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Culture and the Evolution Learning of Social

Culture and the Evolution Learning of Social

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Culture and the of Social Mark V. Flinn Department of , University of Missouri

Applications of modern evolutionary to have generated several different theoretical approaches that challenge traditional anthropological perspectives. “Cultural selection” and “ parasite” model culture as an independent evo- lutionary system because transmission of cultural traits via social learning is distinct from transmission of vla DNA replication. “Dual-” and “co-evolution” theories model culture as an intermediary evolutionary process that involves informa- tion from two inheritance systems: and social learning. “Evolutionary psychol- ogy” theories emphasize that the evolutionary history of on mental pro- cesses links culture and biological ; hence, cultural is viewed as part of the organic and not an independent evolutionary system. Cross-cul- tural universals and scenarios of the “environment of evolutionary adaptedness” are used to identify characteristics of the “evolved mind” (human ). “Behavioral ecol- ogy” theories examine relations between and environmental context. Behav- ioral/cultural variations are viewed as products of flexible decision-making processes (evolved mind) that may respond adaptively to micro-environmental differences. It is difficult to devise empirical tests that distinguish among these theories, because they share many basic premises and make similar predictions about . In- deed, some of the apparent differences may be more semantic than substantive. Social learning is the key process underlying these evolutionary . Here I argue that human learning mechanisms are products of natural selection, and hence process information in ways that reflect evolutionary design. I review common objec- tions to this hypothesis, including (1) learning processes are uncoupled from genetics and biological adaptation; (2) culture (or its effects) is partly extrasomutic; (3) culture, by most definitions, involves mental phenomena, including conscious thought; (4) culture involves the use ofarbitrary to communicate information; (5) culture appears to have emergent properties at the group level, such as shared values and beliefs resulting in political and religious ; (6) culture involves historical processes; and (7) com- plex culture is uniquely human-we need an explanation for why the human alone evolved such extensive social learning aptitudes. I suggest that these controversies can be resolved, although empirical tests are difficult.

Received October 16, 1996; revised October 24, 1996; accepted October 25, 1996. Address reprint requests and correspondence to: Mark V. Flinn, Department of Anthropology, University of Missouri, Columbia, MO 65211.

Evolution and Human Behavior 18: 23-67 (1997) 0 Elsevier Inc., 1997 1090-5138/97/$17.00 655 Avenue of the Americas, New York, NY 10010 PII S10902-5138(96)00046-5 24 M.V. Fhnn

I posit that social was a primary selective pressure on human mental abilities (Alexander 1989) and that this favored -general, constructivist learning capabilities (e.g., Quartz and Sejnowski, in press) that can manage context-dependent analysis and integrate information from domain-specific mechanisms (Hirschfeld and Gelman 1994; MacDonald 1991; Sperber 19%). are unique in the extraordi- nary levels of novelty that are generated by the processing of socially learned informa- tion. Human culture is cumulative; human produces new built upon the old. To a degree that far surpasses that of any other species, human mental processes must contend with a constantly changing information environment of their own cre- ation. Cultural information may be especially dynamic because it is a fundamental as- pect of human social competition. Apparently arbitrary changes in cultural traits, such as clothing styles, music, art, , dialects, and so forth, may reflect information arms races among social competitors. The remarkable developmental plasticity and cross- domain integration of some cognitive mechanisms may be products of selection for spe- cial sensitivity to variable social context. Human “culture” is not just a pool or source of information; it is an arena and theater of social manipulation and competition. Studies of human behavior-including , , relationships, subsistence, economics, and politics-generally are consistent with an evolutionary basis for social learning, but often they fail to add specific new knowledge about the mecha- nisms. Analyses of cognitive aptitudes underlying language, kinship, and so forth, often are inconclusive because cultural information (and consequent behavior) involves com- plex interaction among history, environmental variation, ontogenetic pathways of men- tal processes, and specific context. I suggest that empirical tests of evolutionary must build upon identification of apparent universals and examine variability by incorporating developmental , environmental conditions, and social and historical context. This synthesis would benefit from enhanced cooperation between cognitive psychologists and cultural anthropologists. 0 Elsevier Science Inc., 1997

KEY WORDS: Culture;Evolution; Learning; Psychological mechanisms; Social competition.

ecent modifications of evolutionary theory have kindled renewed inter- est in relations between culture’ and . New concepts of learning as an adaptation (J.L. Gould 1986; Johnston 1982) have inspired new mod- els of culture (e.g., Alexander 1979a, 1990a; Boyd and Richerson 1985; LumsdenR and Wilson 1981; Sperber 1996). Although there is general agreement that evolutionary theory is useful for understanding human behavior and culture, impor- tant issues remain unresolved, and directions for empirical research are enigmatic. Some models, beginning with a genetics , posit that cul- ture is a distinct “co-evolutionary” system because transmission of cultural traits via social learning is separate from transmission of genes via DNA replication (e.g., Cavalli-Sforza and Feldman 1981; Dawkins 1982; Richerson and Boyd 1978; Durham 1991; Rindos 1986b). Culture and genetics are viewed as different, albeit interacting, types of inheritance processes. Grganically evolved (i.e., genetically controlled) “learning biases” may influence culture content, but the property of transmissibility of socially learned information is considered sufficient to distin-

‘By “culture” I refer to common (non-exclusive) usage of the term in . I do not find it important here to distinguish among the various definitions. I use “information” as a more specific term. Culture and the Evolution of Social Learning 25 guish it from other environmental effects on the phenotype. This approach repre- sents as analogous to organic evolution. Other models, originating from a Tinbergian ethological paradigm, emphasize that evolved mental or psychological processes constrain culture. Human psychol- ogy (including learning aptitudes) and, hence, behavior are posited to have been shaped by natural selection (e.g., Alexander 1979a; Barkow 1989b; Irons 1979a; Lumsden and Wilson 1981; Cosmides and Tooby 1995). Sources of controversy in- clude the genetic basis, degree of specificity, , and plasticity (particularly in regard to modem social conditions) of psychological processes (e.g., Daly and Wilson 1995; Symons 1989). The objective of this paper is to examine a key issue underlying this new body of culture theory: the evolutionary basis for social learning. Here I attempt to inte- grate models of culture with knowledge of learning processes from behavioral ecol- ogy and psychology. Empirical studies of culture from evolutionary perspectives are briefly reviewed, and unresolved theoretical problems discussed. I argue that human learning mechanisms are products of natural selection, and hence process informa- tion in ways that reflect evolutionary design. I posit that social competition is a pri- mary selective pressure on human mental abilities (Alexander 1989; Humphrey 1984) that has favored some domain-general, constructivist learning capabilities (e.g., Quartz and Sejnowski, in press) that can manage context-dependent analyses and integrate domain-specific mechanisms (Hirschfeld 1994; MacDonald 199 1; Sperber 1996). This argument remains speculative because it lacks information about (1) the developmental ontogeny and of cognitive processes, (2) the influ- ence of environmental conditions and historical context on information choice, and (3) the dynamics of the use and manipulation of information in social competition.

COMMON GROUND

Anthropology has identified several important relations between biology and cul- ture. First, human “aptitudes” for culture are products of natural selection, evi- denced by (1) increases in cranial capacity from australopithecines to modem humans (Holloway 1975; Spuhler 1959; Wolpoff 1995), and (2) cognitive differ- ences between humans and other (Byrne 1994; Cheney and Seyfarth 1990; Gibson and Ingold 1993; Griffen 1984; Holloway and de la Coste-Lereymondie 1982; Quiatt and Reynolds 1993; Parker and Gibson 1979, 1990). Second, “biologi- cal needs” are facilitated by culture, such as acquiring sufficient food, protection from predators and the elements, , and caring for dependent offspring and kin (Malinowski 1922; Steward 1955). And third, culture (information) is acquired and transmitted via learning processes, usually involving symbolic com- munication (e.g., Murdock 1956). Hence cultural change can occur without accom- panying genetic change, and most cultural differences among human are unlikely to be the result of genetic differences. from one genetic population acquire the cultural traits of another without difficulty (e.g., a child of Yanomamo parentage can readily learn to speak Spanish). These tenets of anthro- 26 M.V. Flinn pology are entirely consistent with the new evolutionary approaches. Most current discussions have moved past old “” issues. The controversy begins when we consider specific content and variability of culture. Although cultural capacities are products of organic evolution, it is possible that cultural information is acquired and transmitted largely or totally independent of biology and natural selection. Organic evolution could be unimportant for analy- sis of specific ideas, beliefs, values, and so forth, that occupy the evolved gray mat- ter. On the other hand, if the evolutionary basis for human neural structures and sub- sequent sensory and cognitive processes significantly influences what particular cultural information is generated, acquired, and/or transmitted, then biological adap- tation would be an important component in a general theory of culture.

Social Learning: Blank Slate, Simple Rules of Thumb, or Complex Evolutionary Design?

Most theories of culture are based upon transmission of information via social leam- ing. There are, however, widely different assumptions about fundamental aspects of social learning processes. , specificity, and of social learning aptitudes are important criteria distinguishing theories of culture (Table 1). Many traditional symbolic, functional, and structural anthropological theories begin with relatively “blank slate” learning. Culture programs the mind, and evolves sui & independent of biopsychological (e.g., Boas 1911; Durkheim 1895; Hallowell 1960; Kroeber 1952; White 1949). Social learning is viewed as a general all-purpose, non-selective by which cultural information is acquired and transmitted. Individuals are passive recipients of , the logical ex- treme positing “cultural standards have no obvious function” (Freilich 1977:91). In- dividual actions reflect programmed by the (for critical reviews, see D’Andrade 1984, 1995; Spiro 1986). Cultural selection theories employ an evolutionary model, with cultural traits or “” as transmitable units (Campbell 1965; Cavalli-Sforza and Feldman 1981; Cloak 1975; Dawkins 1989, 1982; Dunnell 1980; Goodenough 1995; Rondos

Table 1. Learning Aptitudes in Theories of Culture

Theory Learning aptitudes Cultural evolution Function

Traditional Blank slate or arbitrary Culture programs Maintain ; social science the mind provide meaning

Cultural Blank slate, arbitrary, Cultural traits/mind None; selection or evolved; irrelevant compete for replication mechanistic

Dual Simple learning biases: Learning biases and Mix of cultural inheritance , transmission processes and biological determine culture content adaptation

Evolutionary Adaptive, complex, Cumulative result of individual Individual psychology & multiple learning actions, influenced by evolved inclusive Behavioral aptitudes, some specific, mental processes, constrained maximization others more general by history and environment (gene replication) Culture and the Evolution of Social Learning 27

1986a; Ruyle 1973). Cultural evolution is viewed as analogous to organic evolution, with cultural traits taking the place of genes. Social learning is the whereby cultural traits are transmitted and replicated. Learning biases, constraints, or specific aptitudes, however, usually are not important factors in cultural selection models. Hence “a has its own opportunities for replication and its own pheno- typic effects, and there is no why in a meme should have any connec- tion whatever with biological success” (Dawkins 1982: 116). Dual inheritance theories are similar to cultural selection theories in that men- tal information bits underlying cultural traits are viewed as distinct entities (replica- tors) analogous to genes. “Human beings are possessed of &JO major information systems, one genetic, and one cultural” (Durham 1991:9). Culture is an independent inheritance system, albeit one with significant connections to genetic evolution. Dual inheritance models emphasize understanding how cultural transmission oc- curs, and the extent to which such transmission is influenced by organically evolved psychological mechanisms (e.g., learning biases): “If such genetically transmitted rules [predispositions to learn adaptive cultural traits] cannot readily evolve, then other forces may predominate in cultural evolution, forces which do not necessarily act to maximize fitness” (Boyd and Richerson 1985: 133). This approach models the transmission of culture via a relatively (see below) limited set of learning principles, such as imitating successful (high-status) individuals (cf. Flinn and Alexander 1982:394). “Maladaptive” cultural traits that lower individual fitness (e.g., celibacy among priests) are viewed as products of cultural selection or unavoidable results of “low-cost” choice mechanisms. Evolved psychological mechanism theories, including “gene-culture coevolu- tion,” “, ” “human ,” and “” propose that evolved mental processes provide a critical connection between genes and (e.g., Alexander 1990a; Barkow 1989a; Daly and Wilson 1995; Lumsden and Wilson 1981; MacDonald 1988b; Tooby and Cosmides 1992). This approach emphasizes that the apparatus for acquisition, anal- ysis, and transmission of culture-that is, the mind-is a product of natural selec- tion. The mind is predicted to produce adaptive in appropriate environ- ments. Human psychology and cultural transmission are posited to involve a complex array of evolved learning aptitudes developed and modified throughout on- togeny. The body of socially transmitted information traditionally defined as “cul- ture” is viewed as both a part of the environment (a source of information), and part of the of individuals (information in the mind). “Maladaptive” cultural traits that lower individual fitness (e.g., addiction to opium) are viewed in the same way that evolutionary view “non-cultural” maladaptive traits: They result from constraints on perfection such as novel environments or manipulation by com- petitors (e.g., Dawkins 1982; Reeve and Sherman 1993). Socially learned informa- tion may result in “anti-imitation” if appropriate (e.g., avoidance of cigarette smok- ing after reading reports of cancer ). Connections between specific behaviors and information are variable, context dependent, and constructed by individual ; hence, socially learned information-bits are not considered independent rep- licators or mind parasites with behavioral phenotypes. 28 M.V.Flinn

The evolved psychological mechanisms model is distinguished from traditional and cultural selection models by its emphasis on the prediction that the human mind and behavior will show evidence of evolutionary design. The is postulated to be an integrated network of task-specific modules that produce adaptive behavior in appropriate environments, rather than a uniform-blank-slate or simple-learning-biases information processor. The and terminology of this approach is similar to that of evolutionary biological explanations of animal behavior (e.g., Alcock 1994; Daly and Wilson 1983; Tinbergen 1963).

The Evolutionary Basis for Human Mental Processes

Most evolutionary approaches emphasize that the central is not a blank slate. The apparatus for acquisition, analysis, and transmission of cultural information is hypothesized to have been designed by natural selection to produce -maximizing behavior.2 As a product of evolution, the central ner- vous system and its learning capabilities are unlikely to acquire, analyze, or trans- mit cultural traits randomly with respect to natural selection. Hence the choice of cultural traits by individuals is predicted to reflect adaptive (in the modem biologi- cal sense)3 decision-making processes (Alexander 1974, 1990a; Badcock 1991; Barkow 1989a, 1989b; Borgerhoff Mulder 1987; Boyd and Richerson 1985; Cosmides and Tooby 1995; Daly 1982; Donald 1991; Durham 1991; Flinn and Alexander 1982; Irons 1979a, 1983; Konner 1982; Lumsden and Wilson 1981; MacDonald 1988b, 1991; Reynolds 1984; Smith and Winterhalder 1992; Spiro 1986, 1987; Symons 1989; Tooby and Cosmides 1989; Turke 1984, 1990). Differences among these evolutionary approaches center on (1) the extent to which the selection of cultural traits involves forces other than the evolved design of the human mind, and (2) what selective pressures are primarily responsible for the design of the human mind. The evolved psychological mechanisms approach views cultural behavior, mental representations, and material artifacts as part of the human phenotype, sub- ject to natural selection. The central nervous system and its ontogenetic develop- ment of learning mechanisms and other cognitive abilities are the hypothesized con- nections between genes and cultural behavior. Evolutionary design is posited to be “the method to the madness” of mental processes used by individuals to generate, analyze, and choose cultural information and hence determine culture content. The logic of this evolutionary approach hinges on the assumption that humans did not evolve large and associated learning capabilities by random chance. Cultural aptitudes were selected for because they caused human ancestors to acquire

*“Inclusive fitness” is a measure of an individual’s genetic contribution to future generations via direct (personal) reproduction and indirect (effect of the individual on the reproduction of genetic relatives) reproduction (Hamilton 1964). ‘The “new” concept of adaptation is defined in terms of differential genetic representation in future generations (see Alexander 1979a. Dawkins 1982, 1989; Mayr 1982; Reeve and Sherman 1993; Sober 1984, Williams 1966, 1992). Theories combining cultural and organic evolution prior to the emphasis on individual- or gene-level selection are useful but outdated antecedents to current evolutionary models of culture (e.g., Schwartz and Mead 1961; Wallace 1961). Culture and the Evolution of Social Learning 29 information resulting in behavior (e.g., assisting kin, seeking mates, forming advan- tageous social alliances, communicating effectively, and accruing resources) that in- creased genetic representation in future generations relative to less successful alter- natives. Human cultural abilities are the cumulative result of millions of years of competition among millions of heritable variations in central nervous system learn- ing programs. The “winners” (what we have today) are those learning programs that out-reproduced the alternatives (via inclusive fitness-maximizing behavior). Hence the prediction that cultural transmission is unlikely to be random or arbitrary with respect to biological adaptation. This hypothesis is sometimes referred to as “the ar- gument from natural origins” (Boyd and Richerson 1985: 13). This evolved psychological mechanisms approach does not require that each and every choice of a cultural trait must be advantageous for the individual chooser. We all make mistakes. Nor does it imply that the best possible (i.e., most adaptive or optimal) cultural trait will be “invented” by individuals whenever needed. The available choices are constrained by history, difision, ontogenetic environments, and competitor S strategies, as well as by limitations of the central nervous system. The evolutionary hypothesis does require, however, that cultural choice processes must have resulted, on average, in choice by individuals of more adaptive cultural traits over less adaptive ones. It further requires individual differences (however minute) in effectiveness of choice processes, and that some under- lay such differences during the evolutionary history of the human central nervous system (e.g., Lumsden and Wilson 1981). It does not suggest that genetic variation determines contemporary , nor that there are “genes for” specific behaviors such as matrilineal inheritance or cross-cousin rules. What it does suggest is that natural selection favored mental abilities (psychological mecha- nisms)4 to make adaptive (inclusive fitness-maximizing) decisions about costs and benefits of alternative actions under varying contingencies of individual-specific micro-environments (cf. Symons 1992). In normal environments, humans develop cognitive processes that result in ap- parently adaptive behavior, such as what path is used to walk from one village to the next, what areas are searched for wild yams, how to speak, how much time is spent on a hunt, who food is shared with, who is perceived as sexually attractive, who kin- attachment bonds are formed with, and so forth (e.g., Alexander 1990a; Buss 1994; Kaplan and Hill 1992; Pinker 1994). Like all hypothesized , evolved mental processes can result in mistakes. in judgment are no more evidence of a distinct or competing cultural evolutionary process than are moths flying into can- dle flames (Alexander 1979a). I do not agree with the opinion that the evolved psychological mechanisms model proposes that culture “is simply a prisoner of genetic constraints” (Boyd and Richerson 1989:27; see also Montagu 1981; Rogers 1988). The terminologies used by some “sociobiological” models (e.g., Lumsden and Wilson 1981; cf. Lumsden 1989) suggesting that evolved psychological mechanisms (ultimately, genes) “put

‘Other terms used to refer to “psychological mechanisms” that have overlapping definitions (PMs) in cultural evolution literature include “Darwinian ,” “epigenetic rules,” “,” “learning biases,” “cultural aptitudes,” and “constraints on learning.” 30 M. V. Flinn culture on a leash” can be misleading because they imply learning restrictions (sug- gesting that some things could not be learned) rather than aptitudes.’ Use of social learning by human psychological mechanisms allows for creative responses to novel conditions and development of cumulative information. Human mental abilities do not just constrain culture; they generate its options in the first place! Regardless of terminology, the important question is whether or not mental processes have been shaped by natural selection to preferentially use (learn, invent, transmit to relatives and allies, manipulate competitors) cultural information adaptively (Alexander 1979b; Boyd and Richerson 1985). It is difficult to conceive of a process responsible for systematic, incremental development of the over several million years that did not involve dif- ferential replication of genes (i.e., natural selection), and yet, this is what a “blank slate” model of the central nervous system requires.6 If is uncou- pled from natural selection (i.e., is random with respect to inclusive fitness), then once culture developed, the brain (more specifically, those parts of the brain in- volved with cultural information) would no longer evolve. Individuals with larger or more complex brains would not be favored by natural selection unless such brains caused acquisition of cultural information that resulted in behavior that increased in- clusive fitness. Increasingly refined aptitudes for social learning via symbolic com- munication would not have evolved if the information obtained did not result in more adaptive behavior. The assumption that human cultural aptitudes are products of natural selection is shared by all of the new evolutionary theories of culture. The theories differ in predictions about the specificity of cultural aptitudes and how cul- tural variation is described and interpreted. In the following section I discuss a number of objections and unresolved con- troversies that continue to separate evolutionary models of culture from mainstream social and behavioral science.

