Evolutionary psychology


Evolutionary psychology is not simply a subdiscipline of psychology but its evolutionary theory can provide a foundational, metatheoretical framework that integrates the entire field of psychology in the same way evolutionary biology has for biology.[2][3][4]

Evolutionary psychologists hold that behaviors or traits that occur universally in all cultures are good candidates for evolutionary adaptations[5] including the abilities to infer others' emotions, discern kin from non-kin, identify and prefer healthier mates, and cooperate with others. There have been studies of human social behaviour related to infanticide, intelligence, marriage patterns, promiscuity, perception of beauty, bride price, and parental investment, with impressive findings.[6] The theories and findings of evolutionary psychology have applications in many fields, including economics, environment, health, law, management, psychiatry, politics, and literature.[7][8]





Evolutionary psychology is the long-forestalled scientific attempt to assemble out of the disjointed, fragmentary, and mutually contradictory human disciplines a single, logically integrated research framework for the psychological, social, and behavioral sciences – a framework that not only incorporates the evolutionary sciences on a full and equal basis, but that systematically works out all of the revisions in existing belief and research practice that such a synthesis requires.[11]

Just as human physiology and evolutionary physiology have worked to identify physical adaptations of the body that represent "human physiological nature," the purpose of evolutionary psychology is to identify evolved emotional and cognitive adaptations that represent "human psychological nature." According to Steven Pinker, it is "not a single theory but a large set of hypotheses" and a term that "has also come to refer to a particular way of applying evolutionary theory to the mind, with an emphasis on adaptation, gene-level selection, and modularity." Evolutionary psychology adopts an understanding of the mind that is based on the computational theory of mind. It describes mental processes as computational operations, so that, for example, a fear response is described as arising from a neurological computation that inputs the perceptional data, e.g. a visual image of a spider, and outputs the appropriate reaction, e.g. fear of possibly dangerous animals. Under this view, any Domain-general learning is impossible because of the Combinatorial explosion. This implies Domain-specific learning. Evolutionary Psychology specifies the domain as the problems of survival and reproduction.[12]



Nikolaas Tinbergen's four categories of questions can help to clarify the distinctions between several different, but complementary, types of explanations.[13] Evolutionary psychology focuses primarily on the "why?" questions, while traditional psychology focuses on the "how?" questions.[14]

Sequential vs. Static Perspective
Explanation of current form in terms of a historical sequence
Current Form
Explanation of the current form of species
How vs. Why Questions Proximate
How an individual organism's structures function
Mechanistic explanations for how an organism's structures work
Why a species evolved the structures (adaptations) it has


Evolutionary psychology is founded on several core premises.

  2. The brain's adaptive mechanisms were shaped by natural and sexual selection.[2][15]
  3. Different neural mechanisms are specialized for solving problems in humanity's evolutionary past.[2][15]
  4. The brain has evolved specialized neural mechanisms that were designed for solving problems that recurred over deep evolutionary time,[15] giving modern humans stone-age minds.[2][16]




Darwin, Charles (1859). The Origin of Species. p. 488 via Wikisource.


The content of evolutionary psychology has derived from, on the one hand, the biological sciences (especially evolutionary theory as it relates to ancient human environments, the study of paleoanthropology and animal behavior) and, on the other, the human sciences, especially psychology.

Evolutionary biology as an academic discipline emerged with the modern synthesis in the 1930s and 1940s.[20] In the 1930s the study of animal behavior (ethology) emerged with the work of the Dutch biologist Nikolaas Tinbergen and the Austrian biologists Konrad Lorenz and Karl von Frisch.

W.D. Hamilton's (1964) papers on inclusive fitness and Robert Trivers's (1972)[21] theories on reciprocity and parental investment helped to establish evolutionary thinking in psychology and the other social sciences. In 1975, Edward O. Wilson combined evolutionary theory with studies of animal and social behavior, building on the works of Lorenz and Tinbergen, in his book Sociobiology: The New Synthesis.

In the 1970s, two major branches developed from ethology. Firstly, the study of animal social behavior (including humans) generated sociobiology, defined by its pre-eminent proponent Edward O. Wilson in 1975 as "the systematic study of the biological basis of all social behavior"[22] and in 1978 as "the extension of population biology and evolutionary theory to social organization."[23] Secondly, there was behavioral ecology which placed less emphasis on social behavior; it focused on the ecological and evolutionary basis of animal and human behavior.

In the 1970s and 1980s university departments began to include the term evolutionary biology in their titles. The modern era of evolutionary psychology was ushered in, in particular, by Donald Symons' 1979 book The Evolution of Human Sexuality and Leda Cosmides and John Tooby's 1992 book The Adapted Mind.[5] David Buller observed that the term "evolutionary psychology" is sometimes seen as denoting research based on the specific methodological and theoretical commitments of certain researchers from the Santa Barbara school (University of California), thus some evolutionary psychologists prefer to term their work "human ecology", "human behavioural ecology" or "evolutionary anthropology" instead.[24]


Theoretical foundations


Evolutionary psychologists say that natural selection has provided humans with many psychological adaptations, in much the same way that it generated humans' anatomical and physiological adaptations.[26] As with adaptations in general, psychological adaptations are said to be specialized for the environment in which an organism evolved, the environment of evolutionary adaptedness.[26][27] Sexual selection provides organisms with adaptations related to mating.[26] For male mammals, which have a relatively high maximal potential reproduction rate, sexual selection leads to adaptations that help them compete for females.[26] For female mammals, with a relatively low maximal potential reproduction rate, sexual selection leads to choosiness, which helps females select higher quality mates.[26] Charles Darwin described both natural selection and sexual selection, and he relied on group selection to explain the evolution of altruistic (self-sacrificing) behavior. But group selection was considered a weak explanation, because in any group the less altruistic individuals will be more likely to survive, and the group will become less self-sacrificing as a whole.

