abortions than women who do not become ill (Weigel & Weigel, 1989).
Profets functional analysis of pregnancy sickness
demonstrates the benefits that an evolutionary perspective can have. What has typically been viewed as
a dysfunctional state, for which medication is frequently prescribed, is actually a well-adapted mechanism that serves to foster the development of the
unborn child. Although the discomfort associated
with pregnancy sickness is real, its consequence is an
embryo/fetus protected from environmental toxins
that would impair its development. It is ironic to note
that thalidomide, the drug that led to serious deformations of childrens limbs when taken early in
pregnancy, was sometimes prescribed to alleviate
pregnancy sickness.
Not all current aspects of cognition, behavior, or
morphology are the result of adaptation. Evolution
produces at least three products (Buss, Haselton,
Shackelford, Bleske, & Wakefield, 1998): adaptations, by-products, and noise. Adaptations refer to reliably developing, inherited characteristics that came
about as a result of natural selection and helped to
solve some problems of reproduction or survival in
the environment of evolutionary adaptedness. The
umbilical cord would be an example of an adaptation.
By-products are characteristics that did not solve
some recurring problem and have not been shaped
by natural selection but are a consequence of being
associated with some adaptation. The belly button
would be an example of a by-product. Finally, noise
refers to random effects that may be attributed to

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mutations, changes in the environment, or aberrations of development, such as the shape of ones
belly button. As this tripartite classification indicates, a characteristic may have evolved in a species
but not have been designed by the forces of natural
selection. The evolutionary psychologists task is to
identify and describe psychological mechanisms
that may have served to solve survival or reproduction problems in our species evolutionary past and
to differentiate those mechanisms from characteristics that may be better classified as by-products or
noise.
Furthermore, some adaptations may have negative
effects (by-products) associated with them. For example, the enlarged skull of a human fetus is surely an
adaptation (housing a large brain, associated with
greater learning ability and behavioral flexibility);
however, because of the size of the babys head, birth is
difficult (because of limits on the width of a womans
hips that result from constraints of bipedality), and
many women and infants have died in childbirth. The
cost/benefit trade-off, however, was such that the benefits of an enlarged brain outweighed the detriments
of neonatal and maternal death.
It is worth commenting briefly here on the concept of cost benefit analysis and its significance to
evolutionary psychology. Cost benefit analyses assume that behaviors have both benefits, or functions, and costs, or risks. Behaviors will be naturally
selected if the benefits outweigh the costs: The benefits do not have to be absolutely high but only
greater than associated costs (Krebs & McCleery,
1984). Also, from a developmental perspective, the
benefits associated with costs/risks can be either immediate or delayed. For example, childrens play,
which will be discussed in greater detail below, can
have substantial costs, sometimes resulting in injury
or death (e.g., Cataldo et al., 1986; Peterson, Brezeal,
Oliver, & Bull, 1997). Physical play also requires energy, and the energy demands of play must be considered in light of other caloric requirements (e.g.,
calories required for basic metabolism, growth, and
more direct learning tutorials; see Pellegrini, Hovart,
& Huberty, 1998). What benefits do children reap
from taking such risks? Some appear to be immediate, such as the fostering of muscle and skeletal development, whereas others appear to be delayed,
such as developing social skills that will be important in adult life. Regardless of when the benefit is
realized, most adaptations have some risks associated with them and do not reflect perfect solutions
to recurrent problems but rather trade-offs that have
produced, on average, over evolutionary time, more
benefits than costs.

The Role of the Environment

Counter to some common misconceptions, evolved
psychological mechanisms exist in transactional relations with environmental factors. Believing that certain behaviors are under the influence of evolved psychological mechanisms does not imply that aspects of
the physical and social environment do not play a
critical role in the development or form of behavior.
In fact, quite the opposite is true; most evolved mechanisms are quite sensitive to variations in environments and are expressed differently depending on
ones surroundings (see Gottlieb, 1992, 1998). This
point is critical, because humans live in a wide variety
of environments and require flexible cognitive and
behavioral systems to survive. Moreover, because
evolved mechanisms will be expressed differently in
different environments, evolutionary psychology can
contribute to our understanding of individual differences. For the most part, evolutionary psychology has
emphasized what is universal about the human species. The recognition, however, that genes are differentially activated by different experiences in development (see discussion of developmental systems
approach below), coupled with the idea of natural selection, provides a model for predicting how different
environmental conditions will result in different behavioral phenotypes.

EVOLUTIONARY DEVELOPMENTAL
PSYCHOLOGY
The value of a behavior can be understood in terms of
ultimate function (i.e., fitness, or producing offspring, who, in turn, survive to reproduce) or in terms
of beneficial consequences of that behavior to the organism during its lifespan (Hinde, 1980). Tinbergen
(1963) stated the benefit of asking four questions to
understand the value of behavior: What is the immediate benefit (internal and external to the organism)?
What is the immediate consequence? How does it develop within the species (ontogeny)? How did it evolve
across species (phylogeny)? To answer these questions,
we must take a developmental perspective; we must
appreciate the adaptive value of a particular behavior
at a specific time in development. This implies that different behaviors or characteristics of an animal may be
selected at different times in ontogeny. In other words,
over the course of evolution, natural selection has functioned to adapt organisms to their current environments, and the environments and selective pressures
experienced by our ancestors early in their ontogeny
differed from the environments and selective pressures
experienced by our ancestors later in their lifespan.

Bjorklund and Pellegrini

The Role of the Environment across Ontogeny

in Evolutionary Psychological Perspective:
The Developmental Systems Approach
If evolved psychological mechanisms underlie contemporary behaviors and thought patterns, what role
can culture, or experience in general, play? A common
misconception that was held by many psychologists until recently about evolutionary explication (and may still
be held by some today) is that, if an ability is said to have
evolved or to have an innate component, the result is
one of biological, or genetic, determinism (see Charlesworth, 1992; Morss, 1990). If it is in the genes (which it
must be if it evolved), it cannot be changed. This is not
the case, and evolutionary psychologists are explicit
about the role that the environment plays (and did play)
in the expression of evolved psychological mechanisms.
Evolutionary psychologists assume that organisms
adapt and evolve, through natural selection, by their
transactions with the environment. Organisms affect
their environment (e.g., by choosing and then furnishing their niches), and environments, in turn, affect the organism (e.g., by changing behaviors to meet
the particular demands of a setting). Because of this
transactional relation between organism and environment, we must study organisms interacting with their
environments if we want to understand adaptation
and development. This position rejects any simplistic
biological determinism such as genetic endowment
having a main effect on cognitive functioning (see
Pellegrini & Horvat, 1995, for a discussion) or on social development (Pellegrini & Smith, 1998). More
specifically, we believe that the developmental systems
approach provides a proper appreciation of how biology and environment, at a variety of levels, interact to
produce behavior and development and that such a
model can be used to explain how evolved psychological mechanisms are translated into behavior.
The core concept of the developmental systems approach is that of epigenesis, which Gottlieb (1991a, p. 7)
defined as the emergence of new structures and
functions during the course of development. Gottlieb (1991a, 1998; Gottlieb, Wahlsten, & Lickliter, 1998)
stated that epigenesis reflects a bidirectional relation
between all levels of biological and experiential factors, such that genetic activity both influences and is
influenced by structural maturation, which is bidirectionally related to function and activity. This relation
can be expressed as follows:
genetic activity (DNA RNA proteins)
structural maturation function, activity.
From this perspective, functioning at one level influences functioning at adjacent levels. For example,