THORNY ISSUES

Human culture has important and unique characteristics that present difficult prob- lems for evolutionary models. They include the following: (1) Culture content is transmitted by learning processes (i.e., cognitive information transfer), and not by the transfer of genetic materials; hence, culture appears to be a separate inheritance system, uncoupled from genetics. (2) Culture (or its effects) is partly extrasomatic, cultural traits (e.g., stonepoints, political monuments) exist outside the soma (physi- cal body) of the culture-bearing . (3) Human culture, by most definitions, involves menttiZ phenomena, including conscious thought. (4) Human culture involves the use of arbitrary symbols to communicate information. (5) Culture

5A more appropriate analogy would be that social learning aptitudes let humans off the leash (because they allow for creative, novel behavior), and assist them to find and flush game (as opposed to running randomly). 61t is possible that human cultural capabilities were a “preadaptation” (see Bock 1970 for definition) that fortuitously evolved for other . The novelty of socially transmitted information makes such an unparalleled evolutionary jump worthy of consideration. Culture and the Evolution of Social Learning 31 appears to have emergent properties at the group level, such as shared values and beliefs resulting in political and religious institutions. (6) Culture involves historical processes. History constrains the options (cultural traits) available for individual choice and modification, and culture can change rapidly, apparently outracing genetic evolution. And, (7) complex culture is uniquely human; we need an explana- tion for why humans alone evolved such extensive social learning aptitudes. These characteristics of human culture make it a most challenging and difficult aspect of to understand in scientific terms. However, they do not necessarily cause culture to become a “non-evolutionary” or “separate evolutionary” phenome- non, independent of biological adaptation.

Culture Is Learned

Of the forementioned characteristics of culture, the “culture is learned” problem is the most fundamental (viz. Table 1). It is the source of the greatest amount of confu- sion and disagreement. Hence, I devote more space to discussion of learning than to other issues listed above. Organic or Darwinian evolution is usually defined as a change in gene or frequencies over time. Cultural evolution is usually defined as a change in cultural trait frequencies (or mental information) over time. Because cultural transmission (e.g., imitating a song, or adopting a technique for making stone tools) occurs with- out concomitant genetic transmission, it has been argued that cultural evolution is independent of genes. This leads to the conclusion that cultural evolution is inde- pendent of organic evolution. If biology = genetics, and culture = learning, then if learning # genetics, culture # biology. The logic underlying the above conclusion is perhaps the most significant rea- son why modem social science has not accepted organic evolution as a general the- ory of human behavior. It distinguishes learning from other flexible responses (e.g., physiological changes) to environmental influences in the production of the pheno- type. This is a difficult and important issue, and is a critical assumption underlying “dual-inheritance” and “cultural selection” theories of cultural evolution that treat culture as a partially autonomous inheritance system (e.g., Boyd and Richerson 1985:3, 1989: 27; Cavalli-Sforza and Feldman 1981; Durham 1991; Laland, Kumm, and Feldman 1995; Rindos 1986b).’ Coevolutionary models distinguish “cultural”

‘For example, Dunnell (1989b:40) states that “direct application of biological evolution to cultural phenomena requires genetic transmission of all parts of the human phenotype.” This is difficult to interpret. The environment always participates in the production of all aspects of the phenotype. Culture is part of the environment as well as part of the individual’s phenotype. Dunnell states further (p. 40) that “assumes genetic determination, either directly or indirectly, of human behavior.” All behavior of all species is “determined,” directly or indirectly (?!), by the interaction of genes and environment. His statement that sociobiology “further assumes that transmission operates on particulate ‘bodies’ analogous to genes (e.g., the ‘memes’ of Dawkins [ 19891 and the ‘culturgens’ of Lumsden and Wilson [ 19811)” is ironic and puzzling given Dunnell’s focus on “culturally selected” artifacts or traits as analogous to “naturally selected” genes. Both approaches (meme evolution and cultural selection) neglect the significance of the evolutionary development of human psychology. Both depend upon the effective parasitization of human minds/phenotypes by culturally transmitted replicators (cf. Humphrey, in Dawkins 1989). 32 M.V. FIinn evolution from “biological” evolution on the basis that cultural evolution involves a distinct mode of information transmission (learning on the one hand, genetics on the other). The analogy between cultural evolution and organic evolution, however, may be inappropriate if cultural information is exclusively mediated via organically evolved psychological mechanisms, including social learning processes. Phenotypes are generally accepted to be the products of genes + environment. The directions and degrees to which modify their phenotypes in response to environmental conditions result from a past history of natural selection on abili- ties to modify phenotypes in response to environmental changes (Alexander 1979a; Hill 1993; Stearns 1989). Arctic hares have seasonal changes in fur color, humans develop calluses on their hands and feet, fig alter the -ratios of their broods, sweat learn by association and similarity whom to let into the , chimps observe and imitate termiting with sticks, and so forth. The point here is that phenotypic modifications, whether achieved via learning or physiological changes, are not random. The environment is a causal factor during the development of the phenotype in the context of an evolutionary history of selec- tion for modifications in response to environmental changes (Williams 1966). Envi- ronments do not have random effects on phenotypes. Among Arctic hares, winter conditions result in white coats, summer in brown coats. Other species of rabbit lack this capability; they do not have genetic materials that allow for this adaptive re- sponse. Similarly, lack aptitudes for some aspects of human culture, presumably because chimps do not have genetic materials necessary for develop- ment of the requisite central nervous system and associated psychological mecha- nisms. Regardless of environment, chimpanzees cannot acquire and transmit knowl- edge of differential equations, etc., despite their remarkable cognitive abilities (Rumbaugh and Washburn 1993; Savage-Rumbaugh and Lewin 1994; Tomasello, Kruger, and Ratner 1993). Clearly there are evolved differences between chimps and humans in regard to mathematical reasoning and other mental processes that un- derly human culture (e.g., Geary 1995). Genes evolved to produce phenotypes, including capabilities for learning, be- cause phenotypes provide a means of responding to changing environments (Alex- ander 1979a: 14):

The whole reason for phenotypes having evolved is that they provide flexibility in meeting environmental contingencies that are only predictable on short-term bases. Learned behavior is the ultimate of all such flexibilities. Not just humans and higher but animals in general develop their behavior, or “learn” to do what is appro- priate in their particular life circumstances. Even the remarkably distinctive castes of the social are in nearly all cases determined not by genetic differences, but by variations in experiences with food or chemicals while they are growing up. The ranges of variation, and the adaptive “peaks” along the axes of such variation (in the case of the social insects, the actual worker and soldier castes), are finite and predictable (e.g. Oster and Wilson, 1979). I believe that we will eventually discover that exactly the same is true for the range and relative likelihoods of composites of learned behavior (or “learning phenotypes”) in humans.

Learning capabilities (and psychological mechanisms that utilize information acquired by learning) would not have evolved if they produced behaviors that were Culture and the Evolution of Social Learning 33 random with respect to biological adaptation. Organisms have evolved to learn in ways that maximize inclusive fitness; they have evolved to learn nothing else. This is not to deny that learning can result in maladaptive behavioral modifications; im- perfection is the bedfellow of unpredictable environments and novelties that favor learning capabilities in the first place. Hence, perhaps, our evolved abilities to ana- lyze and learn from mistakes and successes of others (Flinn and Alexander 1982). The degree of with which we view teachings of others-particularly non-relatives, or others whose interests do not jibe with our own-suggests that de- ception and manipulation are additional concerns. Learning is a type of phenotypic modification, one way that environmental conditions are utilized to adjust responses of the organism:

The alternative to cultural behavior is not “genetically transmitted” behavior: the envi- ronment always participates in ontogenesis [development of the phenotype], even when it [the environment] is invariable. Plasticity is the rule rather than the exception for all aspects of phenotypes, and imitation and other learning are not restricted to human culture (Flinn and Alexander 1982:384).

Cultural differences are due not to genetic differences but to a history of learned responses to different environmental conditions. To take a simple example, just as the white coat is advantageous for the Arctic hare in winter, so too knowl- edge of seal hunting is advantageous for the Eskimo. In both cases there is an evolved ability to respond in a flexible way to varying environmental conditions. In both cases the phenotype is adjusting itself in a way that is consistent with inclusive fitness maximization. In the case of seal hunting, psychological mechanisms using social transmission of information are involved. Learning capabilities involve specific propensities and constraints (e.g., Al- berts 1987; Bolles and Beecher 1988; Brown 1991; Carey and Gelman 1991; Craw- ford and Anderson 1989; Gallistel 1990; Geary 1995; Gelman 1990; J.L. Gould 1986; Gould and Marler 1987; Hinde 1987; Marler and Terrace 1984; McNaughton 1989; Oyama 1985; Pinker 1994; Shepard 1987, 1994; Siegler and Crowley 1994). Cognitive capacities, including complex features such as personality development, are influenced by genetic factors (e.g., Perusse et al. 1993; Plomin 1989, 1990; Plo- min, DeFries, and Fulker 1988; Rowe 1994; Starr 1992). The important questions here are whether learning propensities and constraints are adaptations produced by natural selection, and whether they influence transmission of cultural information (Boyd and Richerson 1985). Learning allows modification of behavior based on experience. Behavioral modifications are no less “biological” than physiological modifications; both in- volve chemical-neurological mechanisms, and both are products of evolution. There are several general “methods” or pathways by which adaptive learning can occur (Alexander 1990a; Bolles and Beecher 1988; Carey 1989; Cheney and Seyfarth 1990; Crawford and Anderson 1989; Gallistel 1990, 1995; Gelman 1990; J.L. Gould 1986; Heyes 1993, 1994; Johnston 1982a, 1982b; Mackintosh 1983; Marler and Terrace 1984; McDonald 1988a; Plotkin 1988; Roper 1983; Shettleworth 1984; Staddon 1983; Tomasello et al. 1993; Whiten and Ham 1992; Zentall and Galef 1988). The simplest method is behavior modification based on the “trial and ” 34 M. V. Flinn

of individual experience within specific domains (e.g., rats develop aversions to that result in nausea-Garcia 1974). More complex learning methods use information transfer from one individual to another via “imitation” (for more specific use of the term, see Heyes 1994; Galef 1988, 1996; Tomasello 1996). The ability to (1) observe behavior (of , etc.), (2) produce a mental image, and (3) reproduce the behavior (imitate or copy) can have tremendous advantages over trial and error (Heyes and Galef 1996). Social learning allows one to benefit from the experiences of others, and to provide co-res- ident offspring with a “head start.” Aversions to poisonous mushrooms can be de- veloped by following the example set by others rather than by direct experience. Problems with imitative learning arise if inappropriate (maladaptive) behaviors are acquired (Mayr 1963). For example, it would not suit nest parasitic species such as European to learn their mating songs from their hosts. Imitative learning re- quires learning “templates” or “innately tuned sensory systems” that may be time and/or situation restricted (e.g., : Alexander 1990a; Hirschfeld 1988; Sherman and Holmes 1985; songs: Nottebohm 1984, Petrinovich 1988; : Money 1987). Note that it is the presence of changing and unpredictable (Edelman 1987 uses the term “surprises”) environmental conditions (including so- cial interactions) that favors imitative learning. Imitative learning need not be “blind.” Evaluation of the relative success and failure of one’s peers may allow for more sophisticated behavioral modification. For example, in some species of , selective imitation of songs of males that are suc- cessful in attracting females can be more advantageous than random imitation of a singing male (Baker and Cunningham 1985; Payne 1982). Learning via selective imitation can lead to cumulative directional change if successful innovations are passed along to the next generation (e.g., Maestripieri 1995; Menzel, Davenport, and Rogers 1972; Tomasello, Gust, and Frost 1989). Intergenerational social learn- ing with cumulative modifications can result in “progressive” historical develop- ment of information.8 Because the current information pool is based on experiences of past generations, this type of learning involves historical constraints. Most defini- tions of “culture” involve learned information of this sort, and the behavior it pro- duces. A further modification of learning involves selective imitation of behavior based on an individual’s specific micro-environment and life-history stage. Abilities to “custom-fit” acquisition of information to an individual’s specific life-history cir- cumstances are advantageous if there are significant differences in individual strate- gies. What is best for one individual to learn at a certain time in life may be inappro- priate for another. You want to learn the right thing at the right time from the right (cf. Bandura 1977; Crawford and Anderson 1989; MacDonald 1988b; Piaget 1973). Specialized learning of this sort requires the ability to analyze the in-

‘Note that cultural “progess” (e.g., increasing social complexity, , and group size) is an incidental effect (Ingold 1986; Wright 1990). “Intentionality” of human reason is a product of evolution, not the driving force (Dennett 1983). Competition within a cultural environment, however, may be a zero-sum, “ queen” game in which “winners” must continually advance their tactics beyond their competitors’. Arms races are a good example. Culture and the Evolution of Social Learning 35 dividual’s position in current social-environmental conditions (perhaps involving -awareness and -see Alexander 1989; Cheney, Seyfarth, and Smuts 1986; Dennett 1983, 1995; Damasio 1994; de Waal 1989; Griffen 1984; Humphrey 1984; Tomasello et al. 1993). Such analysis probably requires extensive information storage () in order to have a basis for comparison and perhaps for delayed usage (e.g., young chimps observing an alpha male may store images of his behavior for imitation later in their ). Humans (and perhaps other hominoids, especially chimpanzees) have devel- oped complex forms of learning that involve behavior modification based on mental scenario building (Povinelli 1993, 1996). We use mental “games” to predict possi- ble outcomes of alternatives. Should I get Aunt Leila the red sweater or the blue? Which color would she prefer? Such decisions are based on “foresight” (i.e., pre- dicted outcomes) and “mind-reading” (i.e., comprehension of thought processes, and likely behavioral strategies) of others. Decision-making based on mental scenar- ios allows for experience without the cost. Such enhanced psychological mecha- nisms require expanded neural capacity to develop increased memory (providing a database for evaluation of alternative courses of action) and increased cognitive ca- pabilities (allowing for more extensive analysis, framing, categorization, etc.). Humans use a combination of learning methods in day-to-day living. We ob- tain information from direct observation and symbolic . We “think over” acquired information (consciously or unconsciously), and evaluate whether it is useful (e.g., Chibnik 1981). We modify our behavior accordingly. Unfortunately, quantitative models of cultural transmission do not yet include complex learning and information manipulation that exemplifies human culture and behavior. Several dual inheritance models incorporate simple learning biases or “guided variation” in- volving selective imitation of successful individuals (Boyd and Richerson 1985, 1988a, 1995; Cavalli-Sforza and Feldman 1981; Richerson and Boyd 1989a, 1989b; Rogers 1988a).9 Although shortcuts might economize on the costs of information processing (e.g., Tversky and Kahneman 1974), identification of specific algorithms is tenuous. Humans appear “smart”; we do not randomly imitate cultural traits, even from apparently successful role models. Nor are cultural traits employed randomly; individuals strategically use different behaviors to suit particular contexts. The complexity of social learning processes is the key issue separating evolu- tionary theories of culture. Evolved psychological mechanisms theories emphasize adaptive decision-making, whereas traditional and cultural selection theories focus on non-psychological aspects of information transmission such as structure of diffu- sion (vertical via parents, horizontal via peers, etc.). The issue boils down to whether learning “rules of thumb” are simple or sophisticated, and the importance

‘cf. Boyd and Richerson 1985 and Rogers 1988, who propose that blind imitation social learning abilities may be sufficient to account for cultural evolution if they produce adaptive behavior on average. The critical issue here is whether or not refinements of learning abilities and PMs improve batting averages (IFM behavior). The blind imitation player has a blindfold and no choice of bats, but theoretically hits a pitch and chances upon the base paths often enough to occasionally score. The evolved PMs player has refined skills, chooses a bat carefully, watches the pitches received by previous batters, spits, argues with the umpire, swings differently with runners on base, and so forth. 36 M. V. Flinn

of “other forces” besides evolutionary design of cognitive processes that affect cul- ture content. The ubiquity of apparently nonadaptive or maladaptive behaviors such as tat- toos, arbitrary food taboos, religious beliefs, celibacy, ethnic markers, dress style, and so forth, is interpreted by dual inheritance models as evidence that forces be- sides evolved psychological mechanisms may be influencing culture choice. Cul- tural traits that are maladaptive nonetheless become common by virtue of “cultural” processes such as society-level functions, conformity, or blind imitation, because human psychological mechanisms are not sophisticated enough to discriminate among cultural options. There are alternative explanations for the existence of non- adaptive and maladaptive behaviors, including historical lag, deception and manipu- lation by competitors (Hirshleifer 1987; Krebs and Dawkins 1984); experimenta- tion, selective conformity, and chance (see Alexander 1979a; Dawkins 1982:33-54 for discussion of “constraints on perfection”). Some of the apparent differences be- tween dual-inheritance and evolved psychological mechanisms theories are waning because of a trend in dual-inheritance theory to include more complex learning models (e.g., Boyd and Richerson 1988a, 1995). A main point separating evolutionary theories of culture is whether culture is partially independent of biological adaptation. Evolved psychological mechanisms theory emphasizes that learning capabilities are evolved aspects of the phenotype, and as such, have been designed by natural selection. The information bits in human minds are generated by processing of observations of the behaviors of others, and arc not viable replicators. Detailed descriptions of psychological mechanisms (and their ontogenetic development) are important research objectives. Dual-inheritance and mind-parasite the- ories propose that there are additional forces besides the evolutionary design of the hu- man mind that affect culture content, and that information bits in human minds should be considered distinct entities (replicators) with their own evolutionary interests.

Culture Is Partly Extrasomatic

“Culture is man’s extrasomatic means of adaptation” (White 1959:8).