In 1964, William D. Hamilton proposed inclusive fitness theory, emphasizing a gene-centered view of evolution. Hamilton noted that genes can increase the replication of copies of themselves into the next generation by influencing the organism's social traits in such a way that (statistically) results in helping the survival and reproduction of other copies of the same genes (most simply, identical copies in the organism's close relatives). According to Hamilton's rule, self-sacrificing behaviors (and the genes influencing them) can evolve if they typically help the organism's close relatives so much that it more than compensates for the individual animal's sacrifice. Inclusive fitness theory resolved the issue of how altruism can evolve. Other theories also help explain the evolution of altruistic behavior, including evolutionary game theory, tit-for-tat reciprocity, and generalized reciprocity. These theories help to explain the development of altruistic behavior, and account for hostility toward cheaters (individuals that take advantage of others' altruism).[28]


Evolved psychological mechanisms


Evolutionary psychologists seek to understand psychological mechanisms by understanding the survival and reproductive functions they might have served over the course of evolutionary history. These might include abilities to infer others' emotions, discern kin from non-kin, identify and prefer healthier mates, cooperate with others and follow leaders. Consistent with the theory of natural selection, evolutionary psychology sees humans as often in conflict with others, including mates and relatives. For instance, a mother may wish to wean her offspring from breastfeeding earlier than does her infant, which frees up the mother to invest in additional offspring.[28][29] Evolutionary psychology also recognizes the role of kin selection and reciprocity in evolving prosocial traits such as altruism.[28] Like chimpanzees and bonobos, humans have subtle and flexible social instincts, allowing them to form extended families, lifelong friendships, and political alliances.[28] In studies testing theoretical predictions, evolutionary psychologists have made modest findings on topics such as infanticide, intelligence, marriage patterns, promiscuity, perception of beauty, bride price and parental investment.[6]

Historical topics

Proponents of evolutionary psychology in the 1990s made some explorations in historical events, but the response from historical experts was highly negative and there has been little effort to continue that line of research. Historian Lynn Hunt says that the historians complained that the researchers:

have read the wrong studies, misinterpreted the results of experiments, or worse yet, turned to neuroscience looking for a universalizing, anti-representational and anti-intentional ontology to bolster their claims.[30]

Hunt states that, "the few attempts to build up a subfield of psychohistory collapsed under the weight of its presuppositions." She concludes that as of 2014 the "'iron curtain' between historians and psychology...remains standing."[31]


Not all traits of organisms are evolutionary adaptations. As noted in the table below, traits may also be exaptations, byproducts of adaptations (sometimes called "spandrels"), or random variation between individuals.[32]


AdaptationExaptationByproductRandom variation
DefinitionOrganismic trait designed to solve an ancestral problem(s). Shows complexity, special "design", functionalityAdaptation that has been "re-purposed" to solve a different adaptive problem.Byproduct of an adaptive mechanism with no current or ancestral functionRandom variations in an adaptation or byproduct
Physiological exampleBones / Umbilical cordSmall bones of the inner earWhite color of bones / Belly buttonBumps on the skull, convex or concave belly button shape
Psychological exampleToddlers' ability to learn to talk with minimal instructionVoluntary attentionAbility to learn to read and writeVariations in verbal intelligence

One of the tasks of evolutionary psychology is to identify which psychological traits are likely to be adaptations, byproducts or random variation. George C. Williams suggested that an "adaptation is a special and onerous concept that should only be used where it is really necessary."[34] As noted by Williams and others, adaptations can be identified by their improbable complexity, species universality, and adaptive functionality.

Obligate and facultative adaptations


Cultural universals

Evolutionary psychologists hold that behaviors or traits that occur universally in all cultures are good candidates for evolutionary adaptations.[5] Cultural universals include behaviors related to language, cognition, social roles, gender roles, and technology.[36] Evolved psychological adaptations (such as the ability to learn a language) interact with cultural inputs to produce specific behaviors (e.g., the specific language learned).

Basic gender differences, such as greater eagerness for sex among men and greater coyness among women,[37] are explained as sexually dimorphic psychological adaptations that reflect the different reproductive strategies of males and females.[28][38] Research has also shown participants were able to recognize the facial expression of fear significantly better on a male face than on a female face. Females also recognized fear generally better than males.[39]


Environment of evolutionary adaptedness


The idea of an environment of evolutionary adaptedness was first explored as a part of attachment theory by John Bowlby.[42] This is the environment to which a particular evolved mechanism is adapted. More specifically, the environment of evolutionary adaptedness is defined as the set of historically recurring selection pressures that formed a given adaptation, as well as those aspects of the environment that were necessary for the proper development and functioning of the adaptation.