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genes code for the production of protein molecules,

which in turn determine the formation of structures,
such as muscle or nerve cells. But activity of these and
surrounding cells can serve to turn on or off a particular gene, thereby causing the commencement or cessation of genetic activity. Also, self-produced activity
or stimulation from external sources can alter the development of sets of cells. From this viewpoint, there
are no simple genetic or experiential causes of behavior; all development is the product of epigenesis, with
complex interactions occurring among multiple levels
(see also Johnson, 1998).
Evolved psychological mechanisms can be thought
of as genetically coded messages that, following
epigenetic rules, interact with the environment to
produce behavior. The experiences of each individual, however, are unique, beginning before birth, and
if the developmental systems account of ontogeny
closely mirrors reality, there should be substantial
plasticity in development. Yet, despite the fact that
genes will be expressed differently in different environments, almost all members of a species (human or
otherwise) develop in a species-typical pattern. How
can this be so and the developmental systems perspective still be valid?
The answer lies in the fact that humans (or chimpanzees or ducks) inherit not only a species-typical
genome but also a species-typical environment. According to Lickliter (1996, pp. 9091), . . . the organism-environment relationship is one that is structured
on both sides. That is, it is a relation between a structured organism and a structured environment. The organism inherits not only its genetic complement, but
also the structured organization of the environment
into which it is born. To the extent that an organism
grows up under conditions similar to that in which its
species evolved, development will follow a speciestypical pattern. Tooby and Cosmides (1992) have argued that complex, psychological mechanisms evolve
only under circumstances when the environments are
relatively stable over many generations. Thus, over
long periods of time, members of a species could expect certain types of environments, and they evolved
species-typical solutions to deal with such stable environments. For example, in the wild, a mother duck
will lay several eggs together in a nest and stay close
by the eggs until they hatch. While in the egg, the
ducklings begin to vocalize and so hear themselves,
the vocalizations of their brood mates, and those of
their mother. How might these experiences influence later species-typical behavior?
In a procedure developed by Gottlieb (1976, 1991b),
ducklings, while still in the egg, were isolated from
other eggs and their mother so that they could not hear

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the vocalizations of other animals. Their vocal chords

were also treated so that they could not produce any
sound (a condition that wears off several days after
hatching). Following hatching, these animals were
placed in a large container and heard the maternal call
of two speciestheir own and anotherplayed
through speakers on opposite sides of the container.
Most untreated birds in this situation approached the
call of their own species, seeming to know instinctively which call is that of their species and which is
not. However, ducklings who were prevented from
hearing any duck vocalizations, either their mothers,
those of other ducklings still in their eggs, or their own,
failed to make this discrimination and were just as
likely to approach the call of an alien species as that of
their own. Thus, prehatching experience plays a critical
role in posthatching species-typical behavior. The reason that nearly all ducks approach the species-typical
call after hatching is that nearly all ducks inherit not
only the genetic disposition to make such a selection
but also the species-typical environment that provides
the necessary experiences for such a pattern to develop.
A related example demonstrates how providing an
animal with species-atypical experience (rather than
depriving it of experience as in the Gottlieb studies)
can disrupt development. Lickliter (1990) removed
part of the eggshell 2 to 3 days before hatching of bobwhite quail and provided visual experience (patterned light) to these animals. Following hatching,
the quail chicks were tested in a situation similar to
that used by Gottlieb, with the maternal call of a quail
coming from one speaker and that of a chicken coming from another. A group of control animals that had
the egg shell removed but did not receive any additional visual experience displayed the species-typical
pattern: They approached the maternal call of their
own species on most occasions. In contrast, most of
the experimental animals showed no preference or
approached the maternal call of the chicken. The animals that received extra visual stimulation showed
enhanced visual discrimination abilities relative to
control animals, thus demonstrating a facilitory effect
of the early visual stimulation; but this came at a cost
to auditory discrimination abilities. Other research,
using ducks, quail, and rats as subjects, has demonstrated that providing young animals with stimulation that is outside the species norm has negative consequences for development (e.g., Gottlieb, Tomlinson,
& Radell, 1989; Kenny & Turkewitz, 1986; Lickliter &
Lewkowitz, 1995; Spear, 1984). What results such as
these demonstrate is that behaviors (here related to
infantmother attachment) that are found in almost
all normal members of a species are influenced by often subtle characteristics of the environment. Evolved

psychological mechanisms at the human level can be

similarly viewed. Strong species-wide biases may
exist for certain behaviors, but how any particular
evolved mechanism is expressed will vary with environmental conditions experienced at certain times in
development.
The substantial plasticity characteristic of early development provides a behavioral route for evolutionary change (Bateson, 1988; Gottlieb, 1992). For example, Gottlieb (1992) proposed that large-brained
animals with extended juvenile periods display substantial behavioral and cognitive malleability and
that this malleability can result in drastic changes in a
phenotype when a young animal is exposed to a
species-atypical environment. In this way, changes in
developmental rate or expressions of novel behavior,
brought about by changes in environmental conditions,
can serve as the fodder for natural selection, and lead,
eventually, to species-wide changes in a phenotype.
Some examples of how modified early environments can alter species-typical behavior that are particularly pertinent to human evolution come from observations of human-reared (enculturated) great apes.
Great apes (mostly common chimpanzees) who have
been raised by humans, much as human children, often
display more human-like cognitive abilities than
those displayed by mother-reared animals (see Call &
Tomasello, 1996). For example, the most successful of
the language-trained apes have been enculturated
(e.g., Gardner & Gardner, 1969; Savage-Rumbaugh et
al., 1993). Similarly, mother-reared apes rarely demonstrate imitation of tool use, particularly deferred imitation (i.e., imitating a behavior following a significant
delay). In contrast, enculturated common chimpanzees, bonobos, and orangutans have all been shown
to display above-chance levels of deferred imitation
of object manipulation (Bering, Bjorklund, & Ragan, in
press; Bjorklund, Bering, & Ragan, 2000; Tomasello,
Savage-Rumbaugh, & Kruger, 1993). Deferred imitation has traditionally been interpreted as requiring
symbolic representation (e.g., Meltzoff, 1995; Piaget,
1962), and aspects of these apes atypical, human-like
rearing history apparently prompted the emergence of
representational skills, at least in limited contexts,
which are absent from their mother-reared conspecifics.
It is not possible at this time to say what aspects of the
apes experiences are responsible for the change in their
cognitive abilities and behavior toward more humanlike thinking. One attractive candidate, however, has
been joint-attentional strategies, whereby adults draw
the attention of the young animal to an object (Call & Tomasello, 1996). An important aspect of this research is
that it provides an experiential vehicle by which our
hominid ancestors (using contemporary great apes as a

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model) could have begun to modify their cognition in

the direction that resulted in Homo sapiens.
The Influence of Natural Selection
at Different Times in Ontogeny
Ontogenetic adaptations and adaptive immaturity. In
keeping with the basic argument that there are different selection pressures on organisms at different
times in development is the idea that some aspects of
infancy and childhood are not preparations for later
adulthood but were selected in evolution to serve an
adaptive value for that specific time in development
(Bjorklund, 1997a; Oppenheim, 1981). As a result, certain immature aspects of a young animal often have
adaptive value. They were selected in evolution to
help keep the animal alive at that time in ontogeny.
This perspective has long been held by developmental psychobiologists, whose typical subjects are birds
or infrahuman mammals (e.g., Gottlieb et al., 1998;
Spear, 1984; Turkewitz & Kenny, 1982), but has been
less popular with developmental psychologists who
study human ontogeny and whose focus has often
been to find behaviors or traits early in life that are
predictive of later development.
Many adaptations are limited to a particular time in
development; they facilitate the young organisms
chances of surviving to adulthood and eventually reproducing. This is reflected by the concept of ontogenetic
adaptations neurobehavioral characteristics that serve
specific adaptive functions for the developing animal
(see Oppenheim, 1981). These are not simply incomplete versions of adult characteristics but have specific
roles in survival during infancy or youth and disappear when they are no longer necessary. For example,
embryos of most species have specializations that
serve to keep them alive but that disappear or are discarded once they serve their purpose, such as the yolk
sac, embryonic excretory mechanisms, and hatching
behaviors in embryonic birds (Oppenheim, 1981).
Ontogenetic adaptations in human infancy. Such adaptations are not limited to prenatal behaviors. Infant reflexes, such as the sucking reflex in mammals, are obvious postnatal behaviors that serve a specific
function and then disappear. Some aspects of human
infants cognition have also been interpreted as serving a specific function, only to disappear or to become
reorganized later in life. For example, the imitation of
facial gestures by newborns (e.g., Meltzoff & Moore,
1985) has been characterized by some as an ontogenetic adaptation (e.g., Bjorklund, 1987). Under the appropriate conditions, newborn infants will imitate a
range of facial gestures, although imitation of facial
expressions decreases to chance levels by about 2