Although most anthropological theories view culture as a “mental” phenomenon (Geertz 1957; Goodenough 1970; cf. Keesing 1974), some include behavior and “physical” products of human thought and action, such as tools, housing, clothing, speech, art, and musical instruments (e.g., Kroeber 1944; White 1949; cf. Cloak 1975). Here I do not wish to engage this definitional argument. The issue at hand is whether or not phenomena outside the body (i.e., “extrasomatic”) are subject to evo- lutionary adaptation. This question has been troublesome for biologists as well. Consider a spider’s web. Even though it is not a part of the spider’s body, it is undoubtedly a product of evolution. Web production evidently has been designed by natural selection and hence may be considered part of the spider’s phenotype (Dawk- ins 1982). Most webs are species-specific, and appear well-suited to a particular spe- cies’ niche. Bird , beaver dams, hives, and caddis- houses are similarly viewed as phenotypic adaptations. The difference between a hermit crab using learned Culture and the Evolution of Social Learning 37 behavior to acquire a shell and a snail using internal to build a shell is at the level of mechanism. Both are phenotypic adaptations produced by natural selection. But is there a fundamental difference between a bird’s nest and an Eskimo ig- loo? Or between a callus and a glove? The information and responsible for the construction of these flexible modifications of the phenotype reside within the central nervous system. In the case of the igloo and the glove, there are impor- tant “constraints” imposed by cultural history. Knowledge of how to construct an ig- loo or sew a glove, based on the past experiences of other humans, is a critical as- pect of human adaptation. (I discuss the importance of historical constraints in a following section). Here the point is that even though igloos and gloves are not part of the human body, this does not necessarily remove them from the realm of evolu- tionary adaptation. Natural selection can operate on bird nest or igloo design just as surely as it can operate on snail shell design. Heritable differences in nest or igloo building skills can affect survival and just as surely as morpho- logical differences in shells. This is the primary justification for considering some extrasomatic phenomena part of the phenotype (Dawkins 1982).‘O Whether or not the extrasomatic trait itself is subject to natural selection is an important source of controversy. Some dual-inheritance and cultural selection mod- els (e.g., Ball 1984; Cavalli-Sforza and Feldman 1981; Cloak 1975; Dawkins 1982; Dunnell 1989a; Durham 1982, 1991; Goodenough 1995; Mundinger 1980; Pulliam and Dunford 1980; Rindos 1985, 1986a, 1986b, 1988; Ruyle 1973) posit that cul- tural traits (or “memes”) are independent evolutionary entities or “replicators” (see the following section, “Culture Is Mental”). Social learning adds a new wrinkle to the extrasomatic phenotype. If bird nest types are partially or fully “borrowed” or imitated from individual to individual, then frequencies of different nest types depend not only on differential reproduction of individuals with different nest types, but also on whether nest types are differen- tially copied. The rate at which specific nest types are copied will depend on interaction among (1) features of the nest type, (2) distribution of nest types in the environment, (3) evolved psychological mechanisms, and (4) previously learned information (his- tory and individual mental development). The success (spread and persistence) of phenomena that are acquired by social learning depends, at least in part, upon men- tal decision-making processes, There is, however, an additional process independent of mental choice that may affect extrasomatic traits. Consider two variants of a sword, A and B. A is un-

“IThere are additional problems with defining phenotype, such as: Where does it end? Are footprints part of the phenotype? Are bee hives or other types of cooperative housing to be considered phenotypes? These are unresolved issues. It is also important to recognize that selection may design flexibility into the nest-phenotype; different conditions may warrant different responses. Hence the evolution of an “ sense” that can respond to variable environmental puzzles, as for example with different geometric configurations of branch structure within which the nest must be built. There are additional thorny unanswered issues. Is a farm part of the farmer’s phenotype? How is it different from the spider’s web or the hermit crab’s shell? If we allow for the farm to be part of the farmer’s phenotype, then are the and animals part of his or her phenotype? Where does the phenotype end? All of these examples are modifications of the environment engineered by the for its reproductive advantage. The effects of a hailstorm on a spider’s web and a farmer’s wheat crop are the same: Caloric intake is diminished and reproduction potentially reduced. 38 M. V. Flinn breakable in battle, and saves its owner from death. A is likely to become more com- mon than the alternative variant, B, whose unlucky possessor perished, for at least two reasons. First, people may recognize A as a superior weapon and copy it; this fits with “adaptive psychological mechanisms” selection of cultural traits. Second, by virtue of its longer life (as well as that of its owners), A is more likely to be observed and hence potentially copied. Tea cups may be substituted for swords with essentially the same result (unbreakable tea cups would have greater opportunity for observation and copying). The issue here is whether or not cultural traits have qualities that affect their distribution independent of choices made by human minds. If so, then we need a model of cultural evolution that includes such characteristics (e.g., Boyd and Richerson 1985; Dawkins 1989; Rindos 1986a, 1986b), and that can determine how important such “ex- trasomatic cultural selection” is relative to choice based on psychological mechanisms. The issue is beginning to be addressed by empirical studies (e.g., Aunger 1992).

Culture Is (Partly or Totally) Mental

Most definitions of human culture involve use of the human brain. Indeed, some definitions emphasize culture as a mental phenomenon, that is, information stored in the central nervous system (e.g., Geertz 1957; Goodenough 1970). Others (especially in archeology) focus on physical or social artifacts (Dunnell 1980; Leonard and Jones 1987; Rindos 1988). The issue here is not definitional, but whether or not information, and/or the mental processes used to analyze and transmit information (and hence modify behavior), are subject to evolutionary adaptation. Does the fact that culture is at least partly mental remove it from the realm of organic evolution? Is there a fundamental difference between mental phenomena (e.g., “ideas”) that are based on “information” and phenomena that are based on some physical form or structure? The information responsible for culture is processed and stored by physiologi- cal/chemical means. Culture is ultimately rooted in neurobiology. There is no basic physical difference between mechanisms of, for example, locomotion, and mecha- nisms of thought. Both involve chemical structures that are ontogenetic products of interaction between genes and environment. Both are products of evolution. At this basic level there are no grounds for treating mental phenomena differently than other aspects of life. The brain uses electrochemical structures for information stor- age; the liver uses chemical structures (e.g., glycogen) for storage. But there is, of course, an important difference between informational and most morphological structures that is of great significance for understanding culture. In- formation can be communicated. An can spread rapidly throughout a popula- tion, transmitted from one brain to another. Transmission and replication of ideas in- dependent of physical reproduction is the basis for traditional anthropological, cultural selection/mind parasite, and dual-inheritance theories of culture.”

“The primary differences between coevolutionary theories and traditional anthropological theories in this context are (1) traditional anthropological theories involve function or “meaning” at the level of the group rather than the cultural trait itself (e.g., Harris 1974), and (2) traditional anthropological theories do not specify the mechanisms by which cultural traits change in frequency (Murdock 1956, and Barth 1967 are notable exceptions). Culture and the Evolution of Social Learning 39

Cultural traits transmitted by social learning seem to belong to a distinct pro- cess, partially independent of genetic evolution. This is a central tenet of traditional anthropological theories of culture (e.g., Hallpike 1986; Spencer 1880; White 1949)-for reviews, see (Keesing 1974; Kroeber and Kluckhohn 1952). It has emerged in a more explicit form in cultural selection and dual-inheritance evolution- ary models of culture (e.g., Boyd and Richerson 1985; Campbell 1965; Cavalli- Sforza and Feldman 1981; Cloak 1975; Dawkins 1982, 1989; Goodenough 1995; Mundinger 1980; Richerson and Boyd 1978; Rindos 1985). These new theories pro- pose fundamental similarities between the process whereby genes are transmitted from generation to generation, and the process whereby cultural traits/information are transmitted from person to person, eventually passing down through history.‘* A new joke, for example, may be told and retold, replicating its way through a popula- tion, in a way analogous to the replication of genes or the spread of a . The crit- ical issue here is whether or not the mode of transmission and inheritance results in a distinct evolutionary system with its own emergent properties that are at least par- tially independent of “biological” underpinnings. provides clear descriptions of the idea that culture traits are subject to a separate evolutionary process. Less explicit models are more difficult to contrast. Dawkins (1982:290) hypothesizes that the basis for a cultural trait is a “meme.” defined as:

A unit of cultural inheritance, hypothesized as analogous to the particulate gene, and as naturally selected by virtue of its “phenotypic” consequences on its own survival and replication in the cultural environment.

Dawkins’ theory is based on the concept that memes are “replicators,” defined as “any entity in the universe of which copies are made” (Dawkins 1982:293). Dawkins requires that memes have a “definite structure, realized in whatever physi- cal medium the brain uses for storing information” (Dawkins 1982:109). So the mental chemical structure is equivalent to a gene (DNA), and its behavioral effects are equivalent to phenotypic effects (cf. Cloak 1975). However, no chemical or syn- aptic structures in the central nervous system have been found that correspond to

‘*Boyd and Richerson (1988) refer to this type of analysis of cultural evolution as “methodological ,” as opposed to “substantive Darwinism.” Methodological Darwinism is based on the possibility that “behavioral strategies are transmitted culturally instead of genetically .” Boyd and Richerson 1988:339). I view this position as analogous to stating that “arms are transmitted morphologically instead of genetically.” Their approach analyzes the transmission of cultural traits as analogs of genes. If natural selection produced the mechanisms by which behavioral strategies are transmitted culturally, then there can be no “instead of,” and the utility of “methodological Darwinism” is restricted to description, not explanation. Note that evolved PMs theory does not suggest that the medium of information transfer cannot have significant effects on culture; witness the dramatic changes accompanying the development of and television! The medium may even influence the success of certain types of cultural traits over others. For example, the success of science fiction and fantasy characters such as “Mutant Ninja Turtles” are probably dependent upon visual media. And ideas contained in the Bible may have benefited from having been stored and conveyed via written language. The effects of such transmission mechanisms would seem to be a useful addition to coevolutionary models. But I doubt our understanding of the cultural success of, say, the Ten Commandments, would be much improved. The nature of human social relationships and the specific historical context in which the commandments were developed and popularized seem much more significant determinants of their content and appeal than the “details of cultural transmission.” Clearly we cannot ignore in this regard either. 40 M. V. Flinn specific behaviors or cultural traits (e.g., Gazzaniga 1989, 1992; Pribram 1971; cf. Dawkins 1989). The physical composition of information storage and manipulation probably varies from individual to individual and changes throughout ontogeny (e.g., Black 1995). It is unlikely that there are structures in the brain that store cul- tural information in a way analogous to the DNA structure of genes. There are no evident brain chemicals/structures that are replicators (more precisely, no “germ- line” replicators). Indeed, Dawkins (1982: 112) notes several weaknesses in the meme/gene anal- ogy. First, “it is not clear that they [memes] occupy and compete for discrete ‘loci,’ or that they [memes] have identifiable ‘’.” Second, the meme “copying pro- cess is probably much less precise.” And third, “new ‘’ may be ‘directed’ rather than random.” He concludes, “These differences may prove sufficient to ren- der the analogy with genetic natural selection worthless or even positively mislead- ing.” I agree. In spite of this caveat, Dawkins (1982: 116) argues that cultural or “memic” evolution may go on quite independently of organic evolution: “A meme has its own opportunities for replication and its own phenotypic effects, and there is no reason why success in a meme should have any connection whatever with biological suc- cess.” I disagree. Dawkins’ meme theory is based on the premise that memes culturally evolve to maximize their survival. Like genes, memes are “selected” on the basis of their ef- fects: “If the phenotypic effect of a meme is a tune, the catchier it is the more likely it is to be copied” (Dawkins 1982: 110). But what is the basis for some tunes being “catchier” than others (and why are some ways of making stone points or clay pots copied more than others)? What are the mechanisms of cultural/psychological infor- mation selection, and where did they come from? This is a critical question that all models of cultural evolution must contend with. For cultural evolution to be an au- tonomous process, cultural selection must be independent of (past) organic evolution. The logic of the autonomous cultural system approach is tenuous if the mecha- nisms by which cultural traits increase or decrease in frequency are products of a history of natural selection on human abilities to utilize the accumulated body of in- formation that comprises culture. Selection of cultural traits by an individual may be independent of genetic transmission of that individual, but it can never be indepen- dent of the past history of natural selection that produced psychological mechanisms of cultural selection in the first place (Alexander 1979a; Flinn and Alexander 1982). Cognitive aptitudes underlying cultural selection-choice, imitation, learning, syn- thesis, teaching, comparison, foresight, , inference, and so forth-are capa- bilities that evolved by natural selection. Ultimately such mechanisms are based on neurological processes (e.g., Gazzaniga 1992). Daw,kins (1989: 194) likens social learning capabilities to primeval soup (meme: human brain:: DNA: primeval soup). This is true in the trivial sense that brains are a required medium for the transmission and replication of ideas. But the issue is whether or not minds develop psychological mechanisms designed by natu- ral selection to preferentially select some memes over others (and further, to use in- ference, i.e., past experience, current conditions, and mental scenarios of the future, Culture and the Evolution of Social Learning 41 to modify meme selection-see Staddon 1988). This is where the brain: primeval soup analogy breaks down. Primeval soup was not designed by natural selection for the function of DNA replication; the brain is. Meme theory suggests that cultural traits have evolved via cultural selection to “parasitize” human brains (Humphrey in Dawkins 1989:207; cf. Goodenough 1995; Goodenough and Dawkins 1994; Hull 1982; Sperber 1985). Such an epidemiologi- cal process is possible if cultural traits/memes are valid replicators, but it still would be best understood using knowledge of what human psychological mechanisms have evolved to do (Sperber 1991). From this “culture trait = mind parasite” per- spective we would study the capabilities of minds to be preferentially “parasitized’ by memes that are consistent with evolved mental properties. I find this way of thinking awkward. Cultural traits are replicators in the ,same limited sense as skin cells in a callus, glycogen in a liver, or cortisol in an adrenal gland. All involve flex- ible phenotypic responses and are not meaningful germ-line replicators. They are most appropriately considered as part of the individual’s phenotype. Other “cultural selectionists” (e.g., Cavall-Sforza and Feldman 1981; Dunnell 1980; Rindos 1986a; cf. early formulations by Boehm 1978; Campbell 1965; Cloak 1975; Ruyle 1973; Schwartz and Mead 1961; Simpson 1962; Wallace 1961) employ a logical argument similar to Dawkins’ in that they propose an independent process of cultural selection. Unlike Dawkins, however, they do not require that cultural traits have specific mental templates (i.e., transmissible replicators); instead, it is the cultural traits themselves that are analogous to genes. For example, specific variants of artistic design found on cooking pots may increase or decrease in frequency. This trait-evolution approach may be a useful descriptive tool for examining cultural change, but it is not a functional theory unless it incorporates selective mechanisms (e.g., mental choice) underlying changes in trait frequencies. The fundamental distinction among cultural selection, dual-inheritance, and evolved psychological mechanisms models is the of adaptation. In cultural se- lection theory, cultural traits/memes are the adaptive units that determine culture content by virtue of their own success or failure. In evolved psychological mecha- nisms theory, psychological mechanisms influence culture content (given the infor- mation provided by history and diffusion). Dual-inheritance theory allows for both. In effect there is an “arms race” between cultural traits and psychological mecha- nisms, with cultural traits that remain attractive to psychological mechanisms being saved, while psychological mechanisms evolve to select adaptive cultural traits. The crux of the issue is the extent to which evolved psychological mechanisms exist. Only if the central nervous system is a blank slate would culture content be deter- mined by a selective diffusion process independent of organic evolution.‘” Evolved psychological mechanisms theory begins with the evolutionary basis for cultural selection, that is, evolutionary design of human psychological mecha- nisms (Alexander 1979a, 1990a; 1990b; Tooby and Cosmides 1989). Culture in- volves mental processes that are products of evolutionary adaptation. Unfortunately

“Except for trivial effects of differential reproduction of trait carriers (Durham 1979). 42 M. V. Flinn this assumption does not tell us much about the specifics of human psychological mechanisms, but it seems a more viable starting point than apsychological and non- evolutionary theories have provided.

Culture Is Transmitted Via Arbitrary Symbols

If others possess information that is useful (i.e., results in adaptive decision making, e.g., when to corn, how to fashion a stone point, or whom to trust as an ally), then individuals can benefit greatly from communication/transmission of informa- tion (Krebs and Dawkins 1984; Petrinovich 1995; Pinker 1994). Human language allows for learning on a grand scale. Individuals can acquire information about events and concepts that are distant in time and space, rather than being limited to imitative learning based on direct observation. Complex scenarios (and “intentions” in the sense of Dennett, 1983) can be rapidly communicated and analyzed. Group decision-making can become much more complex, based on mutual discussion (Boehm 1978) and “mind-reading” (Cheney and Seyfarth 1990; Whiten 1991). The auditory and visual cues that humans use to communicate are commonly termed “symbols” because of the apparent arbitrariness of the connection between a specific and its “meaning.” This arbitrary quality of symbols could provide the basis for culture as an autonomous process (White 1949, 1959). However, if abilities to communicate using arbitrary symbols are products of natural selection (Burling 1986; Pinker and Bloom 1990), then both communication/linguistic skills and the content of information transfer should be designed to produce inclusive fitness-maxi- mizing behavior. Arbitrary symbols are particularly efficient means of information transfer for humans because of the need to communicate novel meanings/information (e.g., Wallmann 1992). Arbitrary symbols are used to represent information, but the spe- cific information itself is not arbitrary. Bird songs and cricket chirps are equally arbitrary in this sense; there is no logical connection between particular wavelengths of a cricket’s call and communication of, say, readiness to mate. The particular wavelength is an arbi- trary symbol, albeit highly constrained by phylogenetic history. Indeed, there is no evo- lutionary basis for symbols not to be arbitrary. The particular choice of symbols, however, should be influenced by functional considerations of time, error preven- tion, historical antecedents, and sensory and central nervous system capabilities. The general point here is that although the specific symbols used to represent infor- mation are partially arbitrary (given morphological and neurological constraints) and hence best understood as a diffusion process, this does not mean that the content of in- formation transmitted is arbitrary or independent of evolved psychological mechanisms.