The environment of evolutionary adaptedness is significantly different from modern society.[44] The ancestors of modern humans lived in smaller groups, had more cohesive cultures, and had more stable and rich contexts for identity and meaning.[44] Researchers look to existing hunter-gatherer societies for clues as to how hunter-gatherers lived in the environment of evolutionary adaptedness.[28] Unfortunately, the few surviving hunter-gatherer societies are different from each other, and they have been pushed out of the best land and into harsh environments, so it is not clear how closely they reflect ancestral culture.[28] However, all around the world small-band hunter-gatherers offer a similar developmental system for the young ("hunter-gatherer childhood model," Konner, 2005; evolved developmental niche" or "evolved nest;" Narvaez et al., 2013). The characteristics of the niche are largely the same as for social mammals, who evolved over 30 million years ago: soothing perinatal experience, several years of on-request breastfeeding, nearly constant affection or physical proximity, responsiveness to need (mitigating offspring distress), self-directed play, and for humans, multiple responsive caregivers. Initial studies show the importance of these components in early life for positive child outcomes.[45][46]

Evolutionary psychologists sometimes look to chimpanzees, bonobos, and other great apes for insight into human ancestral behavior.[28] Christopher Ryan and Cacilda Jethá argue that evolutionary psychologists have overemphasized the similarity of humans and chimps, which are more violent, while underestimating the similarity of humans and bonobos, which are more peaceful.[47]


Since an organism's adaptations were suited to its ancestral environment, a new and different environment can create a mismatch. Because humans are mostly adapted to Pleistocene environments, psychological mechanisms sometimes exhibit "mismatches" to the modern environment. One example is the fact that although about 10,000 people are killed with guns in the US annually,[48] whereas spiders and snakes kill only a handful, people nonetheless learn to fear spiders and snakes about as easily as they do a pointed gun, and more easily than an unpointed gun, rabbits or flowers.[49] A potential explanation is that spiders and snakes were a threat to human ancestors throughout the Pleistocene, whereas guns (and rabbits and flowers) were not. There is thus a mismatch between humans' evolved fear-learning psychology and the modern environment.[50][51]

This mismatch also shows up in the phenomena of the supernormal stimulus, a stimulus that elicits a response more strongly than the stimulus for which the response evolved. The term was coined by Niko Tinbergen to refer to non-human animal behavior, but psychologist Deirdre Barrett said that supernormal stimulation governs the behavior of humans as powerfully as that of other animals. She explained junk food as an exaggerated stimulus to cravings for salt, sugar, and fats,[52] and she says that television is an exaggeration of social cues of laughter, smiling faces and attention-grabbing action.[53] Magazine centerfolds and double cheeseburgers pull instincts intended for an environment of evolutionary adaptedness where breast development was a sign of health, youth and fertility in a prospective mate, and fat was a rare and vital nutrient.[54] The psychologist Mark van Vugt recently argued that modern organizational leadership is a mismatch.[55] His argument is that humans are not adapted to work in large, anonymous bureaucratic structures with formal hierarchies. The human mind still responds to personalized, charismatic leadership primarily in the context of informal, egalitarian settings. Hence the dissatisfaction and alienation that many employees experience. Salaries, bonuses and other privileges exploit instincts for relative status, which attract particularly males to senior executive positions.[56]

Research methods


Evolutionary psychologists use several strategies to develop and test hypotheses about whether a psychological trait is likely to be an evolved adaptation. Buss (2011)[57] notes that these methods include:

Current evolutionary Adaptiveness. In addition to evolutionary models that suggest evolution occurs across large spans of time, recent research has demonstrated that some evolutionary shifts can be fast and dramatic. Consequently, some evolutionary psychologists have focused on the impact of psychological traits in the current environment. Such research can be used to inform estimates of the prevalence of traits over time. Such work has been informative in studying evolutionary psychopathology.[58]

Evolutionary psychologists also use various sources of data for testing, including experiments, archaeological records, data from hunter-gatherer societies, observational studies, neuroscience data, self-reports and surveys, public records, and human products.[59] Recently, additional methods and tools have been introduced based on fictional scenarios,[60] mathematical models,[61] and multi-agent computer simulations.[62]

Main areas of research


Survival and individual level psychological adaptations

Problems of survival are clear targets for the evolution of physical and psychological adaptations. Major problems the ancestors of present-day humans faced included food selection and acquisition; territory selection and physical shelter; and avoiding predators and other environmental threats.[63]


Consciousness meets George Williams' criteria of species universality, complexity,[64] and functionality, and it is a trait that apparently increases fitness.[65]

In his paper "Evolution of consciousness," John Eccles argues that special anatomical and physical adaptations of the mammalian cerebral cortex gave rise to consciousness.[66] In contrast, others have argued that the recursive circuitry underwriting consciousness is much more primitive, having evolved initially in pre-mammalian species because it improves the capacity for interaction with both social and natural environments by providing an energy-saving "neutral" gear in an otherwise energy-expensive motor output machine.[67] Once in place, this recursive circuitry may well have provided a basis for the subsequent development of many of the functions that consciousness facilitates in higher organisms, as outlined by Bernard J. Baars.[68] Richard Dawkins suggested that humans evolved consciousness in order to make themselves the subjects of thought.[69] Daniel Povinelli suggests that large, tree-climbing apes evolved consciousness to take into account one's own mass when moving safely among tree branches.[69] Consistent with this hypothesis, Gordon Gallup found that chimps and orangutans, but not little monkeys or terrestrial gorillas, demonstrated self-awareness in mirror tests.[69]


Sleep may have evolved to conserve energy when activity would be less fruitful or more dangerous, such as at night, and especially during the winter season.[69]

Sensation and perception

Many experts, such as Jerry Fodor, write that the purpose of perception is knowledge, but evolutionary psychologists hold that its primary purpose is to guide action.[70] For example, they say, depth perception seems to have evolved not to help us know the distances to other objects but rather to help us move around in space.[70] Evolutionary psychologists say that animals from fiddler crabs to humans use eyesight for collision avoidance, suggesting that vision is basically for directing action, not providing knowledge.[70]

Building and maintaining sense organs is metabolically expensive, so these organs evolve only when they improve an organism's fitness.[70] More than half the brain is devoted to processing sensory information, and the brain itself consumes roughly one-fourth of one's metabolic resources, so the senses must provide exceptional benefits to fitness.[70] Perception accurately mirrors the world; animals get useful, accurate information through their senses.[70]