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months of life (e.g., Abravanel & Sigafoos, 1984; Jacobson, 1979). Rather than serving to acquire new behaviors, which seems to be the primary function of imitation in later infancy and childhood, several researchers
have speculated that imitation has a very different
and specific function for the neonate. For example, Jacobson (1979) suggested that imitation of facial gestures is functional in nursing; Legerstee (1991) proposed that it serves as a form of prelinguistic
communication; and Bjorklund (1987) suggested that
it facilitates motherinfant social interaction at a time
when infants cannot intentionally direct their gaze and
control their head movements in response to social
stimulation. Heimann (1989) provided support for
these latter interpretations by reporting significant
correlations between degree of neonatal imitation
and subsequent quality of motherinfant interaction
at 3 months. Thus, early imitation appears to have a
specific adaptive function for the infant (i.e., to facilitate communication and social interaction) that is presumably different from the function that imitation
will serve in the older infant and child (but see Meltzoff & Moore, 1992, for a different interpretation).
Presumably, these different functions for similar behavior at different times in ontogeny were selected
over evolutionary time.
Play as an ontogenetic adaptation. There are similar
examples from social development, of which play is
perhaps the most obvious. Play is in many ways a
quintessential developmental construct. For instance,
it has been used to define, relationally, a developmental
period: The juvenile/childhood period is often defined as the period during which playful behavior is
dominant. Correspondingly, play is sometimes defined as that behavior which is exhibited by juveniles
(Martin & Caro, 1985). Thus, play has been considered to be an integral and important part of childhood
and one which accounts for a substantial portion of
childrens time and energy budgets (Hinde, 1974).
The ubiquity of play in juveniles lives has led many
scholars to assume that play serves a very important
developmental function. For example, some scholars
have listed over 30 possible functions of play (Baldwin & Baldwin, 1977).
More exact definitions of play have been proffered
by both ethologists (e.g., Martin & Caro, 1985) and
child developmentalists (Rubin, Fein, & Vandenberg,
1983), and they agree on a common consequential
definition of play: It is behavior that appears to have
no apparent function or where the means of a behavior are more important than the ends. In the ethological literature, this sort of purposeless behavior has
typically been divided into object play, social play,
and physical play (Fagen, 1981; Martin & Caro, 1985).

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Child Development

Specifically, play can occur with objects, as in the case

of Piagets (1962) sensorimotor play where very
young children and juveniles from a number of primate species (e.g., Kohler, 1925) perform a variety of
novel behavioral routines with objects. Play can be
solitary, as in cases where individuals play with materials, or social, where they play with an adult or a
peer. Physical play is vigorous and can be either solitary (e.g., swinging) or social (e.g., wrestling with a
peer or parent). For children, the paradigmatic case of
play is social-fantasy play, a uniquely human variety
of play (McCune-Nicholich & Fenson, 1984; Smith &
Vollstedt, 1985), although symbolic play has been
inferred in nonhuman primates reared by humans
(i.e., enculturated; see Tomasello & Call, 1997).
Some students of both animals and childrens play
have seen it as a source of creativity that may eventually lead to discovering new ways to solve old problems (Biben, 1998; Oppenheim, 1981); and, because of
the youthful tendency toward play and curiosity in
animals, it is likely that innovations will be introduced by the young rather than by adults. Support
for this contention comes from observations of the
skill of potato washing in Japanese macaque monkeys (Kawai, 1965). A group of Japanese scientists
provisioned a troop of wild monkeys with sweet potatoes, which were often sandy. One juvenile monkey
learned to wash potatoes in sea water before eating
them, and this was subsequently learned by other juveniles, and then some adult females. (Few adult
males ever learned this.) This innovation was then
passed on to infants as part of the culture. Although it
is unlikely that important cultural innovations will be
made through the play of human children, the discoveries children make through play may serve as the
basis of later innovations or true creativity, which become important later in life.
The functional question of play is particularly interesting in light of its most common definitional
attributeserving no apparent purpose. How can a
behavior be both developmentally important yet serve
no apparent purpose? Most theories, especially those
in the child development literature, assume that the
benefits of play are deferred until after the period
of childhood (Groos, 1898, 1901; Vygotsky, 1978). As
Kagan (1996) notes, this assumption may be due to
the bias toward the importance of early experience in
human development. In these theories, childrens
play is a way in which to learn skills important in
adulthood. Consequently, play is viewed as an imperfect version of adult behavior. In Batesons (1976)
terms, this is the scaffolding view of play: Play functions in the assembly of skills and is disassembled
when the skills are complete (e.g., Bruner, 1972). The

classic example of play serving deferred benefits is

where the play-fighting characteristic of juvenile
males is seen as practice for adult hunting and fighting skills (Smith, 1982).
An alternative view of play, labeled the metamorphic
view by Bateson (1976), holds that play is not an incomplete or imperfect version of adult behavior but is beneficial immediately and specialized to the niche of
childhood. In this way, play can be considered a specific adjustment to the context of childhood (Bateson,
1976; Bjorklund, 1997a; Gomendio, 1988; Pellegrini &
Bjorklund, 1997; Pellegrini & Smith, 1998). This view is
also consistent with the perspective that natural selection exerts functional pressure during the period of
childhood. An example of play serving an immediate
function holds that the sense of mastery and selfefficacy associated with play probably relates to children experimenting with new and different activities
and roles. Once activities are chosen, they should be sustained, which in turn affords opportunities for learning
specific skills (Bjorklund & Green, 1992). In a similar
vein, boys rough-and-tumble play may serve as a way
in which to learn and practice social signaling (Martin
& Caro, 1985), with exaggerated movements and a
play face communicating playful intent. Furthermore,
it is used as a way in which boys establish leadership in
their peer group and assess others strength (Pellegrini
& Smith, 1998). Rough-and-tumble play also has immediate nonsocial benefits; it provides opportunities
for the vigorous physical exercise that is important for
skeletal and muscle development (Bruner, 1972; Dolhinow & Bishop, 1970).
The adaptive nature of cognitive immaturity. Infants
and young childrens immature cognition may also
provide some adaptive value that is often overlooked
by developmental psychologists and educators (see
Bjorklund, 1997a; Bjorklund & Green, 1992; Bjorklund
& Schwartz, 1996). For example, young childrens poor
metacognition, particularly their poor ability to judge
the competency of their own performance, may be
adaptive in some contexts. Children who overestimate their own abilities may attempt a wider range
of activities and not perceive their less-than-perfect
performance as failure (e.g., Bjorklund, Gaultney, &
Green, 1993).
Other researchers have speculated that young childrens limited working-memory capacity may facilitate language acquisition. For example, Newport (1991)
and Elman (1994) have each proposed that children
initially perceive and store only component parts of
complex stimuli. They start with single morphemes
(usually a single syllable) and gradually increase the
complexity and the number of units they can control.
This results in a simplified corpus that actually makes