Culture Has Emergent Properties

Information may develop emergent properties when it is shared and transmitted among individuals. For example, new ideas may be generated by synthetic processes when individuals “share” (communicate) information, and information “pools” can become cumulative over time (hence, knowledge can “advance” or build upon prior Culture and the Evolution of Social Learning 43 knowledge). Perhaps more importantly, the combined knowledge and behavior of indi- viduals can produce group phenomena such as shared “institutions” of religious belief, legal rules, and language. These properties of culture-shared, integrated, “meaningful” information-seem to give culture a life of its own, independent of the individual. Indeed, it is difficult to imagine how cultural “systems” could appear to “func- tion” so well if they are mere incidental by-products of individual behavior. Hence the hypothesis that culture functions to preserve itself, rather than serving the ex- plicit needs of individuals possessing it (e.g., Durkheim 1938; Hallpike 1986; Par- sons 1949; Radcliffe-Brown 1952). Or further, that human psychology is itself de- signed to produce “socializable” individuals that perpetuate their (e.g., Freud 1938). This society level functionalism parallels the group, species, and eco- system level functionalism prevalent in biology and ecology during the first half of the 20th century. Williams (1966; Williams and Williams 1957), Fisher (1958), Hamilton (1964), Lack (1966), and Lewontin (1970) challenged the concept of group-level functionalism in evolutionary theory, noting that natural selection is rel- atively weak at levels higher than the individual. The basic problem with explana- tions based on group- or cultural-level adaptation (e.g., Harris 1974:66) is that they require rapid rates of and replacement among isolated groups. In biology, most behavioral characteristics are currently interpreted as adaptations for individ- ual survival and reproduction, and not for group or species survival (e.g., Alcock 1994; cf. Wilson and Sober 1994).14 This dramatic shift in evolutionary theory and its ramifications for theories of social organization are beginning to their way through and other social sciencesI (Brumfiel 199255 1):

We must recognize that culturally based behavioral “systems” are the composite out- comes of negotiation between positioned social agents pursuing their goals under both ecological and social constraints.

Current evolutionary models in propose that culture may be understood as the cumulative outcome of individual actions, which ultimately are products of evolved psychological mechanisms (Cosmides and Tooby 1989:5 l-52):

The study of culture is the study of how different kinds of information from each individ- ual’s environment, especially from his or her social environment, can be expected to affect that individual’s behavior. The behavior elicited by this information reverberates through- out the individual’s , as information that other individuals may act on in turn. The ongoing cycle that results is the generation of culture. By directly regulating individ- ual learning and behavior, those psychological mechanisms that select and process infor- mation from the individual’s social environment govern the resulting cultural dynamics. Information, or culture, thus involves not only social interaction, but also an endless history of social interaction, filtered and analyzed at each step by psycho-

14Boyd and Richerson (1988) argue that cultural institutions such as and social rules may arise from a process of of cultural traits because it is difficult to develop a model in which is advantageous for individuals in large groups. However the Alexander (1987) hypothesis that indirect reciprocity and observed reciprocity are important cogs in human has yet to be incorporated into mathematical models, perhaps because of the enormous complexity this would create. 15George Peter Murdock (1972) criticized society-level functionalism as “anthropology’s mythology.” apparently without knowledge of these changes in evolutionary theory. 44 M.V. Flinn logical mechanisms that are themselves developed during ontogeny in response to the particular subset of information that each individual is exposed to. This ap- proach parallels studies of human motivation in cultural anthropology (e.g., D’An- drade and Strauss 1992). But culture content cannot be explained by reference to evolved psychological mechanisms alone. How could we predict and explain the directions and pace of cultural changes, given that all humans have roughly the same psychological mech- anisms? Environmental differences appear insufficient to account for cultural differ- ences without including some type of interaction with historical context. A complete theory of culture must include the effects of (1) social integration and shared infor- mation, “reverberation throughout the individual’s social group”; (2) history, “the ongoing cycle”; (3) individual psychological and informational development (on- togeny); (4) the non-cultural environment (e.g., flora and fauna, geography, demog- raphy); (5) evolved psychological mechanisms that influence culture trait choice; and (6) chance or accidental events. There are two critical problems for such a general theory. First, it is not clear whether there are emergent properties in the history-psychology-genetics-history- psychology-genetics . . . cycle (Boyd and Richerson 1985; White 1949; cf. Alex- ander 1979a:69). And second, it is not clear whether the pool of information (cul- ture) can develop (over time-history) into something that “overtakes” the psycho- logical mechanisms (Alexander 1979a:80; Irons 1979a:38, Irons 1983:171). If the information that is available during ontogenetic development of psychological mechanisms is sufficiently different from information that such mechanisms evolved to use for appropriate development, then maladaptive psychological devel- opment (and subsequent cultural evolution) is likely to occur. Social competition among individuals involving exploitation or manipulation may also be an important source of maladaptive behavior (Alexander 1987; Chagnon 1982; Fredlund 1985; Harpending, Rogers, and Draper 1987). These two problems may be partially resolved by empirical studies of the rela- tionship between psychological mechanisms and behavior in contemporary environ- ments (e.g., Daly and Wilson 1988, 1995). Comparative studies of from different cultures are likely to be particularly useful for teasing apart the dynamic between psychological mechanisms and the cultural information pool (Belsky, Steinberg, and Draper 1991; Chisholm 1996; Worthman 1992). We need to better understand how psychological mechanisms are affected by informa- tion acquired during ontogeny. An additional problem that the psyche must contend with is the ability to juggle input from multiple psychological mechanisms, and to endure short-term pain for perceived long-term benefits (e.g., finishing a Ph.D. dissertation). Such abilities suggest the existence of a hierarchical (executive) decision structure. Complex judg- ments are required when several conflicting options co-exist, for example, to get ripe fruit you must first climb a dangerous tree, or, if you eat all the cookies, you might be reprimanded by mom for not sharing. There is no clear uniform evolutionary model for psychological mechanisms. It is humbling to remember that they are constructs or heuristics that may not have Culture and the Evolution of Social Learning 45 precise discoverable physical neurological forms. Some might be usefully distin- guished as specific, others more general. Some might be highly sensitive to minor environmental differences, others more robust. Some might be more likely to result in maladaptive responses to cultural novelty, others less so. All are hypothesized to be (ultimately) products of organic evolution. The issue of whether evolved psycho- logical mechanisms fall behind cultural evolution depends on understanding what such mechanisms are, what selective pressures created them, and what the pace and directions of cultural change are.

Culture Is Historical

Social learning allows for a “progressive” or “ratchet-effect” historical development of information: culture is cumulative.” But each step of cultural evolution, each bit of change in the information pool, must be generated and processed by an individual psyche. The “early” theories of cultural evolution based on a rule of unfolding (e.g., Morgan 1877; Tylor 1871) were wrong because they attributed design to history; later theories repeated the mistake (e.g., White 1949, 1959; see critiques by Ingold 1986; Johnson and Earle 1987). A historical theory of culture without psychology is as incomplete as is a psychological theory of culture without history. History constrains and provokes future directions of both organic and cultural evolution by providing materials upon which subsequent developments must build. It is useful to consider cultural change (originating with individual mental change) as being constrained by adaptive “informational landscapes” analogous to the Se- wall Wright (1931) concept in . Evolutionary pathways are “channeled” by historical constraints, not just evolved phenotypic mechanisms (Mayr 1988: 108). Dawkins (1982:38-39) provides an insightful description of his- torical constraints on evolution:

The jet engine superseded the propeller engine because, for most purposes, it was supe- rior. The designers of the first jet engine started with a clean drawing board. Imagine what they would have produced if they had been constrained to “evolve” the first jet engine from an existing propeller engine, changing one component at a time, nut by nut, screw by screw, rivet by rivet. A jet engine so assembled would be a weird con- traption indeed. It is hard to imagine that an aeroplane designed in that evolutionary way would ever get off the ground. Yet in order to complete the biological analogy we have to add another constraint. Not only must the end product get off the ground; so must every intermediate along the way, and each intermediate must be superior to its predecessor. When looked at in this light, far from expecting animals to be perfect we may wonder that anything about them works at all. A further constraint on organic evolution is imposed by the of mu- tation. Changes in the propeller engine are produced by a process analogous to a blindfolded group of welders and drillers and blasters, or the Shakespearean chimp on a typewriter. Cultural evolution may be a less chaotic process, because our minds filter out some of the less successful /mental mutations before we present

“In some respects this is similar to the problem of the development of species with larger, more complex soma over evolutionary time. 46 M.V.Flii them to others (who might reject them anyway), and because we appear to have the ability to direct our thoughts toward resolving specific problems (within limitations imposed by the information pool provided by history: imaginations are not unlimited). Although information can be transmitted intact and unchanged over periods of time, it also can change rapidly, producing novel challenges for the human psyche. Changing information favors flexibility in the information processing system. For example, canalized symbol-meaning relationships are unlikely because language changes so rapidly (Chomsky 1973; Pinker 1994). Social competitors can constantly update tactics; mental “arms races” of various sorts can develop. In spite of its po- tential advantages, however, learning is risky (Alcock 1994; Williams 1966). Novel environments create opportunities for acquiring the “wrong” information for behav- ioral development (Alexander 1979a; Barkow 1989b; Irons 1983). Some , for example, readily learn to chase cars. Culture appears to be an especially rapid source of change that may alter human environments too quickly for genes underlying psy- chological learning mechanisms to keep pace. Hence the human psyche, having evolved in hunter-gatherer cultural conditions, may not be adapted to current cir- cumstances (Symons 1979:35; cf. MacDonald 1988a). This “ psyche” or “Stone-Age mind” approach posits that human psychological mechanisms are rela- tively specific and inflexible; we have difficulty adapting appropriately to novel en- vironmental conditions generated by cultural change. Alternatively, if some evolved psychological mechanisms are more “domain- general,” or if there are hierarchical decision-making processes that can modify re- sponses from domain-specific modules, then modem environments are more likely to be within the range of appropriate response (Alexander 1990a; Daly and Wilson 1995): For example, if humans evolved psychological mechanisms that specify: “avoid running mastodons,” then such mechanisms would not prevent us from step- ping in front of speeding automobiles. On the other hand, if we evolved to “avoid large, fast-moving objects,” then both Mastodons and Mack trucks would be avoided by the same psychological mechanism (see Turke 1990). An even broader range of adaptive responses might be included under “avoid objects that your elder caretakers (parents) treat as dangerous” (see MacDonald 1996). Such a mechanism would be subject to developmental modification by social learning, and perhaps be capable of contending with extreme novelty (e.g., “avoid drug dealers”), although there are likely to be time lags before sufficient experience occurred for recognition of the novelty as dangerous. For example, consider tobacco, cocaine, and French pastries. All evidently appeal to evolved psychological mechanisms, via altering brain chemistries in ways that some humans find pleasurable. Recognition of such novelties as dangers occurred only after considerable exposure and long-term obser- vation of their effects. Dangers that are consistent and ubiquitous throughout evolu- tion are more likely to produce canalized responses, especially if they are suffi- ciently deadly so as to not allow for learning by experience (e.g., poisonous snakes-Alexander 1990a; Mineka and Cook 1988). The development of new cultural traits may present new opportunities and challenges, sometimes resulting in further cultural change in directions that would not be predictable without knowledge of the historical context. For example, female Culture and the Evolution of Social Learning 47 preferential among high-status groups such as the Rajput Brahmins of Northern India (Dickemann 1981; cf. Kitcher 1985) may have developed over time into a status marker. Initially, occasional female infanticide might have benefited high-status under certain circumstances, such as close birth-spacing. Over time, families with fewer daughters may have been identified with high status. Such status markers may have fostered social arms races that eventually resulted in the extreme of complete female infanticide among the highest status groups. Cultural practices that restrict female sexuality (claustration, veiling, genital mutilation, chaperones, honor doctrines, etc.-see Dickemann 1981) may have devel- oped in similar fashion, with higher status families continually raising the level of restriction in order to distinguish themselves from lower status groups. Hence un- derstanding cultural behavior requires knowledge of historical precedents leading to the current context (Alexander 1979a; Durham 199 1).

Why Is Complex Culture Unique to the Human Species?

Some recent theories of the evolution of human intelligence focus upon social com- petition as a selective force (Alexander 1971, 1989, Brothers 1990; Byrne and Whit- ten 1988; Humphrey 1976, 1984). Once hominids became the “ecologically dominant” species, they were their own most potent selective pressure. Humans uniquely evolved sophisticated brains because humans were the only species to become “their own principal hostile force of nature” (Alexander 1989:469) via inter- and intra- group competition. Increasing intellectual and linguistic capacities were favored because such skills allowed individuals to better anticipate and manipulate social interactions with other increasingly intelligent humans. This “runaway” produced ever greater cerebral capabilities until the reproductive advan- tage of above-average social skills/intelligence diminished (perhaps with the devel- opment of large social groups-see Alexander 1989). This “social competition” hypothesis has important ramifications for under- standing human psychological mechanisms. It suggests that the human psyche is designed primarily to contend with social relationships, whereas the physical (non- social) environment is relatively unimportant. Most differential reproduction in re- gard to brain evolution was a consequence of interactions with conspecifics, not with food and climate. The primary mental chess game was with other intelligent competitors, not with fruits, tools, prey, or snow (although enhanced intelligence is surely useful in dealing with such hostile forces as we11).17Human - ships are complex. Predicting future moves of a social competitor-cooperator, and appropriate countermoves, amplified by multiple relationships, shifting coalitions,

“This might explain the apparent lack of significant populational differences in human intelligence: The uniquely common selective force that all humans contend with is competition with other humans, regardless of physical environment. Interpopulational further eliminates potential differences among populations. 48 M. V. Flinn and deception, make social success a difficult undertaking (Alexander 1987, 199Ob; Axelrod and Hamilton 1981; Dennett 1983; de Waal 1982). Indeed, the potential variety of human social puzzles is apparently infinite; no two social situations are precisely identical, nor are any two individuals ever in ex- actly the same social environment. Moreover, social relationships can change rap- idly, requiring quick modification of strategy. This unpredictable, dynamic social hodgepodge would seem to favor flexible, “open,” “domain-general,” or “execu- tive” psychological mechanisms highly dependent upon social learning and capable of integrating information processed by more restricted, “domain-specific” mecha- nisms (e.g., Lopreato 1984; Meltzoff 1995; Mueller, in press). These complex cog- nitive processes would be more capable of contending with (and producing) novel- ties created by cultural change and culture- and individual-specific differences. Unfortunately such chameleonic psychological mechanisms would be nightmares to document empirically, evidenced perhaps by our meager understanding of the “black box” (but see, e.g., Robin and Holyoak 1995). The dramatic expansion of the neo-cortex during is consistent with the hypothesis that social fi- nesse is a particularly important human skill.

METHODOLOGIES FOR RECOGNIZING EVOLUTIONARY DESIGN

Differences in methodological approaches accompany semantic and conceptual dif- ferences among evolutionary theories of culture. In general this has resulted in a powerful diversity of empirical research (Hinde 1991), but it has not been without acrimony and apparent misunderstanding (cf. Buss 1995; Reeve and Sherman 1993; Symons 1992). Early evolutionary studies examined available ethnographic literature to test for associations between general patterns of human behavior and major theoretical concepts such as , reciprocity, and mate competition (e.g., Alexander 1974, 1977; Daly and Wilson 1983; Hawkes 1983; Lancaster and Lancaster 1983; Van den Berghe 1979). The resolution of apparent anomalies-such as cousin mar- riage asymmetries, , infanticide, and the mother’s brother phenomenon- were key objectives. The success of this preliminary research, and the large number of new hypotheses generated, sparked interest among anthropologists and other be- havioral . Previously collected ethnographic data were reassessed to deter- mine whether individual variation in behavior corresponded to micro-environmental differences, such as economic and social status (e.g., Chagnon and Irons 1979). More recently, a number of ethnographic studies have been conducted that were designed specifically to test evolutionary hypotheses (see reviews in Betzig, Borgerhoff Mulder, and Turke 1988; Cronk 1991; Smith and Winterhalder 1992). Characteristic of these new evolutionary studies is the use of objective, quantitative methods from a range of disciplines such as endocrinology, developmental psychol- ogy, demography, , and microeconomics, in addition to more traditional ethnographic methods. Such research techniques are beginning to provide informa- tion on possible proximate mechanisms that underlie cultural information and behavior. Culture and the Evolution of Social Learning 49

The assimilation of modern evolutionary theory into psychology resulted in a somewhat different and complementary methodological approach to that developed in anthropology. Beginning with a sophisticated knowledge of human cognitive pro- cesses, evolutionary psychology was more focused on identifying psychological mechanisms, and less concerned with environmental variation or demographic data testing whether behaviors are currently associated with fitness. The “species-typical architecture of the human mind” was postulated to be composed of distinct “do- main-specific” modules (cf. Fodor 1983) that are ubiquitous (Tooby and Cosmides 1992:38):

Empirically, of course, the fact that any given page out of Gray S describes in precise anatomical detail individual humans from around the world demonstrates the pronounced monomorphism present in complex human physiological adaptations. Although we cannot yet directly “see” psychological adaptations (except as described neuroanatomically), no less could be true of them. is everywhere the same. The methodology used to identify psychological adaptations is a combination of (I) documenting universality (Brown 1991) and (2) fit with the logic of apparent advantage in the “environment of evolutionary adaptedness” (Bowlby 1969), postu- lated to be a environment that dominated most of human evolutionary his- tory (Cosmides, Tooby, and Barkow 19925):

. the most reasonable default assumption is that the interesting, complex functional design features of the human mind evolved in response to the demands of a hunting and gathering way of life. . . We cannot rely on intuitions honed by our everyday experi- ences in the modem world . . behavior generated by mechanisms that are adaptations to an ancient way of life will not necessarily be adaptive in the modem world. Thus our concern is with adaptations- mechanisms that evolved by natural selection-and not with modem day adaptiveness. . For example, male/female differences in sexual proceptivity are consistent with expectations based on parental expenditure (Buss 1994; Symons 1979), regardless of effects of novelties such as modem birth control. From an anthropological per- spective, these psychological studies have been limited in cross-cultural scope; for example, the widespread phenomenon of cousin marriage rules in non-industrial so- cieties, a unique and important aspect of human mating systems, is not included in most studies of (e.g., Buss 1989; cf. Brown 1991). Empirical proce- dures used to document “Stone Age” psychological adaptations involve a combina- tion of traditional psychological methods and evolutionary logic. The mind is envi- sioned as a “jukebox” or “Swiss army knife” of distinct information processing modules that are requested and played (“evoked”) for specific needs/tasks. The em- pirical job is to determine which songs are included in this -human set of 45s. and how the tunes are played. Cultural variation, ontogenetic contingencies, hierar- chical integration (networking and comparison) of information from task-specific modules, and context specificity are not yet primary foci. A complementary approach (“human behavioral ecology” or “Darwinian an- thropology”) posits that characteristics of the “environment of evolutionary adapt- edness” such as foraging subsistence and group composition (see Foley 1992; Kelly 1995; Mithen 1990, 1994) are less monomorphic than assumed by the evolutionary 50 M. V. Flinn psychology paradigm (see also Nesse and Williams 1994:139). Both evolutionary psychology and human behavioral ecology posit that the human social environment is a critical selective pressure involved in the evolution of human psychology (Alex- ander 1989; Cosmides and Tooby 1992; Humphries 1984), but the behavioral ecol- ogy approach emphasizes the variable and novel aspects of such social environ- ments. More attention is given to developmental contingencies and hierarchical (executive) decision processes or motivational structures (Alexander 1990a; Mac- Donald 1991; Worthman 1992). This model suggests that behavioral variability in contemporary social environments is useful for analysis of human psychological mechanisms (e.g., Daly and Wilson 1995). Subsistence, mate choice, , kin- ship, warfare, and other human behaviors are variable because psychological mech- anisms are adapted to respond to environmental differences (Alexander 1974; Irons 1979a, 1983; Standen and Foley 1989; Winterhalder and Smith 1992). Methodologies used for identification of psychological mechanisms in human behavioral ecology includes examination of associations between behavior, fitness, and environmental situation, parallel to methodologies used in non-human behav- ioral ecology (Hinde 1991; Reeve and Sherman 1993). For example, reproductive success (number of offspring) is associated with family wealth among the Yomut Turkmen (Irons 1979c), hunting success among the Ache (Hill and Kaplan 1985; see also Hill and Hurtado 1996), land ownership in a rural Caribbean village (Flinn 1986), wealth and status among the Mukogodo (Cronk 1989a, 1989b), headmanship among the Yanomamo (Chagnon 1979), and political power cross-culturally (Betzig 1986). Similarly, social status is associated with mating success among French Ca- nadians (Perusse 1993). Unspecified and unknown (but presumed evolved) psycho- logical mechanisms for economic and social striving and use of resources and power to enhance mating opportunities are assumed to underlie such behavior. The lack of association between status and reproduction in modern high-technology (Vining 1986) is attributed to environmental novelties such as birth control and pro- longed education (e.g., Flinn 1987; Irons 1983), and helps flesh out the design of hu- man motivations. The use of social learning to alter behavior in response to ob- served maladaptive effects suggests that the management of conflicting motivations may be flexible. One generation’s mistakes may serve as lessons for the next, beg- ging the question: What are the evolutionary designs of such “corrective” psycho- logical mechanisms that modify decisions based upon processed information from lower-level domain-specific modules? Both research paradigms (evolutionary psychology and human behavioral ecology) have proven useful for testing evolutionary hypotheses, and suggest ave- nues for more detailed analysis of psychological mechanisms, including physiologi- cal processes (e.g., Buss et al. 1992; Flinn and England 1995, in press).”