Scientists who study perception and sensation have long understood the human senses as adaptations to their surrounding worlds.[70] Depth perception consists of processing over half a dozen visual cues, each of which is based on a regularity of the physical world.[70] Vision evolved to respond to the narrow range of electromagnetic energy that is plentiful and that does not pass through objects.[70] Sound waves go around corners and interact with obstacles, creating a complex pattern that includes useful information about the sources of and distances to objects.[70] Larger animals naturally make lower-pitched sounds as a consequence of their size.[70] The range over which an animal hears, on the other hand, is determined by adaptation. Homing pigeons, for example, can hear very low-pitched sound (infrasound) that carries great distances, even though most smaller animals detect higher-pitched sounds.[70] Taste and smell respond to chemicals in the environment that are thought to have been significant for fitness in the environment of evolutionary adaptedness.[70] For example, salt and sugar were apparently both valuable to the human or pre-human inhabitants of the environment of evolutionary adaptedness, so present day humans have an intrinsic hunger for salty and sweet tastes.[70] The sense of touch is actually many senses, including pressure, heat, cold, tickle, and pain.[70] Pain, while unpleasant, is adaptive.[70] An important adaptation for senses is range shifting, by which the organism becomes temporarily more or less sensitive to sensation.[70] For example, one's eyes automatically adjust to dim or bright ambient light.[70] Sensory abilities of different organisms often coevolve, as is the case with the hearing of echolocating bats and that of the moths that have evolved to respond to the sounds that the bats make.[70]

Evolutionary psychologists contend that perception demonstrates the principle of modularity, with specialized mechanisms handling particular perception tasks.[70] For example, people with damage to a particular part of the brain suffer from the specific defect of not being able to recognize faces (prosopagnosia).[70] Evolutionary psychology suggests that this indicates a so-called face-reading module.[70]

Learning and facultative adaptations

In evolutionary psychology, learning is said to be accomplished through evolved capacities, specifically facultative adaptations.[71] Facultative adaptations express themselves differently depending on input from the environment.[71] Sometimes the input comes during development and helps shape that development.[71] For example, migrating birds learn to orient themselves by the stars during a critical period in their maturation.[71] Evolutionary psychologists believe that humans also learn language along an evolved program, also with critical periods.[71] The input can also come during daily tasks, helping the organism cope with changing environmental conditions.[71] For example, animals evolved Pavlovian conditioning in order to solve problems about causal relationships.[71] Animals accomplish learning tasks most easily when those tasks resemble problems that they faced in their evolutionary past, such as a rat learning where to find food or water.[71] Learning capacities sometimes demonstrate differences between the sexes.[71] In many animal species, for example, males can solve spatial problem faster and more accurately than females, due to the effects of male hormones during development.[71] The same might be true of humans.[71]

Emotion and motivation

Motivations direct and energize behavior, while emotions provide the affective component to motivation, positive or negative.[72] In the early 1970s, Paul Ekman and colleagues began a line of research which suggests that many emotions are universal.[72] He found evidence that humans share at least five basic emotions: fear, sadness, happiness, anger, and disgust.[72] Social emotions evidently evolved to motivate social behaviors that were adaptive in the environment of evolutionary adaptedness.[72] For example, spite seems to work against the individual but it can establish an individual's reputation as someone to be feared.[72] Shame and pride can motivate behaviors that help one maintain one's standing in a community, and self-esteem is one's estimate of one's status.[28][72] Motivation has a neurobiological basis in the reward system of the brain. Recently, it has been suggested that reward systems may evolve in such a way that there may be an inherent or unavoidable trade-off in the motivational system for activities of short versus long duration.[73]




Evolutionary psychology is primarily interested in finding commonalities between people, or basic human psychological nature. From an evolutionary perspective, the fact that people have fundamental differences in personality traits initially presents something of a puzzle.[75] (Note: The field of behavioral genetics is concerned with statistically partitioning differences between people into genetic and environmental sources of variance. However, understanding the concept of heritability can be tricky – heritability refers only to the differences between people, never the degree to which the traits of an individual are due to environmental or genetic factors, since traits are always a complex interweaving of both.)

Personality traits are conceptualized by evolutionary psychologists as due to normal variation around an optimum, due to frequency-dependent selection (behavioral polymorphisms), or as facultative adaptations. Like variability in height, some personality traits may simply reflect inter-individual variability around a general optimum.[75] Or, personality traits may represent different genetically predisposed "behavioral morphs" – alternate behavioral strategies that depend on the frequency of competing behavioral strategies in the population. For example, if most of the population is generally trusting and gullible, the behavioral morph of being a "cheater" (or, in the extreme case, a sociopath) may be advantageous.[76] Finally, like many other psychological adaptations, personality traits may be facultative – sensitive to typical variations in the social environment, especially during early development. For example, later born children are more likely than first borns to be rebellious, less conscientious and more open to new experiences, which may be advantageous to them given their particular niche in family structure.[77] It is important to note that shared environmental influences do play a role in personality and are not always of less importance than genetic factors. However, shared environmental influences often decrease to near zero after adolescence but do not completely disappear.[78]




Evolutionary psychologists hold that the FOXP2 gene may well be associated with the evolution of human language.[81] In the 1980s, psycholinguist Myrna Gopnik identified a dominant gene that causes language impairment in the KE family of Britain.[81] This gene turned out to be a mutation of the FOXP2 gene.[81] Humans have a unique allele of this gene, which has otherwise been closely conserved through most of mammalian evolutionary history.[81] This unique allele seems to have first appeared between 100 and 200 thousand years ago, and it is now all but universal in humans.[81] However, the once-popular idea that FOXP2 is a 'grammar gene' or that it triggered the emergence of language in Homo sapiens is now widely discredited.[82]