Bjorklund and Pellegrini

the job of analyzing language easier. With success and

time, maturationally paced abilities gradually increase,
as does language learning. Both Newport (1991) and
Elman (1994) performed computer simulations in
which they restricted the amount of information the
simulations could process at any one time (equivalent
to restricting how much children can hold in working
memory). They each reported that aspects of language were more easily acquired when the input was
initially limited (either by presenting a reduced corpus
or by limiting the working memory of the system).
These researchers concluded that young childrens
limited working-memory capacity restricts how much
language information can be processed, which simplifies what is analyzed, thereby making the task of
language acquisition easier. Preliminary support for
the less is more position also comes from evidence
that adults learn an artificial grammar faster when
presented with smaller units of the language (Kersten
& Earles, in press). (See Bjorklund & Schwartz, 1996,
for a discussion of these ideas applied to remediation
of language disabilities in children.)
Issues of accelerating cognitive development. Research
on these and other topics of cognitive development
(see Bjorklund, 1997a) indicates that certain aspects of
immaturity may be adaptive. This raises the question
about the wisdom of attempts to accelerate intellectual
development, frequently advocated in the United
States (see Bjorklund & Schwartz, 1996; Goodman,
1992), as well as the potential negative side effects of
early medical interventions. For example, Als (1995)
has suggested that preterm human infants have experiences similar in nature to those of Lickliters bobwhite
quail. In an extensive review of research examining factors that influence preterm infants brain development,
Als suggested that the unexpected stimulation that
preterm infants often receive in hospitals disrupts
brain development (particularly the frontal cortex)
during sensitive periods and frequently causes impairments resulting in lowered IQ, attention deficits,
eye-hand coordination difficulties, impulsivity, and
speech problems. These deficits, however, are often
accompanied by accelerated or enhanced abilities in
other areas, such as mathematics. Alss interpretation
is similar to that of Lickliters for bobwhite quail:
Stimulation outside the species-typical range can have
unforeseen consequences on brain and behavior development. Als (1995, p. 452) writes: Social contexts
evolved in the course of human phylogeny are surprisingly fine-tuned in specificity to provide goodenough environments for the human cortex to unfold,
initially intrauterinely, then extrauterinely. . . . With
the advances in medical technology, that is, material
culture, even very immature nervous systems exist

1695

and develop outside the womb. However, the social

contexts of traditional special care nurseries bring
with them less than adequate support for immature
nervous systems . . . leading to maladaptations and
disabilities, yet also to accelerations and extraordinary abilities.
In research with rhesus monkeys, Harlow (1959)
reported that animals who began discrimination
training at 155 days of age or younger actually performed more poorly on the learning tasks later in life
than animals who did not begin training until 190
days of age or older. This was true despite the fact that
the younger animals had more experience on the task
than the older animals. Harlow (p. 472) concluded
that there is a tendency to think of learning or training as intrinsically good and necessarily valuable to
the organism. It is entirely possible, however, that
training can either be helpful or harmful, depending
upon the nature of the training and the organisms
stage of development.
These and other findings (see Bjorklund, 1997a)
suggest that infants and young children respond to
experiences differently than older children and adults
and may be adapted for receiving particular amounts
and types of stimulation at different points in development. This interpretation, we argue, is consistent
with an evolutionary developmental psychological
perspective and is apt to be missed or interpreted
otherwise without such a perspective. This viewpoint
can be worthwhile for evaluating the benefits and the
costs of early education and intervention programs
for infants and young children (e.g., Hyson, HirshPasek, & Rescorla, 1990) and for children with special
needs (e.g., Goodman, 1992).
The Ontogeny of Evolved
Psychological Mechanisms
As we mentioned earlier, foremost in evolutionary
psychology is the idea that psychological mechanisms underlie important social and intellectual behaviors and that these mechanisms have evolved
(Buss, 1995; Tooby & Cosmides, 1992). These are
domain-specific, modular-like mechanisms that
evolved in the environment of evolutionary adaptedness, when our ancestors survived as hunters and
gatherers, and may not be associated with greater reproductive fitness today.
It would be extreme to claim, of course, that all
adaptive behaviors or thought processes have been
explicitly selected for their fitness value; some may
have been associated with another adaptive trait (byproduct) and not selected for themselves, and others
may simply have not been sufficiently maladaptive to

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Child Development

result in extinction. But a core assumption of evolutionary psychology is that psychological mechanisms
evolved to solve specific problems and are modular in
nature. They also did not evolve to deal with the
problems of contemporary humans; our species has
only recently abandoned a nomadic lifestyle for one
of villages, towns, and cities. Rather, these mechanisms evolved over the past several million years to
handle the problems faced by our hominid ancestors.
An important point here is that evolved mechanisms themselves develop. Evolved epigenetic programs are expressed by means of interaction with the
childs physical and social environment. Because of
the commonalities of human environments throughout the world and across time, many aspects of the
human mind and behavior will develop in a speciestypical way. Yet, these programs also reveal a substantial degree of flexibility, which permits individuals to adapt to the specific features of their environments. For example, children acquire language over
the course of 4 or 5 years. For adults, learning a second language is often very difficult, and the ease with
which children learn a first language seems at odds
with (i.e., independent of) their other more general
cognitive abilities. A number of specific evolved psychological mechanisms have been proposed to explain childrens acquisition of language (see Pinker,
1994), although other evolutionary-friendly proposals that posit a domain-general mechanism have also
been suggested (see Elman et al., 1996).
Similarly, aspects of childrens understanding of social functioning has been hypothesized to be modular in
nature (e.g., Baron-Cohen, 1995; Leslie, 1994). For example, by age 4, most children understand that other
people have beliefs and desires, sometimes different
from their own, that motivate their behavior. This
knowledge that peoples behavior is motivated by their
beliefs and desires (belief-desire reasoning; Wellman,
1990) has been referred to as a theory of mind, and it is difficult to imagine how any person could survive in human culture without such a theory. Being able to think
about others thoughts is crucial to detecting deception
and other social strategies that might handicap individuals. Although social intelligence, broadly defined, continues to develop into adulthood, most children by the
age of 4 have developed a belief-desire theory of mind.
Most children much younger than 4 years of age, however, seem to lack the requisite knowledge or conceptual
ability characteristic of belief-desire reasoning.
Theory of mind is illustrated by false-belief tasks.
In the standard false-belief task (e.g., Wimmer &
Perner, 1983), children watch as a treat is hidden in a
specific location (in a box, for example). Another person
(Maxi) is present when the treat is hidden but then

leaves the room, at which time the treat is moved to a

new location. Children are then asked where Maxi will
look for the treat when he returns. Most 4-year-old
children can solve the problem, stating that Maxi will
look where the treat was originally hidden, whereas
most younger children state that Maxi will look for the
treat in the new hiding place, apparently not realizing
that Maxis knowledge is different from their own.
Having a belief-desire theory of mind is required
for everyday exchanges of resources between two
people. For instance, in research by Peskin (1992),
3-year-old children play a game with mean monkey, who always wants the toy that the child wants
most. When children are asked to tell mean monkey which of several toys they really want and
which one they really dont want, mean monkey
(a hand puppet controlled by the experimenter) always takes the most desired toy, leaving the child
with the least desired one. Four-year-old children
catch on very quickly to the trick to deceive mean
monkey by pretending that the least-wanted toy is
really their favorite, thus foiling mean monkeys
evil plan. Most 3-year-olds, in contrast, never catch on
and spend the entire game being honest with mean
monkey and never getting the toys they most desire.
They fail either to monitor their own thinking or to
realize that mean monkey has a different goal in
mind than they do.
Some have argued that primate intelligence evolved
in response to detecting others cheating and cooperation (e.g., Humphrey, 1976), but a fully developed
theory of mind, based on belief-desire reasoning, is
found only in Homo sapiens. Although primatologists
have observed monkeys and apes engaging in tactical
deception, reflecting a suite of advanced cognitive
abilities (see Whiten & Byrne, 1988), such deception
does not necessarily require the ability to read the
mind of another individual (see Bjorklund & Kipp, in
press). For example, using nonverbal false-belief
tasks, Call and Tomasello (1999) found no evidence of
belief-desire reasoning, comparable to that of a human
4-year old, for chimpanzees and orangutans. In other
research, Povinelli and Eddy (1996) demonstrated
that chimpanzees do not understand seeing. In
their experiments, chimpanzees were just as likely to
request food from a naive observer or one who was
blindfolded as they were from an observer who knew
or could see the location of the desired treat. Thus, although deception is an important social skill, it does
not necessarily imply a highly developed theory of
mind. Humans obviously evolved a theory of mind
since our species last shared a common ancestor with
chimpanzees, and researchers have speculated how
other cognitive abilities, including language (e.g.,