“Some additional methods used to evaluate evolutionary hypotheses include general observations from ethnographic literature (e.g., nepotism, avoidance, infanticide: Alexander 1974, 1979a; Dickemann 1979), cross-cultural correlations (e.g., mating systems: Betzig 1986; Flinn and Low 1986) demographic measurements (e.g., sex ratios: Boone 1986; Chagnon, Flinn, and Melancon 1979; Voland 1984). measurements of reproductive success (e.g., Borgerhoff Mulder 1988; Chagnon 1979; Hill and Hurtado 1996). measurements of attitudes (e.g. Buss 1989), measurements of acquisition (e.g., Culture and the Evolution of Social Learning 51

Empirical Evidence for Evolutionary Design: of Mind and Adaptive Patterns of Human Behavior

The evolved psychological mechanisms model would be falsified by lack of evi- dence for task-specific learning mechanisms that reflect evolutionary design. If the human mind is a product of organic evolution, then the psychological mechanisms by which it operates should result in adaptive behavior in environments similar to those of human evolutionary history. Evolutionary psychology and behavioral ecol- ogy models of culture generate specific predictions about (1) adaptiveness of behav- ior, (2) ontogenetic constraints and environmental contingencies, and (3) neurological and/or other physiological “wetware” (Table 2). A list of possible products of evolved psychological mechanisms is presented in Table 3. These postulated behavioral effects of psychological mechanisms use so- cial learning in ways that (hypothetically) allowed for more adaptive behavioral re- sponses than if social learning were not involved. This suggests a more active role for human psychology in cultural evolution than “blank slate” and “cultural selec- tion” models, but does not imply a direct genetic of culture. Know- ledge of the genetics, developmental pathways and contingencies, and individual environmental and historical contexts for expression of these hypothetical adaptive products of evolved psychological mechanisms is limited. Language abilities are probably the best understood (e.g., Neville 1992; Pinker 1994). Most postulated behavioral outcomes of evolved psychological mechanisms are consistent with results of traditional ethnographic research, such as the ubiquity of kinship, competition for mates and resources, language, differences, and (e.g., Murdock 1949). Evolutionary theory provides new ex- planations for these common features (Brown 1991) and their variation among hu- man cultures. However, large gaps remain between general theory and the identifi- cation of specific mechanisms. Indeed, defining “psychological mechanisms” is problematic. Evolutionary- Tinbergen (1963) “function’‘-explanations of culture exceed knowledge of how the evolved mind works. Neurological processes are many steps removed from heu- ristics such as postulated psychological mechanisms for mate choice, reciprocity, or language. Understanding integration of different levels of processed information is an important challenge for evolved psychological mechanisms theories (Barkow 1989b; Lumsden and Wilson 198 1).

Kaplan and Hill 1985, 1992), measurements of behavior of individuals (e.g., Flinn 1988b; Hewlett 1988). measurements of psychological parameters (e.g., reciprocity: Cosmides and Tooby 1989). This scientific methodological emphasis of evolutionary approaches in anthropology contributes to disagreements with humanistic approaches in anthropology. For example, a reading of Oedipus Rex may provide more useful information about the human family than 10,000 quantitative behavioral observations. Both approaches, however, involve a key element of the scientific approach: communication among observers. You and I may interpret Shakespeare somewhat differently, but were there no common understanding between us then I doubt it would be so popular. The fact that “plays” and other forms of aesthetic communication so commonly utilize themes of , hate, greed, selfishness, , honesty. deception, trust, and so forth, is consistent with an evolutionary approach, as is cultural and individual specificity of interpretation. Methodological differences between humanistic and evolutionary anthropology involve issues of quality control and techniques, not subject matter. Appropriate methods for testing evolutionary hypotheses from “text” have yet to be developed (but see Nesse 1995). 52 M. I? Flinn

Table 2. Behavior, Brain Design, and Cultural Variability

Theory Behavior Brain design Cultural variability

Traditional Maintain society None, Random: historical and social science provide meaning irrelevant idealogical constraints

Cultural selection Meme product for memes ? diffusion of memes

Dual Mix of cultural and Simple? Medium for Diffusion; ecological and inheritance biological adaptation transmission of traits historical constraints

Separate “domain-specific” Evolutionary Adaptive- modules integrated by Noise, novelty, some psychology “monomorphic” EEA hierarchical mechanisms ecological constraints

Domain specific and general Behavioral Adaptive- modules integrated by Micro-ecological and ecology variable EEA hierarchical mechanisms historical constraints

Vision, for example, is vital for acquisition of cultural information. Photons strike cells in the retina (transduction), eventually stimulating site-specific modules (orientation columns) in the visual cortex. This information may be used in “higher level” processes (e.g., Gray et al. 1989), such as pattern recognition with fa- cial identity templates. Skipping intermediate steps, this processed information may be used during ontogeny in “even higher level” processes, such as attachment emo- tions, and involve neuroendocrinological systems such as response. The end results-again skipping many steps-include “cultural” activities such as kin relationships, inbreeding avoidance, and so forth. Hearing is also vital for acquisition of cultural information. Sound vi- brate the tympanum, are transduced via the cochlea, eventually stimulating the pri- mary auditory cortex. During development, some specific sounds become associ- ated with specific meanings (e.g., Grossberg and Stone 1986). For example, humans growing up in American culture learn to associate the phoneme “ma” with “mother” (i.e., adult female that takes care of me, etc.). Surely such linguistic skills are prod- ucts of evolution (Dunbar 1993; Pinker 1994; Wallace 1989), and involve connec- tions with visually acquired information (e.g., what ma looks like). Postulated “domain-specific” and “domain-general” psychological mecha- nisms-such as recognition of mother or acquisition of language-are heuristics for function, not specific modules of or chemo-electric activity that are identi- cal in each human. “Cultural” phenomena involve information processing by nu- merous neurological domains that appear hierarchical, fuzzy, networked, mobile, ontogenetically plastic, and perhaps ephemeral and individual-specific (D’Andrade 1995; Sperber 1996). “Epigenetic rules” (Alexander 1990a; Lumsden and Wilson 1981) and “Darwinian algorithms” (Cosmides and Tooby 1989) are not likely writ- ten in genetic neuronal stone. Domain-specific “modules” may occupy specific re- gions of the brain (albeit with minor variations among individuals), but this does not indicate neuronal pre-wiring. Localized modules may be products of developmental processes of selection of operating units that produce “results” judged effective by Culture and the Evolution of Social Learning 53

Table 3. Human Behaviors and Abilities Postulated to Use Adaptive Social Learning Mechanisms

Foraging decisions (Kaplan and Hill 1992; Smith 1991) Recognition of kin (Alexander 1990a) Recognition of individuals (Colgan 1983) Avoidance of inbreeding (Aberle et al. 1961: Alexander 1979a; 1993) Maternal attachment (Bowlby 1969; Chisholm 1996; Petrovich and Gewirtz 1985) Paternal attachment (Hames 1988; Lamb et al. 1987) Paternity recognition (Daly and Wilson 1988; Flinn 1988b) attachment (Turke 1988; Weisner and Gallimore 1977) Sibling rivalry (Daly and Wilson 1988) Kin assistance (Chagnon and Bugos 1979) (Nesse 1990) Variation in kinship behavior (e.g., maternal/paternal biases: Flinn I98 I ; Irons 1983) -offspring conflict (Daly and Wilson 1988) Mate competition (Chagnon 1988; Daly and Wilson 1988; Flinn and Low 1986; Irons 1983) Mate guarding (Daly and Wilson 1983: Dickemann 1981; Flinn 1988a) Mate bonding (Belsky et al., 1991; Chisholm 1993) Mate choice (Borgerhoff Mulder 1988; Buss 1989, 1994: Symons 1979) Sex/gender recognition and attraction (Money 1987; Symons 1979) Sexual (Buss 1994; Daly, Wilson, and Weghorst 1980: Hrdy 1981) Resource aquisition and competition (Flinn 1986; Hill and Kaplan 1989a, 1989b; Irons 1979b) Status aquisition and competition (Chagnon 1979; Betzig 1986; Irons 1979b) Verbal and non-verbal communicaton (Pinker 1994; Pinker and Bloom 1990) Mathematical reasoning (Geary 1995) (Alexander 1987; Eibl-Eibesfeldt 1989; McDonald, in press) Simple reciprocity (Axelrod and Hamilton 1981; Cosmides 1989; Cosmides and Tooby 1995) Complex reciprocity, including political and economic coalitions, group games, , and moral systems (Alexander 1987; Chagnon 1974; Cosmides and Tooby 1992) Religious beliefs (Boyer 1994; Reynolds and Tanner 1995) Technological and artistic tool use (Reynolds and Tanner 1995; Wynn 1989) Appreciation of aethetics, art and music (Alexander 1990b; Nesse 1995) Age-specific learning aptitudes (Piaget 1973; Pinker 1994; Smith 1987) Consciousness, scenario-building, intentionality, and foresight (Alexander 1989: Dennett 1983; Humphries 1986)

higher level executive management modules. Loss of specific function associated with localized brain damage suggests regional modularity; however, the ability to recuperate lost functions, particularly if they occur early in development, suggests that such modularity may be established or modified by developmental processes and not by predetermined localized neural modules. This apparent lack of direct one-to-one associations between neural location and psychological function makes mapping the “architecture of the evolved mind” a more difficult enterprise.

SUMMARY AND CONCLUDING REMARKS

Culture deserves scientific explanation because, like other potential adaptations, it has complex design (Dawkins 1986; Williams 1966). Humans live in a social mileau of politics, kinship, residence, language, art, technology, , economics, and so forth, that has identifiable patterns. The evolutionary approach advocated here views culture as an adaptive part of individual human phenotypes. Information underlying culture is acquired primarily 54 M. V. Flinn

by social learning, which is stored and analyzed in the human central nervous sys- tem. The mechanisms that create, acquire, retain, and analyze socially transmitted information are products of human evolutionary history, and hence are predicted to be designed to produce adaptive behavior. Because an individual’s particular situa- tion in society is dynamic and unpredictable (and has been so throughout human evolutionary history), psychological mechanisms are posited to be highly flexible and able to respond appropriately to a wide array of novel challenges, albeit fre- quently requiring “learning from your mistakes.” Information underlying culture is individual-specific, perhaps even time-spe- cific, but shared sufficiently to allow for varying degrees of communication and mu- tual comprehension. The collective human information pool changes over time in response to complex interactions among environmental, historical, demographic, and psychological variables, muddled further by competitive and cooperative striv- ings of individual humans. Culture is not a simple phenomenon. There is, however, one consistent force acting on the information pool in human minds: the generation and choice of cultural traits (information usage) by evolved psychological mecha- nisms that use social learning (Alexander 1990a; Boyd and Richerson 1985; Tooby and Cosmides 1992). Understanding the evolved functions of such cognitive pro- cesses and their ontogeny in environmental and historical context is critical to a gen- eral theory of culture.

REFERENCES

Aberle, D.F., Broffenbrenner, U., Hess, E.H., Miller, D.R., and Schneider, D.M. The taboo and the mating patterns of animals. 65:253-265, 1963. Alberts, J.R. Early learning and ontogenetic adaptation. In Perinatal Development: A Psychobiological Perspective, N. Krasnegor, E M. Blass, M.A. Hofer, and W. Smotherman (Eds.). Orlando, FL: Academic Press, 1987. Alcock, .I. Animal behavior (fifth edition), Sunderland, MA: Sinauer, 1994. Alexander, R.D. The search for an evolutionary of man. Proceedings of the Royal Society oj Victoria 84:99-120, 197 1. Alexander, R.D. The evolution of . Annual Review of Ecology and 5:325-383, 1974. Alexander, R.D. Natural selection and the analysis of human sociality. In Changing Scenes in the Natural : 1776-1976, C.E. Goulden (Ed.). Philadelphia Academy of Natural Sciences Special Publication 12, 1977, pp. 283-337. Alexander, R.D. Danvinism and Human Aflairs, Seattle: University of Washington Press, 1979a. Alexander, R.D. Evolution and culture. In and Human Social Behavior, N. Chag- non and W. Irons (Eds.). N. Scituate, MA: Duxbury Press, 1979b. pp. 59-78. Alexander, R.D. Evolution, culture, and human behavior: some general considerations. In Natural Selec- tion and Social Behavior. R. Alexander and D. Tinkle (Eds.). New York: Chiron Press, 1981, pp. 509-520. Alexander, R.D. The Biology ofMoral Systems, Hawthorne, NY: Aldine Press, 1987. Alexander, R.D. Evolution of the human psyche. In The Human Revolution, P. Mellars and C. Stringer (Eds.). Chicago: University of Chicago Press, 1989, pp. 455-513. Alexander, R.D. Epigenetic rules and Darwinian algorithms: the adaptive study of learnlng and develop- ment. and Sociobiology 1 l(3) l-63, 1990a. Culture and the Evolution of Social Learning 55

Alexander, R.D. How Did Humans Evolve? Special publication #I, Ann Arbor: Museum of , University of Michigan, 1990b. Aunger, R. The nutritional consequences of rejecting food in the Ituri forest of Zaire. 20:263-291, 1992. Axelrod, R., and Hamilton, W.D. The evolution of cooperation. Science 211: 1390-1396, 1981. Badcock, CR. Evolution and Individual Behavior. Oxford: Basil Blackwell, 1991. Bailey, R.C., and Peacock, N. Efe Pygmies of Northeast Zaire: subsistence strategies in the Ituri Forest. In Coping with Food Supply, I. deGarine and GA. Harrison (Eds.). Oxford: , 1988, pp. 88-l 17. Baker, M.C., and Cunningham, M.A. The biology of bird song dialects. Behavioral and Brain Sciences 8:85-133, 1985. Ball, J.A. Memes as replicators. Ethology and Sociobiology 5: 145-161, 1984. Bandura, A. . Englewood Cliffs, NJ: Prentice Hall, 1977. Barkow, J. The distance between genes and culture. Journal of Anthropological Research 40:367-379, 1984. Barkow, .I. The elastic between genes and culture. Ethology and Sociobiology 10: I1 l-130, 1989a. Barkow, J. Darwin, Sex, and Status: Biological Approaches to Mind and Culture, Toronto: Toronto Uni- versity Press, 1989b. Barkow, J., Cosmides, L., and Tooby, J. TheAdapted Mind, Oxford: Oxford University Press, 1992. Barth, F. On the study of . American Anthropologist 69~661-669, 1967. Baumeister, R.F., and Leary, M.R. The need to belong: desire for interpersonal attachments as a funda- mental human . Psychological Bulletin 117:497-529, 1995. Belsky, J., Steinberg, L., and Draper, P. Childhood experience, interpersonal development, and reproduc- tive strategy: an evolutionary theory of . 62:647-670, 1991. Betzig, L. Despotism and DifSerential Reproduction. A Darwinian View of History, Hawthorne, NY: Al- dine, 1986. Betzig, L. Rethinking : a response to some recent critiques. Ethology and Sociobiology 10:315-324, 1989. Betzig, L., Borgerhoff Mulder, M., and Turke, P. (Eds.) Human Reproductive Behaviour, Cambridge: Cambridge University Press, 1988. Bikchandani, S., and Hirshleifer, D. A theory of fads, fashion, custom, and cultural change as informa- tional cascades. Joumal of Political Economy 100(5): 992-1026, 1992. Black, LB. Trophic interactions and brain plasticity. In The Cognitive , M.S. Gazzaniga (Ed.). Cambridge: MIT Press, 1995, pp. 9-17. Boas, F. The Mind ofPrimitive Man, New York Macmillan, 1911. Bock, P.K. Rethinking Psychological Anthropology, New York: W.H. Freeman, 1988. Bock, W. Microevolutionary sequences as a fundamental concept in macroevolutionary models. Evolu- tion 241704-722, 1970. Boehm, C. Rational pre-selection. American Anthropologist 80:265-296, 1978. Bolles, R.C., and Beecher, M.D. (Eds.) Evolution and Learning, Hillsdale, NJ: Lawrence Erlbaum Asso- ciates, 1988. Boone, J. and elite family structure in preindustrial states: A case study of late medi- eval-early modem Portugese . American Anthropologist 89:619-634, 1986. Borgerhoff Mulder, M. Adaptation and evolutionary approaches to anthropology. Man 22:25-41, 1987. Borgerhoff Mulder, M. Kipsigis bridewealth payments. In Human Reproductive Behaviour, L. Betzig, M. Borgerhoff Mulder, and P. Turke (Ed%). Cambridge: Cambridge University Press, 1988, pp. 65-82. Borgerhoff Mulder, M., and Caro, T. The problem of adaptation in the study of human behavior. Ethol- ogy and Sociobiology, 8:61-72, 1987. Bowlby, J. Attachment. New York: , 1969. Boyd, R., and Richerson, P. Culture and the Evolutionary Process, Chicago: University of Chicago Press, 1985. Boyd, R., and Richerson, P. Simple models of complex phenomena: the case of cultural evolution. In The Latest on the Best: Essays on Evolution and Optima&y, J. Dupre (Ed.). Cambridge: MIT Press, 1987. Boyd, R., and Richerson, P. An evolutionary model of social learning: the effects of spatial and temporal variation. In Social Learning: Psychological and Biological Perspectives, T.R. Zentall and B.G. Galef (Eds.). Hillsdale, NJ: Lawrence Erlbaum Associates, 1988a, pp. 29-48. 56 M. V. Fhm