If the theory that language could have evolved as a single adaptation is accepted, the question becomes which of its many functions has been the basis of adaptation. Several evolutionary hypotheses have been posited: that language evolved for the purpose of social grooming, that it evolved as a way to show mating potential or that it evolved to form social contracts. Evolutionary psychologists recognize that these theories are all speculative and that much more evidence is required to understand how language might have been selectively adapted.[87]


Given that sexual reproduction is the means by which genes are propagated into future generations, sexual selection plays a large role in human evolution. Human mating, then, is of interest to evolutionary psychologists who aim to investigate evolved mechanisms to attract and secure mates.[88] Several lines of research have stemmed from this interest, such as studies of mate selection[89][90][91] mate poaching,[92] mate retention,[93] mating preferences[94] and conflict between the sexes.[95]

In 1972 Robert Trivers published an influential paper[96] on sex differences that is now referred to as parental investment theory. The size differences of gametes (anisogamy) is the fundamental, defining difference between males (small gametes – sperm) and females (large gametes – ova). Trivers noted that anisogamy typically results in different levels of parental investment between the sexes, with females initially investing more. Trivers proposed that this difference in parental investment leads to the sexual selection of different reproductive strategies between the sexes and to sexual conflict. For example, he suggested that the sex that invests less in offspring will generally compete for access to the higher-investing sex to increase their inclusive fitness (also see Bateman's principle[97]). Trivers posited that differential parental investment led to the evolution of sexual dimorphisms in mate choice, intra- and inter- sexual reproductive competition, and courtship displays. In mammals, including humans, females make a much larger parental investment than males (i.e. gestation followed by childbirth and lactation). Parental investment theory is a branch of life history theory.

Buss and Schmitt's (1993) Sexual Strategies Theory[98] proposed that, due to differential parental investment, humans have evolved sexually dimorphic adaptations related to "sexual accessibility, fertility assessment, commitment seeking and avoidance, immediate and enduring resource procurement, paternity certainty, assessment of mate value, and parental investment." Their Strategic Interference Theory[99] suggested that conflict between the sexes occurs when the preferred reproductive strategies of one sex interfere with those of the other sex, resulting in the activation of emotional responses such as anger or jealousy.

Women are generally more selective when choosing mates, especially under long term mating conditions. However, under some circumstances, short term mating can provide benefits to women as well, such as fertility insurance, trading up to better genes, reducing risk of inbreeding, and insurance protection of her offspring.[100]


Another interesting line of research is that which examines women's mate preferences across the ovulatory cycle.[104][105] The theoretical underpinning of this research is that ancestral women would have evolved mechanisms to select mates with certain traits depending on their hormonal status. Known as the ovulatory shift hypothesis, the theory posits that, during the ovulatory phase of a woman's cycle (approximately days 10–15 of a woman's cycle),[106] a woman who mated with a male with high genetic quality would have been more likely, on average, to produce and bear a healthy offspring than a woman who mated with a male with low genetic quality. These putative preferences are predicted to be especially apparent for short-term mating domains because a potential male mate would only be offering genes to a potential offspring. This hypothesis allows researchers to examine whether women select mates who have characteristics that indicate high genetic quality during the high fertility phase of their ovulatory cycles. Indeed, studies have shown that women's preferences vary across the ovulatory cycle. In particular, Haselton and Miller (2006) showed that highly fertile women prefer creative but poor men as short-term mates. Creativity may be a proxy for good genes.[107] Research by Gangestad et al. (2004) indicates that highly fertile women prefer men who display social presence and intrasexual competition; these traits may act as cues that would help women predict which men may have, or would be able to acquire, resources.



Robert Trivers' theory of parental investment predicts that the sex making the largest investment in lactation, nurturing and protecting offspring will be more discriminating in mating and that the sex that invests less in offspring will compete for access to the higher investing sex (see Bateman's principle).[97] Sex differences in parental effort are important in determining the strength of sexual selection.



Family and kin

Inclusive fitness is the sum of an organism's classical fitness (how many of its own offspring it produces and supports) and the number of equivalents of its own offspring it can add to the population by supporting others.[110] The first component is called classical fitness by Hamilton (1964).




  • is the reproductive cost to the altruist,


Although it is generally true that humans tend to be more altruistic toward their kin than toward non-kin, the relevant proximate mechanisms that mediate this cooperation have been debated (see kin recognition), with some arguing that kin status is determined primarily via social and cultural factors (such as co-residence, maternal association of sibs, etc.),[112] while others have argued that kin recognition can also be mediated by biological factors such as facial resemblance and immunogenetic similarity of the major histocompatibility complex (MHC).[113] For a discussion of the interaction of these social and biological kin recognition factors see Lieberman, Tooby, and Cosmides (2007)[114] (PDF).