Bjorklund and Pellegrini

Smith, 1998) and cognitive inhibition (Bjorklund &

Kipp, in press), might have co-evolved, or been prerequisite for, this uniquely human ability.
Consistent with an evolutionary developmental
psychological perspective, research has indicated that
this ability may be composed of a small set of modular-like skills. For example, Baron-Cohen (1995) has
proposed four separate, interacting modules involved in mindreading that develop over infancy and
early childhood. The earliest developing module is
the Intentionality Detector (ID), which interprets moving objects as having some volition or intention. For
example, an object that is moving toward an individual may be perceived as an agent with some intention
toward that individual (for instance, it wishes to
harm me, to be near me). This is a very primitive skill,
likely possessed by all animals with a nervous system. The second module is the Eye-Direction Detector
(EDD), which has three related functions: It detects
the presence of eyes or eye-like stimuli, determines
whether the eyes are looking toward it or toward
something else, and infers that if an organisms eyes
are looking at something then that organism sees that
thing. In other words, this module is responsible for
our belief that knowledge is gained through the eyes
(both ours and the eyes of others). According to
Baron-Cohen, these first two modules develop between birth and 9 months of age. The third module is
the Shared-Attention Mechanisms (SAM). Whereas the
ID and EDD involve only two objects/individuals
(that is, dyadic interactions/representations), the
SAM involves triadic interactions/representations.
For example, if person A is looking at object B, and
person C can see the eyes of person A and can see object B, person C can come to the conclusion that You
(person A) and I (person C) are looking at the same
thing. This module develops between 9 and 18
months. Finally, the Theory-Of-Mind Module (TOMM)
is roughly equivalent to the belief-desire reasoning
described earlier and is reflected by passing falsebelief tasks. This module develops between the ages
of about 18 to 48 months.
Possessing a theory of mind is central to any understanding of what it means to be human; and although monkeys and apes seem not to have a humanlike theory of mind, social primates do possess the
rudiments of an understanding of other conspecifics
as social beings, and these animals serve as models
for what the mind of our hominid ancestors may have
been like (see Byrne & Whiten, 1988; Russon, Bard, &
Parker, 1996). Moreover, researchers have used models
similar to those of Baron-Cohen (1995) described above
to explain primate behavior (e.g., Hauser & Carey,
1998; Tomasello & Call, 1997) in an attempt not only to

1697

understand the mind of primates but also to get a

glimpse at the evolution of the human mind.
Evidence for the modularity of the various components of Baron-Cohens model comes from studies of
children with autism. Baron-Cohen (1995) reviewed
research from his laboratory and those of other scientists suggesting that the more advanced forms of mindreading (SAM and TOMM) are typically absent in
children with autism. Autistic children (and later
adults) often seem to be in a world of their own and
have a difficult time in most forms of social interaction. Baron-Cohen claims that the primary deficit of
these children is an inability to read minds, or what
he calls mindblindness. Evidence for this conclusion
comes from studies in which autistic children are presented with false-belief and other theory-of-mind
tasks and consistently fail them, despite performing
well on other, nonsocial tasks (e.g., Baron-Cohen,
1989; Baron-Cohen, Leslie, & Frith, 1985; Perner, Frith,
Leslie, & Leekam, 1989). This is in contrast to children
with mental retardation, such as Down syndrome,
who perform theory-of-mind tasks easily, despite often
doing poorly on other tasks that assess more general
intelligence (e.g., Baron-Cohen et al., 1985). Most autistic children are able to perform well on the simpler
tasks requiring the ID or EDD modules, but fail tasks
involving the SAM and especially the TOMM modules. According to Baron-Cohen, autistic children are
unable to understand other peoples different beliefs,
even those children who are functioning at a relatively high intellectual level.
PARENTAL INVESTMENT: EVOLUTIONARY
EFFECTS ON CHILDREARING
When thinking of evolutionary (i.e., selective) influences on infancy and childhood, one naturally thinks
of direct genetic effects. What characteristics of childrens behavior have been selected to promote their
survival? There are, however, important factors external to children that may also influence their survival
and reproductive success and that have also been influenced by a long history of natural selection. More
specifically, we refer to the quantity and quality of
parenting, or parental investment, that children receive.
Within evolutionary psychology, the concept of parental investment (Trivers, 1972) has been used primarily to explain differences in behaviors related to
mating and parenting among men and women (see
Bjorklund & Shackelford, 1999). Because ancestral
men and women faced different adaptive problems
surrounding the amount of time, effort, and resources
required to rear an offspring to maturity, they evolved
different adaptive mechanisms. In mammals, fertili-

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Child Development

zation and gestation occur within the female, and

after birth, mothers provide the primary nutritional
support for their offspring until they are weaned. In
contrast, the males investment in the next generation
may be as little as the sperm he contributes. For a
slow-developing species such as Homo sapiens, however, paternal investment, in the form of providing
food and protection for the offspring and mother as
well as child care, increases the likelihood that a
mans offspring will survive and attain relatively
high status in the social group (see Geary, 1998). To
the extent that parenting influences childrens behavior and development (see Collins, Maccoby, Steinberg, Hetherington, & Bornstein, 2000), aspects of parental investment theory can and have been applied
to child development.
There is no doubt that the presence and investment
of parents, particularly in high-stressed environments,
is crucial to a childs survival and eventual social status. For example, Geary (1998) summarized research
from traditional societies, as well as historical data
from Western cultures, indicating that father absence is
associated with higher childhood mortality and, for
those children reaching adulthood, lower social status
than for children who have a father present. The death
rate is even greater when a mother is absent. What are
the factors that contribute to parents investing their
time and resources in a child? How might factors in the
home environment influence the reproductive strategies of the children as they grow up? Are there evolutionarily sound principles to predict under what conditions children will be neglected, abused, and even
killed by a parent (see Daly & Wilson, 1988)?
In light of the diverse social ecological niches that
individuals inhabit, that successful adaptation depends on the ability to choose from a variety of alternative strategies is not surprising. Most basically,
childrearing involves a balance between the caregivers and the offspring. Human newborns, of course,
are helpless, and caregivers must invest heavily in infants to maximize their survival. In terms of a cost
benefit ratio, this can be expressed as high costs (e.g.,
providing food, protection, and thermoregulative
support) and high benefits (nurturing the survival of
ones progeny) to caregivers and low cost and high
benefits to infants. Of course this ratio changes with
development, with benefits to children decreasing
with increasing age.
One aspect of parentchild relationships that has
important consequences for survival is attachment.
Bowlby (1969) integrated Freudian and ethological
concepts of attachment and proposed that mother
infant attachment is a human universal that evolved
to increase the likelihood of survival. Fernald (1992)

has even speculated that the infant-directed speech

that parents around the world use when speaking to
their babies evolved out of the attachment relationship. Infant-directed speech regulates infants emotions, behavior, and attention and also conveys a
mothers own emotional state to her infant, all factors
important to establishing an attachment relationship.
Although all infants become attached, the quality of
those attachment relations varies. Typically, securely
attached infants are viewed as being better adjusted,
both in infancy and in later childhood, than insecurely
attached infants. Although such a characterization
may have some validity from the perspective of mainstream Western society, differences in the quality of
attachment may reflect different adaptive solutions to
different physical and social environments, and insecure attachments, for example, should not automatically be viewed as less optimal than secure attachments.
For instance, children reared in homes characterized by
inadequate resources, high stress, marital discord/
father absence, and harsh and inconsistent childcare
reach puberty early, form short-term and unstable
pair bonds, and invest relatively little in their own offspring (e.g., Belsky, Steinberg, & Draper, 1991; Chisholm, 1999); male children tend to be noncompliant
and aggressive (Draper & Harpending, 1987). In stressful and uncertain environments, there is a tendency to
invest more in mating (for both sexes) than in parenting. In contrast, children from home environments
characterized by adequate resources and spousal
harmony/father present mature later, postpone sexual activity, and show greater investment in the fewer
number of children they produce (e.g., Belsky et al.
1991; Graber, Brooks-Gunn, & Warren, 1995; Kim,
Smith, & Palermiti, 1997; Moffitt, Caspi, Belsky, &
Silva, 1992; Surbey, 1998). Thus, depending on the
availability of resources, which is related to paternal
investment and spousal harmony, different patterns
of socialization occur that result in differential investment in the next generation.
Research examining the effects of environmental
conditions on reproductive maturity generally report
a greater effect for females than for males (e.g., Kim et
al., 1997). This sex difference makes sense, given the
differential investment in offspring by males and females. Because females investment in any conception is greater than males, they should be more sensitive to environmental factors that may affect the
rearing of offspring (such as malnutrition, stress, lack
of resources) than males (Surbey, 1998).
As children grow, they are less dependent on
mothers for their basic needs; thus conflicts between
mothers and childrens interests surface, usually at
the time of weaning (Hinde, 1987). Caregivers must