Boyd, R., and Richerson, P. The evolution of reciprocity in sizable groups. Journal of Theoretical Biol- ogy 132:337-356, 1988b. Boyd, R., and Richerson, P. The evolution of ethnic markers. In Ideas in Anthropology, Santa Fe: School of American Research, 1989. Boyd, R., and Richerson, P. Why does culture increase human adaptability? Ethology and Sociobiology 16(2):125-143, 1995. Bayer, P. The Naturalness of Religious Ideas: A Cognifive Theory of Religion, Berkeley: University of California Press, 1994. Brothers, L. The social brain: a project for integrating behavior and in a new domain. Concepts in 1~27-5 1, 1990. Brown, D. Human Universals, New York: McGraw-Hill, 1991. Bmmflel, E. Weaving and cooking: Women’s production in Aztec Mexico. In Engendering Archeology: Women and Ptzhistory, J.M. Gero and M. Conkey (Eds.). Cambridge: Blackwell, 1992. Burling, R. The selective advantage of complex language. Ethology and Sociobiology 7:1-16, 1986. Buss, D. Evolutionary psychology: A new paradigm for psychological science. Psychological Inquiry 6:1-30, 1995. Buss, D.M. Sex differences in human mate preferences: evolutionary hypotheses tested in 37 cultures. Behavioral and Brain Sciences 12:1-49, 1989. Buss, D.M. , New York: Basic books, 1994. Buss, D.M. The evolutionary psychology of human social strategies. In : Handbook of Basic Principles, E.T. Higgins and A.E. Kmglanski (Eds.). New York: Guilford Press, 1996, pp. 3-38. Buss, D.M., Larsen, R.J., Westen, D., and Semmelroth, J. Sex differences in jealousy: evolution, physiol- ogy, and psychology. Psychological Science 3:251-2X, 1992. Byrne, R. The evolution of intelligence. In Behaviour and Evolution. P.J.B. Slater and T.R. Halliday (Eds.). Cambridge: Cambridge University Press, 1994. Byrne, R., and Whiten, A. Machiavellian Intelligence, Social Expertise and the Evolution of Intellect in Monkeys, , and Humans, Oxford: Oxford University Press, 1988. Calvin, W.H. The Cerebral Symphony, New York: Bantam, 1989. Campbell, D.T. Variation and selective retention in . In Social Change in Devel- oping Areas: A Reinterpretation of Evolutionary Theory, H.R. Barlinger, G.I. Blanksten, and R.W. Mack (Eds.). Cambridge: Shenkman, 1965, pp. 1949. Carey, S., and Gelman, R. The Epigenesis of Mind: Essays on Biology and Cognition, Hillsdale, NJ: Lawrence Erlbaum Associates, 199 1. Car&hers, M. Why Human Have Cultures: Explaining Anthropology and Social Diversity, Oxford: Ox- ford University Press, 1992. Cavalli-Sforza, L.L., and Feldman, M.W. Cultural Transmission and Evolution: A Quantitative Ap- proach, Princeton: Princeton University Press, 1981. Chagnon, N.A. Studying the Yunomamo. New York: Holt, Rinehart, and Winston, 1974. Chagnon, N.A. Is reproductive success equal in egalitarian societies? In Evolutionary Biology and Hu- man Social Behavior, N. Chagnon and W. Irons (Eds.). N. Scituate, MA: Duxbury Press, 1979, pp. 374-401. Chagnon, N.A. Socio-demographic attributes of nepotism in tribal populations: man the rule breaker. In Current Problems in Sociobiology, King’s College Sociobiology Group (Eds.). Cambridge: Cambridge University Press, 1982, pp. 291-318. Chagnon, N.A. Life histories, blood revenge, and warfare in a tribal society. Science 239:985-992, 1988. Chagnon, N.A., and Bugos, P.E., Jr. Kin selection and conflict: An analysis of a Yanomamo ax fight. In Evolutionary biology and human social behavior, N. Chagnon and W. Irons (Eds.). N. Scituate, MA: Duxbury Press, 1979, pp. 213-237. Chagnon, N.A., Flinn, M.V., and Melancon, T. Sex ratio variation among the Yanomamo Indians. In Evolutionary Biology and Human Social Behavior, N. Chagnon and W. Irons @Is.). N. Scitu- ate, MA: Duxbury Press, 1979, pp. 290-320. Chagnon, N.A., and Irons, W. (Eds.) Evolutionary Biology and Human Social Behavior, N. Scituate, MA: Duxbury Press, 1979. Cheney, D., and Seyfarth, R. How Monkeys See the World, Chicago: University of Chicago Press, 1990. Cheney, D., Seyfarth, R., and Smuts, B. Social relationships and social cognition in non-human primates. Science 234:1361-1366, 1986. Chibnik, M. The evolution of cultural rules. Journal of Anthropological Research 37:256-268, 1981. Culture and the Evolution of Social Learning 57

Chisholm, J. Death, hope, and sex: life history theory and the development of reproductive strategies. Current Anthropology 34( l):l-24, 1993. Chisholm, J. The of attachment organization. Human Nature 7(l): l-38, 1996. Chomsky, N. Language and Mind (second edition), New York: Harcourt, Brace and Jovanovich, 1973. Cloak, F.T. Is a cultural ethology possible? Human Ecology 3: 161-182, 1975. Colgan, P. Comparative Social Recognition, New York: John Wiley and Sons, 1983. Cooper, W.S. Decision theory as a branch of evolutionary theory: a biological derivation of the Savage axioms. 94:395-411, 1987. Cosmides, L. The logic of social exchange: has natural selection shaped how humans reason? Studies with the . Cognition 31: 187-276, 1989. Cosmides, L., and Tooby, J. From evolution to behavior: evolutionary psychology as the missing link. In The Latest on the Best: Essays on Evolution and Optimality, J. Dupri (Ed.). Cambridge: MIT Press, 1987, pp. 277-306. Cosmides, L., and Tooby, J. Evolutionary psychology and the generation of culture, part II. Case study: a computational theory of social exchange. Ethology and Sociobiology IO:5 l-98, 1989. Cosmides, L and Tooby, J. Cognitive adaptations for social exchange. In , J.H. Barkow, L. Cosmides, and J. Tooby (Eds.). Oxford: Oxford University Press, 1992, pp. 163- 228. Cosmides, L., and Tooby, J. Origins of : the evolution of functional organization. In Mapping the Mind: Domain Specijicity in Cognition and Culture, L. Hirschfeld and S. Gelman (Eds.). New York: Cambridge University Press, 1994. Cosmides, L., and Tooby, J. From function to structure: the role of evolutionary biology and computa- tional theories in cognitive neuroscience. In The Cognitive Neurosciences, M.S. Gazzaniga (Ed.). Cambridge: MIT Press, pp. 1199-1210, 1995. Cosmides, L., Tooby, J., and Barkow, J.H. Introduction: Evolutionary psychology and conceptual inte- gration. In The Adapted Mind, J.H. Barkow, L. Cosmides, and J. Tooby (Eds.). Oxford: Oxford University Press, 1992, pp. 3-15. Crawford, C., and Anderson, J.L. Sociobiology: an environmentalist discipline? American Psychologist 44(12): 1449-1459, 1989. Crawford, C., Smith, M., and Krebs, D. (Eds.) Sociobiology and Psychology, Hillsdale, NJ: Lawrence Erl- baum Associates, 1987. Crews, D. Diversity and evolution of behavioral controlling mechanisms. In Psychobiology of Reproduc- rive Behavior, D. Crews (Ed.). Englewood Cliffs, NJ: Prentice Hall, 1987, pp. 88-l 19. Cronk, L. From hunters to herders: subsistence change as a reproductive strategy among the Mukogodo. Current Anthropology 30(2):224-234, 1989a. Cronk, L. Low socioeconomic status and female-biased parental investment: the Mukogodo example. American Anthmpologist 91(2):414-429, 1989b. Cronk, L. Human behavioral ecology. Annual Review of Anthropology 20:25-53, 1991. Daly, M. Some caveats about cultural transmission models. Human Ecology 10:401408, 1982. Daly, M., and Wilson, M.I. Sex, Evolution and Behavior (2nd edition), Boston: Willard Grant Press, 1983. Daly, M., and Wilson, M.I. Homicide, Hawthorne, NY: Aldine Press, 1988. Daly, M., and Wilson, M.I. Discriminative parental solicitude and the relevance of evolutionary models to the analysis of motivational systems. In The Cognitive Neurosciences, M.S. Gazzaniga (Ed.). Cambridge: MIT Press, 1995, pp. 1269-1286. D’Andrade, R. Cultural meaning systems. In Culture Theory: Essays on Mind, Self; and Emotion, R.A. Schweder and R.A. Le Vine (Eds.). Cambridge: Cambridge University Press, 1984. D’Andrade, R., and Strauss, C. (Eds.) Human Motives and Cultural Models, Cambridge: Cambridge Uni- versity Press, 1992. D’Andrade, R. The Development of . Cambridge: Cambridge University Press, 1995. Darwin, C. , Cambridge: Harvard University Press, (1859)1967. Dawkins, R. The Extended Phenotype, San Francisco: Freeman, 1982. Dawkins, R. . New York: W.W. Norton, 1986. Dawkins, R. (2nd edition), Oxford: Oxford University Press, 1989 [Original work pub- lished 19761. Dennett, D.C. Intentional systems in : the “Panglossian” paradigm defended. Behav- ioral and Brain Sciences 6:343-390. 1983. 58 M. V. Flinn

de Waal, F. 1982 Politics, New York: Harper and ROW. de Waal, F. 1989. Peacemaking among Primates, Cambridge, MA: Harvard University Press. de Waal, F., and Heltne, P.G. (Eds.). Cambridge: Harvard University Press, 1996, pp. 319-334. Dickemann, M. Female infanticide, reproductive strategies, and social stratification: a preliminary model. In Evolutionary Biology and Human Social Behavior, N. Chagnon and W. Irons (Eds.). N. Sci- tuate, MA: Duxbury Press, 1979, pp. 321-368. Dickemann, M. Paternal confidence and dowry competition: A biocultural analysis of Purdah. In Natural Selection and Social Behavior: Recent Research and New Theory, R.D. Alexander & D.W. Tinkle (Eds.). New York: Blackwell Press, 1981, pp. 417-438. Donald, M. The Origins of the Modem Mind, Cambridge, MA: Harvard University Press, 1991. Draper, P., and Harpending, H. Parent investment and the child’s environment. In Parenting Across the Lifespan: Biosocial Dimensions, J.B. Lancaster, J. Altmann, AS. Rossi, and L.R. Sherrod (Eds.). New York: Aldine de Gruyter, 1987, pp. 217-236. Dunbar, R.I.M. of neocortical size, group size, and language in humans. Behavioral and Brain Sciences 16:681-735, 1993. Dunnell, R. Evolutionary theory and archeology. In Advances in Archeological Method and Theory, Vol. 3, M.B. Schiffer (Ed.). New York: Academic Press, 1980, pp. 35-99. Dunnell, R. The concept of progress in cultural evolution. In Evolutionary Progress, M. Nitecki (Ed.). Chicago: University of Chicago Press, 1989a, pp. 169-194. Dunnell, R. Aspects of the application of evolutionary theory in archeology. In ArcheologicalThought in America. C.C. Lamberg-Karlovsky (Ed.). Cambridge: Cambridge University Press, 1989b, pp. 35-48. Durham, W. The adaptive significance of cultural behavior. Human Ecology 4:89-121, 1976. Durham, W. Toward a coevolutionary theory of and culture. In Natural Selection and Su- cial Behavior, N. Chagnon and W. Irons (Eds.). N. Scituate, MA: Duxbury Press, 1979, pp. 39-59. Durham, W. Interactions of genetic and cultural evolution: models and examples. Human Ecology 10:289-323, 1982. Durham, W. Advances in evolutionary culture theory. Annual Review of Anthropology 19: 187-210, 1990. Durham, W. Coevolution: Genes, Culture, and Human Diversity. Palo Alto, CA: Stanford University Press, 1991. Durkheim, E. The Rules of Sociological Method. S. Solovay and J. Meuller, translators. New York: Free Press, 1938 [1895]. Edelman, G.M. Neural Darwinism, New York: Basic Books, 1987. Edelman, G.M. Topobiology, New York Basic Books, 1989. Edelman, G.M. The Remembered Present: A Biological Theory of Consciousness, New York: Basic Books, 1990. Eibl-Eibesfeldt, I. Human Ethology. Hawthorne, NY: Aldine de Gruyter, 1989. Elder, G. Appearance and education in marriage mobility. American Sociological Review 34:519-533, 1969. Ellison, P.T., Lipson, S.F., and Meredith, M.D. Salivary testosterone levels in males from the Ituri forest of Zaire. American Journal of Human Biology 1:21-24, 1989. Ellison, P.T., Peacock, N.R., and Lager, C. Ecology and ovarian function among Lese women of the Ituri forest, Zaire. American Journal of Physical Anthropology 78:519-526, 1989. Essock-Vitale, S. The reproductive success of wealthy Americans. Ethology and Sociobiology 5:4549, 1984. Fisher, R.A. The Genetical Theory of Natural Selection. New York: Dover, 1958 [1930]. Flinn, M.V. Uterine vs. agnatic kinship variability. In Natural Selection and Social Behavior, R.D. Alex- ander and D.W. Tinkle (Eds.). New York: Blackwell, 1981, pp. 439-475. Flinn, M.V. Correlates of reproductive success in a Caribbean village. Human Ecology 14(2): 225-243, 1986. Flinn, M.V. Resources, reproduction, and mate competition in human populations. Behavioral and Brain Sciences 10(3):305-307, 1987. Flinn, M.V. Parent-offspring interactions in a Caribbean village: daughter guarding. In Human Reproduc- tive Behaviour, L. Betzig, M. Borgerhoff Mulder, and P. Turke (Eds.). Cambridge: Cambridge University Press, 1988a, pp. 189-200. Flinn, M.V. Mate guarding in a Caribbean village. Ethology and Sociobiology 9(1):1-29, 1988b. Culture and the Evolution of Social Learning 59

Flinn, M.V. Step and genetic parent/offspring relationships in a Caribbean village. Ethology and Sociobi- ology 9(3): l-34, 1988~. Flinn, M.V. Household composition and female reproductive strategies. In Sexual and Reproductive Strategies, A. Rasa, C. Vogel, and E. Voland (Eds.). London: Chapman and Hall, 1989, pp. 206233. Flinn, M.V. in a Caribbean village. In Father-Child Relations: Cultural and Biosociul Con- fexts, B. Hewlett (Ed.). Chicago: Aldine press, 1992, pp. 57-84. Flinn. M.V., and Alexander, R.D. Culture theory: the developing synthesis from biology. Human Ecol- ogy 10:383-400, 1982. Flinn, M.V., and England, B.G. Childhood stress and family environment. Current Anthropology 36(5):854-866, 1995. Flinn, M.V., and England, B.G. Social economics of childhood glucocorticoid stress response and health. American Journal of Physical Anthropology (in press). Flinn, M.V., and Low, B.S. Resource distribution, social competition and mating patterns in human soci- eties. In Ecological Aspects of Social Evolution, D.I. Rubenstein and R.W. Wrangham (Eds.). Princeton, NJ: Princeton University Press, 1986, pp. 217-243. Flinn, M.V., Quinlan, R., Turner, M., Decker, S., and England, B. Male-female differences in effects of parental absence on glucocotticoid stress response. Human Nature 7(2): 125-162, 1996. Fodor, J.A. The , Cambridge, MA: MIT Press, 1983. Foley, R.A. Evolutionary ecology of hominids. In Evolutionary Ecology and Human Behavior, E.A. Smith and B. Winterbalder (Eds.). Hawthorne, NY: Aldine de Gruyter, 1992, pp. 131- 164. Fredlund, E.V. The use and abuse of kinship when classifying : a Shitari Yanomamo case study. Ethology and Sociobiology 6: 17-25, 1985. Freilich, M. The meaning of “sociocultural.” In The Concept and Dynamics of Culture,B. Bemardi (Ed.). The Hague, 1977. Freud. S. Totem and Taboo, London: , 1938 [1913]. Galef, B.G., Jr. Imitation in animals: History, definition and interpretation of data from the psychological laboratory. In T.R. Zentall and B.G. Galef, Jr. (Eds.). Social Learning: Psychological and Bio- logical Perspectives. Hillsdale, NJ: Lawrence Erlbaum Associates, 1988, pp. 3-28. Galef, B.G., Jr. Introduction. In C.M. Heyes and B.G. Galef, Jr. (Eds.). Social Learning in Animals. New York: Academic Press, 1996, pp. 3-15. Gallistel, CR. The Organization of Learning, Cambridge, MA: MIT Press, 1990. Gallistel, C.R. The replacement of general-purpose theories with adaptive specializations. In The Cogni- tive Neurosciences, MS. Gazzaniga (Ed.). Cambridge: MIT Press, 1995, pp. 1269-1286. Garcia, J. Behavioral regulation of the milieu intern in man and rat. Science 185:824-831, 1974. Gaulin, S.J.C., and Boster, J.S. Dowry as female competition. American Anthropologist 92:9941005, 1990. Gazzaniga, MS. Organization of the human brain. Science 245:947-952. 1989. Gazzaniga, M.S. Nature’s Mind: New York: Basic Books, 1992. Gazzaniga, MS. (Ed.) The Cognitive Neurosciences, Cambridge, MA: MIT Press, 1995. Geary, D.C. Reflections of evolution and culture in children’s cognition: implications for mathematical development and instruction. American Psychologist, 50:24-37. 1995. Geertz, C. Ritual and social change. American Anrhropologist 59:32-54, 1957. Gelman, R. First principles organize attention to and learning about relevant data: number and the ani- mate-inanimate distinction as examples. 14:79-106, 1990. Gemsbacher, M.A. Language Comprehension as Structure Building. Hillsdale NJ: Lawrence Erlbaum, 1990. Gibbs, H. Cultural evolution of male song types in Darwin’s medium ground finches, Geospiza fortis. Animal Behaviour 39:253-263, 1990. Gibson, K.R., and Ingold, T. (Eds.). Tools, Language and Cognition in Human Evolution. Cambridge: Cambridge University Press, 1993. Gigerenzer, G., and Hug, K. Domain-specific reasoning: social contracts, , and perspective change. Cognition 43: 127-171, 1992. Gigerenzer, G., and Murray, D. Cognition as Intuitive Statisfics, Hillsdale, NJ: Lawrence Erlbaum, 1987. Goodenough, O.R. Mind viruses: culture, evolution and the puzzle of altruism. Social Science fnformn- tion 34(2):287-320, 1995. Goodenough, O.R., and Dawkins, R. The St. Jude mind virus. Nature 37:6492, 1994. 60 M.V. Flinn