Whatever the proximate mechanisms of kin recognition there is substantial evidence that humans act generally more altruistically to close genetic kin compared to genetic non-kin.[115][116][117]

Interactions with non-kin / reciprocity


w > c/b

Reciprocity can also be indirect if information about previous interactions is shared. Reputation allows evolution of cooperation by indirect reciprocity. Natural selection favors strategies that base the decision to help on the reputation of the recipient: studies show that people who are more helpful are more likely to receive help. The calculations of indirect reciprocity are complicated and only a tiny fraction of this universe has been uncovered, but again a simple rule has emerged.[118] Indirect reciprocity can only promote cooperation if the probability, q, of knowing someone's reputation exceeds the cost-to-benefit ratio of the altruistic act:

q > c/b


Trivers argues that friendship and various social emotions evolved in order to manage reciprocity.[120] Liking and disliking, he says, evolved to help present day humans' ancestors form coalitions with others who reciprocated and to exclude those who did not reciprocate.[120] Moral indignation may have evolved to prevent one's altruism from being exploited by cheaters, and gratitude may have motivated present day humans' ancestors to reciprocate appropriately after benefiting from others' altruism.[120] Likewise, present day humans feel guilty when they fail to reciprocate.[120] These social motivations match what evolutionary psychologists expect to see in adaptations that evolved to maximize the benefits and minimize the drawbacks of reciprocity.[120]

Evolutionary psychologists say that humans have psychological adaptations that evolved specifically to help us identify nonreciprocators, commonly referred to as "cheaters."[120] In 1993, Robert Frank and his associates found that participants in a prisoner's dilemma scenario were often able to predict whether their partners would "cheat," based on a half-hour of unstructured social interaction.[120] In a 1996 experiment, for example, Linda Mealey and her colleagues found that people were better at remembering the faces of people when those faces were associated with stories about those individuals cheating (such as embezzling money from a church).[120]

Strong reciprocity (or "tribal reciprocity")

Humans may have an evolved set of psychological adaptations that predispose them to be more cooperative than otherwise would be expected with members of their tribal in-group, and, more nasty to members of tribal out groups. These adaptations may have been a consequence of tribal warfare.[121] Humans may also have predispositions for "altruistic punishment" – to punish in-group members who violate in-group rules, even when this altruistic behavior cannot be justified in terms of helping those you are related to (kin selection), cooperating with those who you will interact with again (direct reciprocity), or cooperating to better your reputation with others (indirect reciprocity).[122][123]

Evolutionary psychology and culture

Though evolutionary psychology has traditionally focused on individual-level behaviors, determined by species-typical psychological adaptations, considerable work has been done on how these adaptations shape and, ultimately govern, culture (Tooby and Cosmides, 1989).[124] Tooby and Cosmides (1989) argued that the mind consists of many domain-specific psychological adaptations, some of which may constrain what cultural material is learned or taught. As opposed to a domain-general cultural acquisition program, where an individual passively receives culturally-transmitted material from the group, Tooby and Cosmides (1989), among others, argue that: "the psyche evolved to generate adaptive rather than repetitive behavior, and hence critically analyzes the behavior of those surrounding it in highly structured and patterned ways, to be used as a rich (but by no means the only) source of information out of which to construct a 'private culture' or individually tailored adaptive system; in consequence, this system may or may not mirror the behavior of others in any given respect." (Tooby and Cosmides 1989).[124]

In psychology sub-fields

Developmental psychology

According to Paul Baltes, the benefits granted by evolutionary selection decrease with age. Natural selection has not eliminated many harmful conditions and nonadaptive characteristics that appear among older adults, such as Alzheimer disease. If it were a disease that killed 20-year-olds instead of 70-year-olds this may have been a disease that natural selection could have eliminated ages ago. Thus, unaided by evolutionary pressures against nonadaptive conditions, modern humans suffer the aches, pains, and infirmities of aging and as the benefits of evolutionary selection decrease with age, the need for modern technological mediums against non-adaptive conditions increases.[125]

Social psychology



Abnormal psychology

Adaptationist hypotheses regarding the etiology of psychological disorders are often based on analogies between physiological and psychological dysfunctions,[131] as noted in the table below. Prominent theorists and evolutionary psychiatrists include Michael T. McGuire, Anthony Stevens, and Randolph M. Nesse. They, and others, suggest that mental disorders are due to the interactive effects of both nature and nurture, and often have multiple contributing causes.[14]

Possible Causes of Psychological 'Abnormalities' from an Adaptationist Perspective
Summary based on information in these textbooks (all titled "Evolutionary Psychology"): Buss (2011),[115] Gaulin & McBurney (2004),[116] Workman & Reader (2008)[132] as well as Cosmides & Tooby (1999) Toward an evolutionary taxonomy of treatable conditions[133]
Causal mechanism of failure or malfunction of adaptationPhysiological ExampleHypothesized Psychological Example
Functioning adaptation (adaptive defense) Fever / Vomiting
(functional responses to infection or ingestion of toxins)
Mild depression or anxiety (functional responses to mild loss or stress[134]/ reduction of social interactions to prevent infection by contagious pathogens)[135]
By-product of an adaptation(s) Intestinal gas
(byproduct of digestion of fiber)
Sexual fetishes (?)
Adaptations with multiple effects Sickle cell disease (Gene that imparts malaria resistance, in homozygous form, causes sickle cell anemia) Schizophrenia or bipolar disorder (May be side-effects of adaptations for high levels of creativity, perhaps dependent on alternate developmental trajectories)
Malfunctioning adaptation Allergies
(over-reactive immunological responses)
(possible malfunctioning of theory of mind module)
Frequency-dependent morphs The two sexes / Different blood and immune system types Personality disorders
(may represent alternative behavioral strategies possibly dependent on its prevalence in the population)
Mismatch between ancestral & current environments Type 2 Diabetes
(May be related to the abundance of sugary foods in the modern world)
Tails of normal distribution (bell curve) Dwarfism or gigantism Extremities of the distribution of cognitive and personality traits
(e.g., extremely introversion and extraversion, or intellectual giftedness and intellectual disability)

Evolutionary psychologists have suggested that schizophrenia and bipolar disorder may reflect a side-effect of genes with fitness benefits, such as increased creativity.[136] (Some individuals with bipolar disorder are especially creative during their manic phases and the close relatives of people with schizophrenia have been found to be more likely to have creative professions.[136]) A 1994 report by the American Psychiatry Association found that people suffered from schizophrenia at roughly the same rate in Western and non-Western cultures, and in industrialized and pastoral societies, suggesting that schizophrenia is not a disease of civilization nor an arbitrary social invention.[136] Sociopathy may represent an evolutionarily stable strategy, by which a small number of people who cheat on social contracts benefit in a society consisting mostly of non-sociopaths.[14] Mild depression may be an adaptive response to withdraw from, and re-evaluate, situations that have led to disadvantageous outcomes (the "analytical rumination hypothesis")[134] (see Evolutionary approaches to depression).