Bjorklund and Pellegrini

balance the costs associated with providing protection, food, and guidance to their offspring with the resources needed for their own survival and future
reproductive needs. Consistent with this argument,
caregivers would expend more resources on only/
last born children. In certain extreme cases (e.g.,
where caregivers resources are limited or where the
potential for offspring survival is low) infanticide can
result (Daly & Wilson, 1984). Fathers investment in
offspring often varies with the degree of paternal certainty (e.g., Daly & Wilson, 1988). These environmental
variations and their effects on parents, in turn, translate into differential treatment, and outcomes, for different children in the same family.
Other individual differences are associated with
childrens relationships with their parents, and these
differences may be related to a specific dimension of
the affectional system, warmth. Warmth has been
conceptualized as a reward system, distinct from the
attachment system, that may have evolved to promote
cohesive family relationships and parental investment
in their children (MacDonald, 1992). Individual differences in the warmth system may underlie parent
child relationships and subsequent personality. Specifically, the affectional system of which warmth is a
component may have evolved in such a way as to
shape our motivation to engage in certain behaviors
(e.g., opioid systems underlie the emotions of social
support and separation; Panksepp, cited in MacDonald, 1992). This reward system may provide the
basis for parents to invest (by providing warmth) in
the prolonged care of their offspring.
Individual differences in the amount and quality of
investment parents provide for their children, as well
as other important aspects of childrens social and
physical environment, can be addressed in terms of
evolutionary developmental psychology. Placing
such emphasis on environmental factors may, upon
initial inspection, seem at odds with a theory based
on the expression of evolved, genetically based, epigenetic programs over the course of ontogeny. But to
the contrary, evolutionary developmental psychology has much to say about the conditions in which
children are reared and the consequences of their
rearing environment on their later development.
The effects that parents have on the personality development and socialization of their children is complicated and not always direct (see Collins et al.,
2000). In fact, other theorists taking an evolutionary
perspective have suggested that forces outside of the
family exert a far greater role on childrens socialization than had previously been believed (e.g., Harris,
1995; Scarr, 1992). For example, Scarr (1992) proposed
that super parenting is not necessary to rear a suc-

1699

cessful child. Rather, children can tolerate a wide range

of parenting styles and still grow up to be successful
(i.e., reproductive) adults. Scarr proposed that patterns of child development are robust to variations in
parenting, with children seeking environments that are
compatible with their genotype, and it is these genotype-compatible environments that are chiefly responsible for shaping childrens behaviors and minds.
There is no single evolutionary account for the role
of parents and other cultural agents on the socialization of children. Taking an evolutionary developmental perspective can, however, provide insight for
understanding the different ways parents treat their
children; it can also help identify and illuminate alternative strategies that children and adolescents use to
deal with adaptive problems (e.g., mating). For example, viewing early physical maturation and sexual activity by some teens as an adaptive reproductive
strategy in response to stressful and uncertain environments may cause policymakers to see the problem
of teenage pregnancy in a different light and propose
different alternatives for its solution.
DEVELOPMENTAL ANTECEDENTS
OF ADULT SEX DIFFERENCES
Evolutionary psychologists have understandably
been interested in sex differences in adults. In particular, evolutionary social psychologists have focused
on sex differences in reproductive strategies, most as
they relate to parental investment theory (e.g., Buss,
1989; Shackelford & Larsen, 1997). Sex differences
have also been a favorite topic of developmental psychologists. Although some cognitive sex differences
are now typically attributed to an interaction of biological and social factors (e.g., Casey, 1996), differences in social behavior between boys and girls are
most typically attributed to the adoption of culturally
imposed gender roles (e.g., Eagly, 1987). Although
ones culture, a proximal mechanism, undeniably exerts a profound effect on ones gender-specific behaviors and roles, evolutionary psychology proposes that
males and females have evolved different strategies relating to mating and childrearing and that
these different evolved strategies, or distal mechanisms, underlie sex differences in associated behaviors across all human cultures (e.g., Bjorklund &
Shackelford, 1999; Geary, 1998). This, of course, does
not imply that adult sex differences arise fully formed
but rather that they emerge over the course of development and follow the precepts of the developmental
systems approach discussed earlier.
From a developmental perspective, many behaviors that reflect sex differences between adults should

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Child Development

have their origins in childhood. Thus, differences in

the social behaviors between boys and girls may reflect
preparations for important reproductively related behaviors observed in adulthood. Although we have
emphasized earlier that many adaptive characteristics of infancy and childhood are selected for this time
in development only and are not (necessarily) preparations for later life, other aspects of childhood do
serve to prepare the way for adulthood, of which sex
differences in social and cognitive abilities are good
examples.
Adult sex differences should be found in mating
strategies, in the degree males and females invest in
the well-being of their offspring, and in intrasexual
competition (see Geary, 1998, 1999). Following the
tenets of parental investment theory (Trivers, 1972),
women, because of the greater potential investment
they have in any potential copulation (i.e., pregnancy
and the principal job of nurturing the resulting child),
should adopt a more conservative mating strategy
than men. They should also be more oriented toward
childcare. With regard to intrasexual competition,
men in all cultures compete with one another for control of resources (e.g., money or cattle) and attainment
of social status. Such competition often involves
physical contests and often results in injury or death
(e.g., Cairns & Cairns, 1994; Daly & Wilson, 1988).
Of course, females also compete with one another
for mates. Our knowledge, however, of the ways in
which females accomplish this goal is extremely limited in both the human and animal literatures. Ethologists who have studied this (Gowaty, 1992; Smuts,
1985, 1995) have found that female primates use alliances with conspecifics for both defense against unwanted sexual overtures and access to desired males.
In gaining access to males, females often compete,
through alliances and deception, with other females
as well. We also know that human females use different forms of aggression than males, specifically relational aggression (Crick & Bigbee, 1998), which involves gossiping, backbiting, and shunning other
women, the goal of which appears to be to disrupt the
social networks of their competitors. In short, and consistent with Darwins original formulations, there is
both within- and between-sex competition for mates.
From this view, it is important to explore further females use of relational aggression or aggression
used in the service of social relations. Knowing the
goal of within- and between-sex relational aggression
would be helpful. That is, relational aggression is
used to manipulate social relations, but we do not
know what these relationships are the vehicle for. Is it
used by preschool girls against both boys and girls to
gain access to favored resources, similar to the way

preschool boys use physical aggression? Is it used in

adolescence against other girls in the service of gaining access to potential mates?
Competition and Aggression
Although females compete with one another and
use aggression, the intensity of that competition is not
as fierce as it traditionally has been for males and
rarely leads to serious physical injury or death. This
pattern of sex differences is particularly critical in a
species, such as humans, that is marginally polygamous, with some males being able to monopolize
more than one female and other males having access
to no females or only to less desirable females (i.e.,
those with low reproductive value). Most mammalian females will find a mate, even if not a highly desirable one; in contrast, the fitness variance is larger
for mammalian males, with many males being totally
excluded from mating. As a result, selection favored a
male psychology in which competitive risk taking
was favored (Daly & Wilson, 1988; Wilson & Daly,
1985). Such risk taking, and the violence that can accompany it, is universal and peaks when males are
entering the reproductive market, which in humans is
in adolescence (Cairns & Cairns, 1994; Daly & Wilson,
1988).
Risk taking and accidents are frequently the result
of competitive or show-off (display) behaviors,
with the purpose being to compete with other members of the same sex or to impress members of the opposite sex. Data from the United States indicate that
death from accidents and injuries resulting from violence are higher in males than females and rise rapidly for males in the late teens and continue to increase into the mid-20s before declining (see Cairns &
Cairns, 1994; Daly & Wilson, 1990). Similar patterns
are observed both for being the victim and the perpetrator of homicide (Cairns & Cairns, 1994; Daly & Wilson, 1988; Wilson & Daly, 1985).
Despite the societal penalties and presumed maladaptiveness of much of this behavior in contemporary culture, human males have inherited a psychology that was adapted to different conditions in which
risky competition during adolescence and young
adulthood, on the average, resulted in increased inclusive fitness. Such behaviors are not, of course,
programmed or inevitable but rather are shaped
by experience over development and are more likely
to be expressed in some environments than others.
For example, when a young male has limited access
to important cultural resources and when life expectancy is low, competing vigorously for mates and
what resources one can attain makes more sense than