Goodenough, W.H. Description and Comparison in Cultural Anthropology, Chicago: Aldine, 1970. Gould, J.L. The biology of learning. Annual Review ofPsychology 37: 163-192, 1986. Gould, J.L., and Marler, P. Learning by . 256:74-85, 1987. Gould, S.J. Cardboard Darwinism: this view of life. Natural History 95: 1421, 1986. Gould, S.J. Tires to sandals: this view of life. Natural History 98:8-16, 1989. Gray, Ch.M., Koenig, P., Engel, A.K., and Singer, W. Oscillatory responses in cat visual cortex exhibit inter- columnar synchronization which reflects global stimulus properties. Nature 338:335-337,1989. Gray, P. Primate Sociobiology, New Haven, CT: HRAF Press, 1985. Griffin, D.R. Prospects for a cognitive ethology. Behavioral and Brain Sciences 1:527-538, 1978. Grifftn, D.R. Animal Thinking, Cambridge: Harvard University Press, 1984. Grossberg, S., and Stone, G. Neuronal dynamics of word recognition and recall: attentional priming, learning, and resonance. Psychological Review 93: 46-74, 1986. Hallowell, AI. Self, society, and culture in phylogenetic perspective. In Evolution after Darwin, S. Tax (Ed.). Chicago: University of Chicago Press, 1960, pp. 309-371. Hallpike, C.R. The Principles of Social Evolution, Oxford: Clarendon Press, 1986. Hames, R. Relatedness and interaction among the Ye’kwana: a preliminary analysis. In Evolutionary Bi- ology and Human Social Behavior, N. Chagnon and W. Irons (Eds.). North Scituate, MA: Dux- bury Press, 1979, pp. 238-250. Hames, R. The allocation of among the Ye’kwana. In Human Reproductive Behaviour, L. Betzig, M. Borgerhoff Mulder, and P. Turke (Eds.). Cambridge: Cambridge University Press, 1988, pp. 227-251. Hames, R. Costs and benefits of monogamy and polygyny for Yanomamo women. Ethology and Sociobi- ology 17:181-199, 1996. Hamilton, W.D. The genetical evolution of social behavior I and II. Journal of Theoretical Biology 7: l- 52, 1964. Harnad, S., Steklis, H., and Lancaster, J. (Eds.). Origins and evolution of language and speech. Annals of the New York Academy of Sciences No. 280, 1976. Harpending, H., Rogers, A., and Draper, P. Human sociobiology. Yearbook of Physical Anthropology 30:127-150, 1987. Harris, M. Cows, Pigs, Wars, and Witches, New York: , 1974. Hartung, J. On natural selection and the inheritance of wealth. Current Anthropology 17:607X122, 1976. Hartung, J. Polygyny and the inheritance of wealth. Current Anthropology 23:1-12, 1982. Hartung, J. Matrilineal inheritance: new theory and analysis. The Behavioral and Brain Sciences 8:661- 688, 1985. Hawkes, K. Kin selection and culture. American Ethnologist 10:345-363, 1983. Hawkes, K., Hill, K., and O’Connell, J. Why hunters gather: optimal foraging and the Ache of eastern Paraguay. American Ethnologist 9:379-398, 1982. Hewlett, B.S. and paternal investment among Aka pygmies. In Human Reproductive Be- haviour, L. Betzig, M. Borgerhoff Mulder, and P. Turke (Eds.). Cambridge: Cambridge Uni- versity Press, 1988, pp. 263-276. Heyes, C.M. Imitation, culture, and cognition. Animal Behavior 49:999-1010, 1993. Heyes, C.M. Social learning in animals: Categories and mechanisms. Biological Reviews 69:207-23 1, 1994. Heyes, C.M., and Galef, B.G., Jr. (Eds.). Social Learning in Animals. New York Academic Press, 1996. Hill, J. Concepts as units of cultural replication. Journal of Social and Biological Structures 12:343-355, 1989. Hill, K. Life history theory and evolutionary anthropology. Evolutionary Anthropology 2(3):78-88, 1993. Hill, K., and Hurtado, A.M. Ache Life History, Hawthorne, NY: Aldine de Gruyter, 1996. Hill, K., and Kaplan, H. Tradeoffs in male and female reproductive strategies among the Ache: part I. In Human Reproductive Behaviour, L. Betzig, M. Borgerhoff Mulder, and P. Turke (Eds.). Cam- bridge: Cambridge University Press, 1989a, pp. 277-289. Hill, K., and Kaplan, H. Tradeoffs in male and female reproductive strategies among the Ache: part II. In Human Reproductive Eehaviour, L. Betzig, M. Borgerhoff Mulder, and P. Turke (Eds.). Cam- bridge: Cambridge University Press, 1989b, pp. 291-305. Hinde, R. Individuals, Relationships, and Culture, Cambridge: Cambridge University Press, 1987. Hinde, R. When is an evolutionary approach useful? Child Development 62:671675, 1991. Hinde, R., and Stevenson-Hinde, J. (eds.) Constraints on Learning: Limitations and Predispositions, New York Academic Press, 1973. Culture and the Evolution of Social Learning 61

Hines, W.G.S., and Bishop, D.T. On learning and the evolutionarily stable strategy. Journal of Applied 20:689-2695, 1983. Hirschfeld, L. On acquiring social categories: and anthropological wisdom. Man 23:61 l-638, 1988. Hirschfeld, L., and Gelman, S. (Eds.) Mapping the Mind: Domain Specificity in Cognition and Culture, New York: Cambridge University Press, 1994. Hirshleifer, J. On the emotions as guarantors of threats and promises. In The Latest on the Best: Essays on Evolution and Optimality, J. Dupre (Ed.) Cambridge: MIT Press, 1987. Holloway, R.L. The role of human social behavior in the . Forty-third James Arthur lecture on the evolution of the human brain. New York: The American Museum of Natural His- tory, 1975. Holloway, R.L., and de la Coste-Lereymondie, MC. Brain endocast asymmetry in pongids and homi- nids: Some preliminary findings on the of cerebral . American Journal of PhysicalAnthropology 58:101-l 10, 1982. Hrdy, S.B. The Woman That Never Evolved, Cambridge: Harvard University Press, 1981. Hull, D.L. The naked meme. In Learning, Development, and Culture, H.C. Plotkin (Ed.). New York: Wiley, 1982. Hull, D.L. versus vehicles. In The Role of Behavior in Evolution, H.C. Plotkin (Ed.). Cam- bridge, MA: MIT Press, 1988. Humphrey, N. The social function of intellect. In Growing Points in Ethology, P.P.G. Bateson and R.A. Hinde (Eds.). Cambridge: Cambridge University Press, 1976. Humphrey, N. Consciousness Regained: Chapters in the Development of Mind. Oxford: Oxford Univer- sity Press, 1984. Hurtado, A.M., and Hill, K.R. Paternal effect on offspring survivorship among Ache and Hiwi hunter- gatherers: Implications for modeling pair-bond stability. In Father-Child Relations: Cultural and Biosocial Contexts, B. Hewlett (Ed.). Hawthorne, NY: Aldine, 1992, pp. 57-84. Ingold, T. Evolution and Social Life, Cambridge: Cambridge University Press, 1986. Irons, W. Natural selection, adaptation, and human social behavior. In Evolutionary Biology and Human Social Behavior. N. Chagnon and W. Irons @is.). North Scituate, MA: Duxbury Press, 1979a. pp. 4-39. Irons, W. Investment and primary social dyads. In Evolutionary Biology and Human Social Behavior, N. Chagnon and W. Irons (Eds.). North Scituate, MA: Duxbury Press, 1979b, pp. 181-212. Irons, W. Cultural and biological success. In Evolutionary Biology and Humun Social Behovior, N. Chagnon and W. Irons (Eds.). North Scituate, MA: Duxbury Press, 1979c, pp. 2.57-272. Irons, W. Why lineage exogamy? In Natural Selection and Social Behavior R. D. Alexander and D. W Tinkle (Eds.). New York: Blackwell, 1981, pp. 476-489. Irons, W. Human female reproductive strategies. In Social Behavior of Female , S. Wasser and M. Waterhouse @is.). New York: Academic Press, 1983, pp. 169-213. Johnson, A.W., and Earle, T. The Evolution of Human Societies, Stanford: Stanford University Press, 1987. Johnston, T. Selective costs and benefits in the evolution of learning. Advances in rhe Study of Behavior 12:65-106, 1982a. Johnston, T. Learning and the evolution of developmental systems. In Learning, Development, and Cul- ture, H.C. Plotkin (Ed.). New York: Wiley, 1982b. pp. 41 l-422. Kagan, J., Resnick, J.S., and Snidman, N. Biological bases of childhood shyness. Science 240: 167-l 7 1, 1988. Kaplan, H. A theory of fertility and parental investment in traditional and modem human societies. Amer- icun Journal of Physical Anthropology Suppl. 23:9 l-135, 1996. Kaplan, H., and Hill, K. Food sharing among Ache foragers: tests of explanatory hypotheses. Current An- thropology 261223-245, 1985. Kaplan, H., and Hill, K.R. Hunting ability and reproductive success among male Ache foragers. Current Anthropology 26(1):131-133, 1985. Kaplan, H., and Hill, K.R. The evolutionary ecology of food acquisition. In Evolutionq Ecology and Human Behavior, E.A. Smith and B. Winterhalder (Eds.). Hawthorne, NY: Aldine de Gruyter, 1992, pp. 167-202. Karmiloff-Smith, A., Klima, E., Bellugi, U., Grant, J., and Baron-Cohen, S. Is there a social module? Language, face processing, and theory of mind in individuals with Williams syndrome. Jounal of Cognitive Neuroscience 7: 196-208, 1995. Keeley, L.H. War Before : The of the Peaceful Savage, New York: Oxford University Press, 1996. 62 M.V. Flinn

Keesing, R.M. Theories of culture. Annual Review of Anthropology 3: 73-98, 1974. Kelly, R.L. The Foraging Spectrum, Washington: Smithsonian Institute Press, 1995. Kimura, D. Sex differences in the brain. ScienrificAmerican 9:119-125, 1992. Kitcher, P. VaultingAmbition: Sociobiology and the Questfor Human Nature, Cambridge: MIT Press, 1985. Konner, M. Evolution of human behavior development. In Handbook of Cross-Cultural Human Develop- ment, R.H. Munroe, R.L. Munroe, and B.B. Whiting (Eds.). New York: Garland STPM, 1981, pp. 3-5 1. Konner, M. The Tangled Wing: Biological Constraints on the Human Spirit, New York: Harper Colo- phon Books, 1982. Krebs, J.R., and Dawkins, R. Animal signals: mind reading and manipulation, In Behavioural Ecology: An Evolurionary Approach (second edition), J.R. Krebs and N.B. Davies (Fds.). Sunderland, MA: Sinauer 1984, pp. 380-402. Kroeber, A. Configurations of Cultural Growth, Berkeley: University of California Press, 1944. Kroeber, A. The Nature of Culrure, Chicago: University of Chicago Press, 1952. Kroeber, A., and Kluckhohn, C. Culture, a critical review of the concepts and definitions. Papers of the Peabody Museum of American Archeology and 47: l-223, 1952. Lack, D. Population Studies of Birds, Oxford: Clarendon Press, 1966. Laland, K.N., Kumm, .I., and Feldman, M.W. Gene-culture coevolutionary theory: a test case. Current Anthropology 36(1):131-156, 1995. Lamb, M., Pleck, J., Chamov, E.L., and Levine, J. A biosocial perspective on paternal behavior and in- volvement. In Parenting Across the Lifespan: Biosocial Dimensions, J.B. Lancaster, J. Alt- mann, A. Rossi, and L. Sherrod (Eds.). Hawthorne, NY: Aldine de Gruyter, 1987, pp. 11 l-142. Lancaster, J.B., and Lancaster, C.S. Parental investment: the hominid adaptation. In How Humans Adapt: A Biocultural Odyssey, D. Ortner (Ed.). Washington: Smithsonian Press, 1983. Lancaster, J.B., and Lancaster, C.S. The watershed: change in parental-investment and family-formation strategies in the course of human evolution. In Parenting Across rhe Life-span: Biosocial Di- mensions, J.B. Lancaster, J. Altmann, A.S. Rossi, and L.R. Sherrod (Eds.). New York: Aldine de Gruyter, 1987, pp. 187-206. Leonard, R.D., and Jones, G.T. Elements of an inclusive evolutionary model for archeology. Journal of Anthropological Archeology 6: 199-219, 1987. Lewontin, R.C. The units of selection. Annual Review of Ecology and Systematics 52:1-18, 1970. Lopreato, 3. Human Nature and Biocultural Evolution, Boston: Allen and Unwin, 1984. Lovejoy, C.O. The origin of man. Science 211:341-350, 1981. Low, B.S. Cross-cultural patterns in the training of children: an evolutionary perspective. Journal of Compararive Psychology 103:31 l-319, 1989. Low, B.S. Marriage systems and pathogen stress in human societies. AmericanZoologist 30:325-339,199O. Low, B.S., and Clarke, A.L. Resources and the life course: patterns through the . Ethology and Sociobiology 13:463-494, 1992. Lumsden, C.J. Gene-culture coevolution and the of . Journal of Social and Biolog- ical Structures 6:101-l 14, 1983. Lumsden, C.J. Does culture need genes? Ethology and Sociobiology 10: 1 l-28, 1989. Lumsden, C.J., and Wilson, E.O. Genes, Mind and Culture, Cambridge: Harvard University Press, 1981. MacDonald, K. (Ed.) Sociobiological Perspectives on Human Development, New York: Springer Verlag, 1988a. MacDonald, K. Social and Personality Development: An Evolutionary Synthesis, New York: Plenum Press, 1988b. MacDonald, K. The plasticity of human social organization and behavior: contextual variables and prox- imal mechanisms. Ethology and Sociobiology 10: 171-194, 1989. MacDonald, K. A perspective on Darwinian psychology: the importance of domain-general mechanisms, plasticity, and individual differences. Ethology and Sociobiology 12:449480, 1991. MacDonald, K. Warmth as a developmental construct: an evolutionary analysis. Child Development 63:753-773, 1992. MacDonald, K. The coherence of individual development: an evolutionary perspective on children’s in- ternalization of parental values. In Parenting Strategies and Children’s Inremalizarion of Kzl- ues: A Handbook of Theoretical and Research Perspecrives, J.E. Grusec and L. Kuczynski (Eds.). New York: Wiley, 1996. Mackintosh, N.J. General principles of learning. In Animal Behavior, Vol. 3, T.R. Halliday and P.J.B. Slater (Eds.). Oxford: Blackwell 1983. Culture and the Evolution of Social Learning 63

Maestripieri, D. Maternal encouragement in nonhuman primates and the question of animal teaching. Hu- man Nature 6(4):361-378, 1995. Malinowski, B. Argonauts of the Western Pa@, New York: E.P. Dutton, 1961 [1922]. Marks, J., and Staski, E. Individuals and the evolution of biological and cultural systems. Human Evolu- tion 3:147-161, 1988. Marler, P. Constraints on learning: development of bird song. In Ethology and Psychiatry, N.F. White (Ed.). Toronto: University of Toronto Press, 1974, pp. 69-83. Marler, P., and Terrace, H. (Eds.) The Biology of Learning, Berlin: Springer-Verlag, 1984. Marshack, A. Evolution of the human capacity: the symbolic evidence. Yearbook ofPhysical Anthropol- ogy 32: l-34, 1989. Mayr, E. Animal Species and Evolution, Cambridge, MA: Harvard University Press, 1963. Mayr, E. The Growth ofBiological Thought, Cambridge, MA: Harvard University Press, 1982. Mayr, E. Toward a New , Cambridge, MA: Harvard University Press, 1988. McEwen, B.S. Stressful experience, brain, and emotions: developmental, genetic, and hormonal influences. In The Cognitive Neumsciences, MS. Gazzaniga (Ed.). Cambridge: MIT Press, 1995, pp. 1117-I 135. McGrew, W. Chimpanzee Material Culiure: Implications for Human Evolution, Cambridge: Cambridge University Press, 1992. McNaughton, N. Biology and Emotion, Cambridge: Cambridge University Press, 1989. Meltzoff, A.N. Understanding the intentions of others: re-enactment of intended acts by 18-month-old children. Developmental Psychology 31:838-850, 1995. Menzel, E.W., Davenport, R.K., and Rogers, C.M. Proto cultural aspects of chimpanzee’s responsiveness in novel situations. Folia primatologica 17: 16 I- 170, 1972. Miller, R. Cortico-hippocampal 1nterpla.y and the Representation of Contexts in the Brain, Berlin: Springer Verlag, 199 1. Mineka, S., and Cook, M. Social learning and the acquisition of snake in monkeys. In Social Leam- ing: Psychological and Biological Perspectives, T.R. Zentall and B.G. Galef Jr. (Eds.). Hills- dale, NJ: Lawrence Erlbaum Associates, 1988. Mithen, S. Evolutionary theory and post processual archeology. AntiquiQ 63:483494, 1989. Mithen, S. Though@ Foragers: A Study of Prehistoric Decision Making, Cambridge: Cambridge Uni- versity Press, 1990. Mithen, S. From domain specific to generalized intelligence: a cognitive interpretation of the MiddleiUp- per Paleolithic transition. In The Ancient Mind: Elements of CognitiveArcheology, C. Renfrew and E.B.W. &brow (Eds.). Cambridge: Cambridge University Press, 1994, pp. 29-39. Money, J. Search for the causes of sexual preference. Contemporary Psychology 27:503-505, 1982. Money, J. Human sexology and psychoneuroendocrinology. In Psychobiology of Reproductive Behavior, D. Crews (Ed.). Englewood Cliffs, NJ: Prentice Hall, 1987, pp. 323-344. Montagu, A. (Ed.) Sociobiology Examined, Oxford: Oxford University Press, 198 1. Morgan, L.H. Ancient Society, New York: Holt, 1877. Mueller, R. A. Innateness, autonomy, universality? Neurobiological approaches to language. Behavioral and Brain Sciences, in press. Mundinger, P.L. Animal cultures and a general theory of cultural evolution. Ethology and Sociobiology 1: 183-223, 1980. Murdock, G.P. . New York: MacMillan, 1949. Murdock, G.P. How culture changes. In Man, Culture and Society, H.L. Shapiro (Ed.). Oxford: Oxford University Press, 1956, pp. 247-260. Murdock, G.P. Anthropology’s mythology. In Proceedings of the Royal Anthropological Institute of Great Britain and Irelandfor 1971, 1972, pp. 17-24. Nesse, M.H. (Ed.) Bioaesthetics: bridging the gap between evolutionary theory and the arts. Human Na- ture 6(2), 1995. Nesse, R. Evolutionary explanations of emotions. Human Nature 1:261-289, 1990. Nesse, R., and Williams, G.C. Why We Get Sick, New York: Randon House, 1994. Neville, H. Fractionating language: different neural subsystems with different sensitive periods. Cerebral Cortex 2:244-258, 1992. Nottebohm, F. Birdsong as a model in which to study brain processes related to learning. Condor 86:227- 236, 1984. O’Brien, M., and Holland, T.D. Variation, selection, and the archeological record. In Archeological Method and Theory, Vol. 2, M.B. Schiffer (Ed.). Tucson: University of Arizona Press, 1990, pp. 3 l-79. 64 M.V.Flinn