Some of these speculations have yet to be developed into fully testable hypotheses, and a great deal of research is required to confirm their validity.[137][138]

Antisocial and criminal behavior

Evolutionary psychology has been applied to explain criminal or otherwise immoral behavior as being adaptive or related to adaptive behaviors. Males are generally more aggressive than females, who are more selective of their partners because of the far greater effort they have to contribute to pregnancy and child-rearing. Males being more aggressive is hypothesized to stem from the more intense reproductive competition faced by them. Males of low status may be especially vulnerable to being childless. It may have been evolutionary advantageous to engage in highly risky and violently aggressive behavior to increase their status and therefore reproductive success. This may explain why males are generally involved in more crimes, and why low status and being unmarried is associated with criminality. Furthermore, competition over females is argued to have been particularly intensive in late adolescence and young adulthood, which is theorized to explain why crime rates are particularly high during this period.[139]

Many conflicts that result in harm and death involve status, reputation, and seemingly trivial insults.[139] Steven Pinker in his book The Blank Slate argues that in non-state societies without a police it was very important to have a credible deterrence against aggression. Therefore, it was important to be perceived as having a credible reputation for retaliation, resulting in humans to develop instincts for revenge as well as for protecting reputation ("honor"). Pinker argues that the development of the state and the police have dramatically reduced the level of violence compared to the ancestral environment. Whenever the state breaks down, which can be very locally such as in poor areas of a city, humans again organize in groups for protection and aggression and concepts such as violent revenge and protecting honor again become extremely important.[139]

Rape is theorized to be a reproductive strategy that facilitates the propagation of the rapist's progeny. Such a strategy may be adopted by men who otherwise are unlikely to be appealing to women and therefore cannot form legitimate relationships, or by high status men on socially vulnerable women who are unlikely to retaliate to increase their reproductive success even further.[140] The sociobiological theories of rape are highly controversial, as traditional theories typically do not consider rape to be a behavioral adaptation, and objections to this theory are made on ethical, religious, political, as well as scientific grounds.

Psychology of religion


Coalitional psychology

Coalitional psychology is an approach to explain political behaviors between different coalitions and the conditionality of these behaviors in evolutionary psychological perspective. This approach assumes that since human beings appeared on the earth, they have evolved to live in groups instead of living as individuals to achieve benefits such as more mating opportunities and increased status.[143] Human beings thus naturally think and act in a way that manages and negotiates group dynamics.

Coalitional psychology offers falsifiable ex ante prediction by positing five hypotheses on how these psychological adaptations operate:[144]

  • Humans represent groups as a special category of individual, unstable and with a short shadow of the future
  • Political entrepreneurs strategically manipulate the coalitional environment, often appealing to emotional devices such as "outrage" to inspire collective action.
  • Relative gains dominate relations with enemies, whereas absolute gains characterize relations with allies.
  • Coalitional size and male physical strength will positively predict individual support for aggressive foreign policies.
  • Individuals with children, particularly women, will vary in adopting aggressive foreign policies than those without progeny.

Reception and criticism


Ethical implications

Critics have argued that evolutionary psychology might be used to justify existing social hierarchies and reactionary policies.[146][147] It has also been suggested by critics that evolutionary psychologists' theories and interpretations of empirical data rely heavily on ideological assumptions about race and gender.[148]

In response to such criticism, evolutionary psychologists often caution against committing the naturalistic fallacy – the assumption that "what is natural" is necessarily a moral good.[147][149][150] However, their caution against committing the naturalistic fallacy has been criticized as means to stifle legitimate ethical discussions.[147]

Contradictions in models

Some criticisms of evolutionary psychology point at contradictions between different aspects of adaptive scenarios posited by evolutionary psychology. One example is the evolutionary psychology model of extended social groups selecting for modern human brains, a contradiction being that the synaptic function of modern human brains require high amounts of many specific essential nutrients so that such a transition to higher requirements of the same essential nutrients being shared by all individuals in a population would decrease the possibility of forming large groups due to bottleneck foods with rare essential nutrients capping group sizes. It is mentioned that some insects have societies with different ranks for each individual and that monkeys remain socially functioning after removal of most of the brain as additional arguments against big brains promoting social networking. The model of males as both providers and protectors is criticized for the impossibility of being in two places at once, the male cannot both protect his family at home and be out hunting at the same time. In the case of the claim that a provider male could buy protection service for his family from other males by bartering food that he had hunted, critics point at the fact that the most valuable food (the food that contained the rarest essential nutrients) would be different in different ecologies and as such vegetable in some geographical areas and animal in others, making it impossible for hunting styles relying on physical strength or risk taking to be universally of similar value in bartered food and instead making it inevitable that in some parts of Africa, food gathered with no need for major physical strength would be the most valuable to barter for protection. A contradiction between evolutionary psychology's claim of men needing to be more sexually visual than women for fast speed of assessing women's fertility than women needed to be able to assess the male's genes and its claim of male sexual jealousy guarding against infidelity is also pointed at, as it would be pointless for a male to be fast to assess female fertility if he needed to assess the risk of there being a jealous male mate and in that case his chances of defeating him before mating anyway (pointlessness of assessing one necessary condition faster than another necessary condition can possibly be assessed).[151][152]