Bjorklund and Pellegrini

taking a more cautious, long-term approach. Under

such conditions, which typify impoverished communities in affluent nations, males can be expected to engage in elevated levels of risk taking and violence
against other males. This is exactly the pattern one sees
in the United States for homicide rates of African American males (see Cairns & Cairns, 1994). The age pattern
is similar to that of Whites, but the absolute rate is
higher and is associated with reduced access to educational and economic opportunities for many inner-city
African American males in comparison with Whites.
Judging Ideal Mates
One interesting sex difference that is apparent in adolescence is concerned with sexual attractiveness. In all
cultures, women (including teenage girls) state that the
ideal mate is someone several years older than they are
(Buss, 1989; Kenrick & Keefe, 1992), and this corresponds to actual marriage practices (Kenrick & Keefe,
1992). Similarly, males in all cultures state that the ideal
mate is someone several years younger than themselves, and the age discrepancy in an ideal mate actually increases as men age (i.e., older men prefer increasingly younger women, whereas womens preferences
shown no such change; Kenrick & Keefe, 1992). Males
preference for younger mates has been interpreted as
reflecting an evolved psychological mechanism for
recognizing reproductive value. Reproductive value,
the number of children a woman can potentially have
in a lifetime, cannot be measured directly but can be inferred from other characteristics, and age is perhaps
the best single predictor. Thus, older mens ideal mate
is not someone just a few years younger than themselves but someone who has high reproductive value.
The only group not to show this pattern is adolescent
boys. Rather, teenage males ideal mate is a woman
several years older than themselves (Kenrick, Keefe,
Gabrielidis, & Cornelius, 1996). According to Kenrick
and his colleagues (1996), the reason for this anomaly
is that adolescent males are making their decision on
the basis of physical cues of reproductive value and
not age, per se. Women in their late teens and early 20s
are the most fertile and thus are selected by adolescent
males as most desirable, despite the fact that such
women express no interest in dating younger males
(Kenrick et al., 1996).
Play as Preparation for Adulthood
Precursors to adult sex differences are readily
found in childhood. Perhaps the most obvious differences are observed in the ways in which boys and
girls segregate themselves. This segregation has im-

1701

plications for the ways in which children interact and

play. For example, as early as 3 years of age, boys engage in more rigorous rough-and-tumble play, particularly in situations not monitored by adults (e.g., see
Pellegrini & Smith, 1998). In fact, there is some suggestion that girls actively avoid contact with boys because of their roughness (e.g., Haskett, 1971), thus
contributing to the universal formation of the samesex play groups that dominate the early school years
(e.g., Edwards & Whiting, 1988). Segregation in play
groups and males play being rougher than females
are also typical of nonhuman primate play (Biben,
1998). Males greater propensity toward rough-andtumble play has been associated with prenatal exposure to male hormones (Collaer & Hines, 1995).
The most common preparatory function proposed
for rough-and-tumble play in the animal literature relates to fighting and hunting in males (Biben, 1998;
Smith, 1982). This argument is based on the similarity
in design features; for example, both playing and
fighting involve hitting movements. In humans,
rough-and-tumble play mirrors the activities associated with malemale competition (i.e., primitive warfare) in hunter-gatherer societies (Keeley, 1996). Further, experiencing both superordinate and subordinate
role characteristics of rough-and-tumble play probably relates to social competence. Evidence from nonhuman primates, for example, suggests that juvenile
squirrel monkeys, Saimiri Sciureus, deprived of opportunities to engage in play fighting where they are
in superordinate (pinner) and subordinate (being
pinned) roles, are later bullies and sissies, respectively
(Biben, 1989). Thus, although boys rough-and-tumble
play may have some immediate benefits, such as establishing social hierarchies and facilitating skeletal and
muscle development, it also appears to teach boys
something about aggression, fighting, and social competition. Further, by engaging in the role alternation
characteristic of playfighting, boys are gaining experience in superordinate and subordinate roles (Biben,
1998), something that is useful in competitive interactions of all sorts but especially useful in their encounters with other males on matters of dominance
and eventual mating choices.
Girls play is also influenced by matters associated
with fitness and mating but in different ways. Girls
engage in more play parenting (i.e., doll play) than
boys, a sex difference that is found even in some primates (Pryce, 1995). Further, there is a relative absence
of dominance-related themes in the play of human and
nonhuman female primates (Biben, 1998). That is, females play is less often centered around roles where
physically based dominance relationships are publically exhibited. This sex differences has been viewed

1702

Child Development

as an evolved tendency that relates to the fact that females take primary responsibility for parenting their
offspring (e.g., Biben, 1998; Geary, 1998).
Sex Differences in Inhibition
Childhood sex differences are also found for some
forms of behavioral and social inhibition that may be
related to mating and childcare strategies. For example, Bjorklund and Kipp (1996) proposed that, because of the greater potential investment women
have in any sexual encounter, it would be in their best
interest to have greater inhibitory control of sexual
and social behaviors relative to men. Thus, compared
to men, women should be better at hiding their true
emotions (so as not to reveal prematurely an interest
in a potential mate). In a literature review, Bjorklund
and Kipp (1996) reported that females displayed
greater inhibitory abilities on tasks potentially related
to mating strategies, such as concealing emotions.
(This is true despite greater female emotional expression.) When sex differences were found consistently in a domain, they were found for children as
well as for adults. For example, in research in which
people are to display a positive emotion after a negative experience (for example, pretending that a
foul-tasting drink tastes good) or vice versa, females
from the age of 4 years are better able to control their
emotional expressions (that is, fool a judge watching
their reactions) than are males (e.g., Cole, 1986;
Saarni, 1984).
Bjorklund and Kipp (1996) also reported sex differences in inhibition abilities, favoring females, on tasks
requiring delay of gratification and resisting temptation, again at all ages tested (e.g., Kochanska, Murray,
Jacques, Koenig, & Vandegeest, 1996; Slaby & Park,
1971). These differences may relate to the greater inhibition skills that women need as principal caregivers
for their children. Effective parents must put the
needs of their infants first, delay their own gratification, resist distractions that would take them away
from their infants, and inhibit many aggressive reflex
responses to an often difficult and aversive infant.
The pattern of sex differences in inhibition abilities
found in both children and adults is consistent with
the pattern that one would predict if pressures associated with taking care of young children were greater
on hominid females than males (see Bjorklund &
Kipp, 1996). No consistent sex differences were found,
at any age, for tasks assessing cognitive inhibition,
which suggests that the sex differences that are observed are relatively domain-specific in nature and
relate to the different mating and childcare strategies
of ancient men and women.