Ortner, D.J. Biocultural interaction in human adaptation. In How Humans Adapt: A Biocultural Odyssey, D.J. Ortner (Ed.). Washington: Smithsonian Institution Press, 1984, pp. 127-145. Oyama, S. The Ontogeny of Information, Cambridge: Cambridge University Press, 1985. Parsons, T. The Structure of Social Action, New York: Free Press, 1949 [1937]. Payne, R.B. Ecological consequences of song matching: breeding success and intraspecific song in indigo buntings. Ecology 63:401411, 1982. Pepperberg, I. The importance of social interaction and observation in the acquisition of communicative competence: possible parallels between avian and human language. In Social Learning: Psy- chological and Biological Perspectives, T.R. Zentall and B.G. Galef Jr. (Eds.). Hillsdale, NJ: Lawrence Erlbaum Associates, 1988. Perusse, D. Cultural and reproductive success in industrial societies: Testing the relationship at the prox- imate and ultimate levels. Behavioral and Brain Sciences 16:267-322, 1993. Petrinovich, L. The role of social factors in white-crowned sparrow song development. In Social Leam- ing: Psychological and Biological Perspectives, T.R. Zentall and B.G. Galef Jr. (Eds.). Hills- dale, NJ: Lawrence Erlbaum Associates, 1988. Petrinovich, L. Human evolution, Reproduction, and , New York: Plenum Press, 1995. Petrovich, S.B., and Gewirtz, J.L. The attachment learning process and its relation to cultural and biolog- ical evolution: proximate and ultimate considerations. In The Psychobiology of Attachment and Separation, M. Reite and T. Fields (Eds.). New York: Academic Press, 1985, pp. 259-291. Piaget, J. The Child and Reality, New York: Grossman, 1973. Pinker, S. : How the Mind Creates Language, New York: William Morrow and co., 1994. Pinker, S., and Bloom, P. Natural language and natural selection. Behavioral and Brain Sciences 13:707- 784, 1990. Plomin, R. Environment and genes: determinants of behavior. American Psychologist 44: 105-l 11, 1989. Plomin, R. The role of inheritance in behavior. Science 248:183-188, 1990. Plomin, R., DeFries, J.C., and Fulker, D.W. Nature and Nurture During Infancy and Early Childhood, New York: Cambridge University Press, 1988. Plomin, R., and McCleamx, G.E. Nature, Nurture, and Psychology, Washington, DC: American Psycho- logical Association, 1988. Plotkin, H.C. (Ed.) Learning, Development, and Culture: Essays in Evolutionary , New York: Wiley, 1982. Plotkin, H.C. Learning and evolution. In The Role of Behavior in Evolution, H.C. Plotkin (Ed.). Cam- bridge, MA: MIT Press, 1988. Plotkin, H.C. Darwin and the Nature of Knowledge, Cambridge, MA: Harvard Press, 1994. , G.G., and Keates, S.G. The evolution of human cognition and cultural capacity: A view from the far east. In Integrative Paths to the Past, R.S. Corrucini and R.L. Ciochon (Eds.). Englewood Cliffs, NJ: Prentice Hall, 1994. Povinelli, D.J. Reconstructing the evolution of mind. American Psychologist 48:493-509, 1993. Povinelli, D.J. What Chimpanzees (might) know about the mind. In Chimpanzee Cultures, R.W. Wrang- ham, WC. McGrew, F.B.M. de Waal, and P.G. Heltne (Eds.). Cambridge: Harvard University Press, 1996, pp. 285-300. Pribram, K.H. of the Brain, Englewood Cliffs, NJ: Prentice Hall, 1971. Pulliam, H.R., and Dunford, C. Programmed to Learn: An Essay on the Evolution of Culture, New York: Columbia University Press, 197 1. Quartz, S.R., and Sejnowski, T.J. The neural basis of cognitive development: a constructivist manifesto. Behavioral and Brain Sciences (in press). Quiatt, D., and Reynolds, V. Primate Behaviour: Information, Social Knowledge, and the Evolution of Culture, Cambridge: Cambridge University Press, 1993. Radcliffe-Brown, A. Structure and Function in Primitive Society, London: Oxford University Press, 1952. Real, L.A. Animal choice behavior and the evolution of cognitive architecture. Science 253:98&986,1991. Reeve, H.K., and Sherman, P.W. Adaptation and the goals of evolutionary research. Quarterly Review of Biology 68:1-32, 1993. Reynolds, P.C. The complementation and tool use. In Tools, Language and Cognition in Human Evolution, K.R. Gibson and T. Ingold (Eds.). Cambridge: Cambridge University Press, 1991, pp. 407-428. Reynolds, V. The relationship between biological and cultural evolution. Journal of Human Evolution 13:71-79, 1984. Culture and the Evolution of Social Learning 65

Reynolds, V., and Tanner, R. The Social Ecology of Religion, Oxford: Oxford University Press, 1995 [1983]. Richerson, P.J., and Boyd, R. A dual inheritance model of the human evolutionary process. I. Basic pos- tulates and a simple model. Journal of Social and Biological Sfrucfures 1: 127-154, 1978. Richerson, P.J., and Boyd, R. Simple models of complex phenomena: the case of cultural evolution. In The Latest on the Best: Essays on Evolution and Optima&v, J. DuprC (Ed.). Cambridge: MIT Press, 1987, pp. 27-52. Richerson, P.J., and Boyd, R. The role of evolved predispositions in cultural evolution: Or human socio- biology meets Pascal’s wager. Ethology and Sociobiology IO: 195-2 19, 1989. Richerson, P.J., and Boyd, R. Cultural inheritance and evolutionary ecology. In Evolufionaty Ecology and Human Behaviour, E.A. Smith and B. Winterhalder (Eds.). Hawthorne, NY: Aldine de Gruyter, 1989b, pp. 61-92. Rindos, D. Darwinian selection, symbolic variation, and the evolution of culture. Currenr Anthropology 26:65-88, 1985. Rindos, D. The genetics of cultural anthropology: toward a genetic model for the origin of the capacity for culture. Journal ofAnthropological Archeology 5: l-38. 1986a. Rindos, D. The evolution of the capacity for culture: sociobiology, , and cultural selection- ism. Current Anthropology 27(4):315-332, 1986b. Rindos, D. Undirected variation and the Darwinian explanation of cultural change. In Archeological Method and Theory, Vol. 1, M.B. Schiffer (Ed.). Tucson: University of Arizona Press, 1988, pp. 1115. Robin, N., and Holyoak, K.J. Relational complexity and the functions of prefrontal cortex. In The cogni- tive Neurosciences. M.S. Gazzaniga, (Ed.). Cambridge: MIT Press, 1995, pp. 1199-1210. Rogers, A. Does biology constrain culture? American Anthropologist 90:8 19-830, 1988. Romney, A.K. Quantitative models, science and cumulative knowledge. Journal of QuantitativeA.nthro- pology 1:153-223, 1989. Roper, T.J. Learning as a biological phenomenon. In Animal Behavior, Vol. 3, T.R. Halliday and P.J.B. Slater (Eds.). Oxford: Blackwell, 1983. Rosenzweig, M.R. Experience, memory, and the brain. American Psychologist 39:365-376, 1984. Rowe, D.C. The Limits of Family InJuence, New York: Guilford, 1994. Rozin, P. The evolution of intelligence and access to the cognitive unconscious. In Progress in Psychobi- ology and Physiological Psychology, J.M. Sprague and A.N. Epstein (Eds.). New York: Aca- demic Press, 1976. Rumbaugh, D.M., Savage-Rumbaugh, E.S., and Scvcik, R.A. Biobehavioral roots of language. In Chim- panzee Cultures, R.W. Wrangham, W.C. McGrew, F.B.M. de Waal, and P.G. Heltne (Eds.). Cambridge: Harvard University Press, 1996, pp. 319-334. Rumbaugh. D.M., and Washburn, D.A. Counting by chimpanzees and ordinality judgments by macaques in video-formatted tasks. In The Development of Numerical Competence: Animal and Human Models, S.T. Boysen and E.J. Capaldi (Eds.). Hillsdale, NJ: Lawrence Erlbaum Associates. 1993, pp. 87-106. Ruse, M. The evolution of intelligence. In Intelligence and Evolutionary Biology, H. Jerrison and I. Jerr- son (Eds.). Berlin: Springer Verlag, 1988, pp. 13-33. Ruyle, E.E. Genetic and cultural pools: some suggestions for a unified theory of biocultural evolution. Human Ecology 1:201-215, 1973. Sahlins, M. Evolution: specific and general. In Evolution and Culture, M. Sahlins and E. Service (Eds.). Ann Arbor: University of Michigan Press, 1960, pp. 1244. Sahlins, M. Sociobiology: The Use and Abuse of Biology, Ann Arbor: University of Michigan Press, 1976a. Sahlins, M. Culture and Practical Reason, Chicago: University of Chicago Press, 1976b. Sapolsky, R. Stress, social status, and reproductive physiology in free-living baboons. In Psychobiology of Reproductive Behavior, D. Crews (Ed.). Englewood Cliffs, NJ: Prentice Hall, 1987, pp. 291-322. Savage-Rumbaugh, S., and Lewin, R. Kanzi. New York: John Wiley, 1994. Starr, S. Developmental theories for the 1990s: development and individual differences. Child Devetop- ment 63:1-19, 1992. Schwartz, T., and Mead, M. Micro- and macro-cultural models for cultural evolution. Anthropological 3: 1-7, 1961. Service, E. The law of evolutionary potential. In Evolution and Culture, M. Sablins and E. Service (Eds.). Ann Arbor: University of Michigan Press, 1960, pp. 93-l 22. 66 M. V. Flinn

Sherman, P.W., and Holmes, W.G. Kin recognition: issues and evidence. Experimental Behavioral Ecol- ogy, Fortschritte der Zoologie 31:431460, 1985. Shettleworth, S. Learning and behavioral ecology. In Behavioral Ecology (second edition), J.R. Krebs and N.B. Davies (Eds.). Oxford: Blackwell, 1984, pp. 170-196. Sieff, D.L. Explaining biased sex ratios in human populations. CurrentAnthropology 31 :2.5-48, 1990. Siegler, R.S., and Crowley, K. Constraints on learning in nonprivileged domains. Cognitive Psychology 21: 194-226, 1994. Silk, J. and kinship in Oceania. American Anthropologist 82:799-820, 1980. Simon, H.A. A mechanism for and successful altruism. Science 250 (12/21): 1665-1668,199O. Simpson, G.G. Comments on cultural evolution. In Evolution and Man’s Progress, H. Hoagland and R.W. Burhoe (Eds.). New York: Columbia University Press, 1962, pp. 104-108. Smith, E.A. Inujjuamiut Foraging Strategies, Hawthorne, NY: Aldine de Gruyter, 1991. Smith, E.A., and Winterhalder, B. Natural selection and decision making: some fundamental principles. In Evolutionary Ecology and Human Behavior, E.A. Smith and B. Winterhalder (Eds.). Haw- thorne, NY: Aldine de Gruyter, 1992, pp. 25-60. Smith, E.A., and Winterhalder, B. (Eds.). Evolutionary Ecology and Human Behavior. Hawthorne, NY: Aldine de Gruyter, 1992. Smith, M.S. Evolution and developmental psychology. In Sociobiology and Psychology, C. Crawford, M. Smith, and D. Krebs (Eds.). Hillsdale, NJ: Lawrence Erlbaum Associates, 1987. Sober, E. The Nature ofSelection, Cambridge: MIT press, 1984. Spencer, H. First Principles (4th edition), New York Appleton, 1880. Sperber, D. Anthropology and psychology: toward an of representations. Man 20:73-89, 1985. Sperber, D. The epidemiology of beliefs. In Psychological Studies ofWidespread Beliefs, C. Fraser (Ed.). Oxford: Oxford University Press, 199 1. Sperber, D. The modularity of thought and the epidemiology of representations. In Mapping the Mind: Domain Spectficity in Cognition and Culture, L. Hirschfeld and S. Gelman (Eds.). New York: Cambridge University Press, 1994. Sperber, D. Explaining Culture: A Naturalistic Approach, Oxford: Blackwell, 1996. Spiro, M. Cultural relativism and the future of anthropology. Cultural Anthropology 1(3):259-286, 1986. Spuhler, J. (Ed.) Evolution of the Capacity for Culture, Chicago: University of Chicago Press, 1959. Squire, L.R., Knowlton, B. and Musen, G. The structure and organization of memory. Annual Review of Psychology 441453-495, 1993. Staddon, J.E. Adaptive Behavior and Learning, Cambridge: Cambridge University Press, 1983. Staddon, J.E. Learning as inference. In Evolution and Learning, R.C. Bolles, and M.D. Beecher, (Eds.). Hillsdale, NJ: Lawrence Erlbaum Associates, 1988. Standen, V., and Foley, R.A. (Eds.). Comparative Socioecology: The Behavioral Ecology of Mammals and Man. Oxford: Blackwell, 1989. Steams, S.C. The evolutionary significance of . Bioscience 39(7):436-445, 1989. Steele, J. and Shennan, S. (Eds.) The of Human Ancestry: Power; Sex and , Lon- don: Routledge, 1996. Steward, J. Theory of , Urbana: University of Illinois Press, 1955. Symons, D. The Evolution of , Oxford: Oxford University Press, 1979. Symons, D. If we are all Darwinians, what’s the fuss about? In Sociobiology and Psychology, C. Craw- ford, M. Smith, and D. Krebs (Eds.). Hillsdale, NJ: Erlbaum, 1987. Symons, D. A critique of Darwinian anthropology. Ethology and Sociobiology 10:131-144, 1989. Symons, D. The use and misuse of Darwinism in the study of human behavior. In The adapted Mind, J.H. Barkow, L. Cosmides, and J. Tooby @Is.). Oxford: Oxford University Press, 1992, pp. 137-159. Thornhill, R., and Thomhill, N. Human : an evolutionary analysis. Ethology and Sociobiology 4:137-173, 1983. Tinbergen, N. On aims and methods of ethology. Zeitschriffiir fierpsychologie 20:410-433, 1963. Tomasello, M. The question of Chimpanzee culture. In Chimpanzee Cultures, R.W. Wrangham, W.C. McGrew, F.B.M. de Waal, and P.G. Heltne (Eds.). Cambridge: Harvard University Press, 1996, pp. 301-317. Tomasello, M., and Call, J. The social cognition of monkeys and apes. Yearbook of Physical Anthropol- ogy 37~273-305, 1994. Tomasello, M., Gust, D., and Frost, G.T. A longitudinal study of gestural communication in young chim- panzees. Primates 30(1):35-50, 1989. Culture and the Evolution of Social Learning 67

Tomasello, M., Kruger, A.C., and Ratner, H.H. Cultural learning. Behavioral and Brain Sciences 16:495-552, 1993. Tooby, J. The emergence of evolutionary psychology. In Emerging Synrheses in Science, D. Pines (Ed.). Santa Fe: Santa Fe Institute, 1985. Tooby, J., and Cosmides, L. Evolutionary psychology and the generation of culture, part I: theoretical considerations. Ethology and Sociobiology 10:29-50, 1989. Tooby, J., and Cosmides, L. The psychological foundations of culture. In The Adapted Mind, J.H. Barkow, L. Cosmides, and J. Tooby (Eds.). Oxford: Oxford University Press. 1992, pp. 19-l 36. Trivers, R.L. Social Evolution, Menlo Park, CA: Benjamin-Cummings, 1985. Turke, P. On what’s not wrong with a Darwinian theory of culture. American Anthropologist 86:663-668, 1984. Turke, P. : childcare networks on Ifaluk. In Human Repmducfive Behaviour, L. Betzig, M. Borgerhoff Mulder, and P. Turke (Eds.). Cambridge: Cambridge University Press, 1988. Turke, P. Which humans behave adaptively, and why does it matter? Ethology and Sociobiology 11:305- 339, 1990. Tversky, A., and Kahneman, D. Judgment under uncertainty: heuristics and biases, Science 185: 1124- 1131. 1974. Tylor, E.B. , New York: Harper, 1958 (I 87 I). Van den Berghe, P.L. Human Family Systems, New York: Elsevier, 1979. Vining, D.R., Jr. Social versus reproductive success. The central theoretical problem of human sociobiol- ogy. Behavioral and Brain Sciences 9(l): 167-2 16. Voland, E. Human sex ratio manipulation: historical data from a German parish. Journal of Human Evo- lution 13:99-107, 1984. Wallace, A.F.C. Culture and Personality, New York: Random House, 1961. Wallace, R. Cognitive mapping and the and mind. CurrentAnthropology 30:5 18-526, 1989. Wallmann, J. Aping Language, Cambridge: Cambridge University Press, 1992. Weisner, T.S., and Gallimore, R. My brother’s keeper: child and sibling caretaking. Current Anthropol- ogy 18:169-191, 1977. West-Eberhard, M.J. Flexible strategy and social evolution. In Animal Societies: Theories and Facts, Y. Ito, J. Brown, and J. Kikkowa (Eds.). Tokyo: Japan Science Society Press, 1987. White, L. The Science of Culture: A Study of Man and Civilization, New York: Grove Press, 1949. White, L. The Evolution of Culture, New York: McGraw Hill, 1959. Whiten. A. (Ed.) Natural Theories of Mind: The Evolution. Development, and Simulation of Everyday Mind-reading. Oxford: Blackwell, 1991. Whiten, A., Ham, R. On the nature and evolution of imitation in the animal : A reappraisal of a century of research. Advances in the Study of Behavior 21:239-283, 1992. Williams, G.C. Adaptation and Natural Selection, Princeton: Princeton University Press, 1966. Williams, G.C. Natural Selection: Domains, Levels, and Challenges, Oxford: Oxford University Press, 1992. Williams, G.C., and Williams, D.C. Natural selection of individually harmful social adaptations among sibs with special reference to social insects. Evolution 11:32-39, 1957. Wilson, E.O. The Societies, Cambridge: Harvard University Belknap Press, 197 1. Wilson, E.O. Sociobiology, Cambridge: Harvard University Belknap Press. 1975. Wilson, E.O. , Cambridge: Harvard University Press, 1978. Wilson, MI., Daly, M., and Weghorst, S.J. Household composition and the risk of child abuse and ne- glect. Journal of Biosocial Science 12:333-340, 1980. Wolf, A.P. Westermarck Redivivus. Annual Review ofAnthropology 22: 157-175, 1993. Wolpoff, M. Human Evolution, New York: McGraw-Hill, 1995. Worthman. C. Cupid and Psyche: investigative syncretism in biological and psychosocial anthropology. In New Directions in Psychological Anthropology, T. Schwartz, G. White, and C. Lutz (Eds.). Cambridge: Cambridge University Press, 1992. Wright, R. The intelligence test (review of Wonderful Life, by S.J. Gould). The New Republic Jan. 29, 1990: 28-36, 1990. Wright, S. Evolution in Mendelian populations. Genetics 28: 114-l 38, 193 1. Wynn, T. The Evolution of Spatial Competence, Champaign: The University of Illinois Press, 1989. Zentall, T.R., and Galef, B.G. Jr. (Eds.) Social Learning: Psychological and Biological Perspectives, Hillsdale, NJ: Lawrence-Erlbaum Associates, 1988.

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