Standard social science model

Evolutionary psychology has been entangled in the larger philosophical and social science controversies related to the debate on nature versus nurture. Evolutionary psychologists typically contrast evolutionary psychology with what they call the standard social science model (SSSM). They characterize the SSSM as the "blank slate", "relativist", "social constructionist", and "cultural determinist" perspective that they say dominated the social sciences throughout the 20th century and assumed that the mind was shaped almost entirely by culture.[149]

Critics have argued that evolutionary psychologists created a false dichotomy between their own view and the caricature of the SSSM.[153][154][155] Other critics regard the SSSM as a rhetorical device or a straw man[150][153][156] and suggest that the scientists whom evolutionary psychologists associate with the SSSM did not believe that the mind was a blank state devoid of any natural predispositions.[150]

Reductionism and determinism

Some critics view evolutionary psychology as a form of genetic reductionism and genetic determinism,[157][158] a common critique being that evolutionary psychology does not address the complexity of individual development and experience and fails to explain the influence of genes on behavior in individual cases.[41] Evolutionary psychologists respond that they are working within a nature-nurture interactionist framework that acknowledges that many psychological adaptations are facultative (sensitive to environmental variations during individual development). The discipline is generally not focused on proximate analyses of behavior, but rather its focus is on the study of distal/ultimate causality (the evolution of psychological adaptations). The field of behavioral genetics is focused on the study of the proximate influence of genes on behavior.[159]

Testability of hypotheses

A frequent critique of the discipline is that the hypotheses of evolutionary psychology are frequently arbitrary and difficult or impossible to adequately test, thus questioning its status as an actual scientific discipline, for example because many current traits probably evolved to serve different functions than they do now.[5][160] While evolutionary psychology hypotheses are difficult to test, evolutionary psychologists assert that it is not impossible.[161] Part of the critique of the scientific base of evolutionary psychology includes a critique of the concept of the Environment of Evolutionary Adaptation (EEA). Some critics have argued that researchers know so little about the environment in which Homo sapiens evolved that explaining specific traits as an adaption to that environment becomes highly speculative.[162] Evolutionary psychologists respond that they do know many things about this environment, including the facts that present day humans' ancestors were hunter-gatherers, that they generally lived in small tribes, etc.[163] Edward Hagen argues that the human past environments were not radically different in the same sense as the Carboniferous or Jurassic periods and that the animal and plant taxa of the era were similar to those of the modern world, as was the geology and ecology. Hagen argues that few would deny that other organs evolved in the EEA (for example, lungs evolving in an oxygen rich atmopshere) yet critics question whether or not the brain's EEA is truly knowable, which he argues constitutes selective scepticism. Hagen also argues that most evolutionary psychology research is based on the fact that females can get pregnant and males cannot, which Hagen observes was also true in the EEA.[164][165] John Alcock describes this as the "No Time Machine Argument", as critics are arguing that since it is not possible to travel back in time to the EEA, then it cannot be determined what was going on there and thus what was adaptive. Alcock argues that present day evidence allows researchers to be reasonably confident about the conditions of the EEA and that the fact that so many human behaviours are adaptive in the current environment is evidence that the ancestral environment of humans had much in common with the present one, as these behaviours would have evolved in the ancestral environment. Thus Alcock concludes that researchers can make predictions on the adaptive value of traits.[166]

Modularity of mind

Evolutionary psychologists generally presume that, like the body, the mind is made up of many evolved modular adaptations,[167] although there is some disagreement within the discipline regarding the degree of general plasticity, or "generality," of some modules.[159] It has been suggested that modularity evolves because, compared to non-modular networks, it would have conferred an advantage in terms of fitness[168] and because connection costs are lower.[169]

In contrast, some academics argue that it is unnecessary to posit the existence of highly domain specific modules, and, suggest that the neural anatomy of the brain supports a model based on more domain general faculties and processes.[170][171] Moreover, empirical support for the domain-specific theory stems almost entirely from performance on variations of the Wason selection task which is extremely limited in scope as it only tests one subtype of deductive reasoning.[172][173]

Cultural rather than genetic development of cognitive tools

Cecilia Heyes has argued that the picture presented by some evolutionary psychology of the human mind as a collection of cognitive instincts  organs of thought shaped by genetic evolution over very long time periods[174][16]  does not fit research results. She posits instead that humans have cognitive gadgets  "special-purpose organs of thought" built in the course of development through social interaction.

Evolutionary psychology defense

Evolutionary psychologists have addressed many of their critics (see, for example, books by Segerstråle (2000), Defenders of the Truth: The Battle for Science in the Sociobiology Debate and Beyond,[175] Barkow (2005), Missing the Revolution: Darwinism for Social Scientists,[176] and Alcock (2001), The Triumph of Sociobiology[177]). Among their rebuttals are that some criticisms are straw men, are based on an incorrect nature versus nurture dichotomy, are based on misunderstandings of the discipline, etc.[159][177][178][179][180][181][182][183][184] Robert Kurzban suggested that "...critics of the field, when they err, are not slightly missing the mark. Their confusion is deep and profound. It’s not like they are marksmen who can’t quite hit the center of the target; they’re holding the gun backwards."[185]

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