An evolutionary account of sex differences, either

in adulthood or childhood, does not minimize the
contribution of culture in affecting the roles that men
and women adopt. Rather, the proposal here is that
adult sex differences, in all cultures, are built upon
evolved, epigenetic programs, based on the differential self-interest of ancestral men and women. Moreover, these abilities develop, with differences observed in adulthood being influenced by experiences
over the juvenile period.
Evolutionary theory provides a reason for assessing
sex differences on a wide range of social and cognitive tasks. Evaluating sex differences in our studies
for their own sake rarely seemed worthwhile. But
having an overarching theory that proposes that
males and females have evolved different strategies
for maximizing their inclusive fitness and that these
strategies develop in interaction with a childs social
and physical environment affords a motivation for
thinking that, sometimes, sex differences are something substantially more than error variance.
EVOLUTIONARY DEVELOPMENTAL
PSYCHOLOGY AND CONTEMPORARY CULTURE
Despite the many differences of lifestyle between contemporary humans and their Pleistocene ancestors,
there has been too little time for evolved psychological mechanisms to have changed since the advent of
civilization 10,000 to 12,000 years ago. As a result,
mechanisms evolved to adapt ancient humans to their
environments may not always be beneficial to modern people.
This insight is relevant to formal educational practices. Given that the modern human mind evolved to
solve problems faced by small groups of nomadic
hunters and gatherers, it is no wonder that many children balk at attending school. From the perspective of
evolutionary psychology, much of what we teach children in school is unnatural in that teaching involves
tasks never encountered by our ancestors (e.g., Jensen
et al., 1997; Pellegrini & Bjorklund, 1997). For example,
although our species has apparently been using language for tens of thousands of years, reading is a skill
that goes back only a few thousands of years, and it is
only in this century that a majority of people on the
planet are literate. Geary (1995) has referred to cognitive abilities that were selected in evolution, such as
language, as biologically primary abilities and to skills
that build upon these primary abilities but that are
principally cultural inventions, such are reading, as biologically secondary abilities. Biologically primary abilities are acquired universally and children typically
have high motivation to perform tasks involving them.

Bjorklund and Pellegrini

Biologically secondary abilities, on the other hand, are

culturally determined, and often tedious repetition
and external motivation are necessary for their mastery. It is little wonder that reading, a supposed language art, and higher mathematics give many children substantial difficulty.
On a related issue, Jensen and his colleagues (1997)
have proposed that a common childhood disorder
that impacts formal schooling, attention-deficit/
hyperactivity disorder (ADHD), may, in fact, reflect an
adaptational problem, for at least some afflicted children. Impulsivity, rapid scanning, and hyperactivity
may have been advantageous traits to early Homo sapiens. For example, high levels of motor activity may
have served (and still do) to gain information about
the immediate environment, which can be of benefit
to foraging, anticipating dangers, and spotting new
opportunities. Rapid scanning, as opposed to the
highly focused scanning useful in modern schools,
may have served to monitor threats and changing environmental conditions, particularly in stressful or
highly varied environments. And quick responses
without the benefit of reflection (impulsivity) may
have been favored when the likelihood of a delayed
response would have resulted in losing food or becoming the victim of a predator. Jensen et al. (1997)
proposed that the relatively high incidence of ADHD
today (3% to 5%) suggests that it has been maintained
by natural selection. Rather than all cases of ADHD being a disorder, many cases may reflect normal variation in a suite of cognitive/behavioral characteristics
or developed responses to early environmental conditions (e.g., high-threat or highly novel environments
yielding rapid scanning). Modern schools, with their
emphasis on highly focused instruction, provide a
poor fit to what were once adaptive evolved mechanisms. Similarly, Panksepp (1998) and Pellegrini and
Horvat (1995) have suggested that most children diagnosed with ADHD may simply be highly active and
playful youngsters who have a difficult time adjusting
to the demands of school. The widespread use of psychostimulant drugs to reduce the hyperactivity and increase the attentional focusing of children with ADHD
may reduce the desire and opportunity to play, which
may, in turn, reduce neural and behavioral plasticity.
An evolutionary perspective to developmental
psychology may also provide insights into some contemporary social issues, such as male-on-male violence during adolescence and young adulthood (e.g.,
Daly & Wilson, 1988), teenage pregnancy (e.g., Weisfeld & Billings, 1988), the effects of different parenting
styles on later behavior (e.g., Belsky et al., 1991), parent
child conflict (e.g., Surbey, 1998; Trivers, 1974), sibling
rivalry (Sulloway, 1996), changing patterns of social

1703

dominance over childhood (Hawley, 1999), and child

abuse (e.g., Daly & Wilson, 1996), among others. For
example, cases of child abuse and child homicide are
much more frequent in stepfamilies, with the abuse
often perpetrated by the stepfather (see Daly & Wilson, 1988, 1996). In fact, Daly and Wilson (1996, p. 79)
state that the step relationship itself is the single
most important risk factor for severe child maltreatment yet discovered. Because stepparents have no genetic investment in their stepchildren, from a strictly
inclusive-fitness perspective, stepparents should seek
to invest few of their resources on their stepchildren.
In many animal species, males will kill the offspring
of a new mate, thereby bringing the female into estrus
sooner and eliminating the investment of resources
he would have had to devote to his mates offspring
from a previous male. Humans, in fact, are exceptional in the nurturing that stepparents do provide for
stepchildren. Yet, differential patterns of child abuse
and expressed motivations for child homicides between genetic parents and stepparents suggest that
much of the violence against children is rooted in ancient evolved adaptations, and knowing this can result
in societal solutions to a pervasive problem.
CONCLUSION
An evolutionary perspective provides a common
ground for interpreting all aspects of human behavior
social, emotional, cognitiveand may serve to integrate the often disparate subfields of psychology. As
developmental psychologists, we have long believed
that the best way to understand any aspect of human
functioning is to look at its ontogeny. But developmental psychology has been as fractionated as its parent discipline. Many developmental scientists often
talk past one another because they fail to share a common view of what is important about development
(see Bjorklund, 1997b). Evolutionary psychology, we
believe, provides a metatheory for developmentalists
assessing a wide range of topics and ages (Baltes,
1997; Bjorklund, 1997b; Fishbein, 1976). As evolutionary theory is the foundation for modern biology, we
believe that it needs to be the foundation for modern
psychology (see Daly & Wilson, 1988; Tooby &
Cosmides, 1992; Wilson, 1998).
We also believe that an explicitly developmental
perspective can have a positive influence on the field
of evolutionary psychology. Much of evolutionary
psychology has been concerned with the natural selection of mature behaviors. Given this perspective,
it is easy to see why some evolutionary psychologists
have not looked at child development for interesting
phenomena. Evolution proceeds when successful in-

1704

Child Development

dividuals reproduce. These are the most progressed

members of the species, and factors that promote
their reproduction obviously characterize adulthood
and not infancy and childhood.
We must admit that, on the surface, this seems to
be a reasonable argument, and, quite obviously,
much of what does contribute to individual success
at reproduction, both today and in our evolutionary
past, is found in the adult. But our ancestors also developed, and before organisms can reproduce to get
their genes into the next generation, they must first
reach adulthood. For a slow-developing species such
as humans, that can be a long and treacherous path.
How people develop is important to eventual reproductive success, and, as we have noted previously,
we have every reason to believe that evolution has
worked to select characteristics of infancy and childhood that are adaptive to surviving to adulthood,
just as it has worked to make adults responsive to the
appropriate social and sexual cues that are so important in getting ones genes into the next generation.
Moreover, important characteristics of adulthood,
such as different sexual strategies of men and
women, should not be seen as preformed, springing
into existence with the first blast of pubertal hormones. Rather, even these characteristics have a developmental history, which can alter the expected
course of adult behavior. We believe that an evolutionary perspective is important for a new science of
developmental psychology.
ACKNOWLEDGMENTS
The authors thank six anonymous reviewers and Barbara R. Bjorklund for helpful comments on earlier
drafts of this manuscript. They also acknowledge the
W. T. Grant and Spencer Foundations for their support of this work.
ADDRESSES AND AFFILIATIONS
Corresponding author: David F. Bjorklund, Department of Psychology, Florida Atlantic University,
Boca Raton, FL 33431; e-mail: dbjorklund@fau.edu.
Anthony D. Pellegrini is at the University of Minnesota at Minneapolis.
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