What Cultural Primatology Can Tell Anthropologists about ...users.clas.ufl.edu/...CulturalPrimatology_ARA_2006.pdf · that cultural primatology has made in deﬁn-ing how cultural

ANRV287-AN35-10 ARI 13 August 2006 6:39

What Cultural PrimatologyCan Tell Anthropologistsabout the Evolutionof CultureSusan E. PerryCultural Phylogeny Group, Max Planck Institute for Evolutionary Anthropology,Deutscher Platz 6, 04103 Leipzig, Germany; Department of Anthropology/Behavior,Evolution, and Culture Group, University of California, Los Angeles, California90095-1553; email: [email protected]

Annu. Rev. Anthropol. 2006. 35:171–90

First published online as a Review inAdvance on May 10, 2006

The Annual Review of Anthropology isonline at anthro.annualreviews.org

This article’s doi:10.1146/annurev.anthro.35.081705.123312

Copyright c© 2006 by Annual Reviews.All rights reserved

0084-6570/06/1021-0171$20.00

Key Words

primates, traditions, cultural transmission, social learning

AbstractThis review traces the development of the field of cultural primatol-ogy from its origins in Japan in the 1950s to the present. The field hasexperienced a number of theoretical and methodological influencesfrom diverse fields, including comparative experimental psychology,Freudian psychoanalysis, behavioral ecology, cultural anthropology,and gene-culture coevolution theory. Our understanding of culturaldynamics and the evolution of culture cannot be complete withoutcomparative studies of (a) how socioecological variables affect cul-tural transmission dynamics, (b) the proximate mechanisms by whichsocial learning is achieved, (c) developmental studies of the role ofsocial influence in acquiring behavioral traits, and (d ) the fitnessconsequences of engaging in social learning.

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INTRODUCTION

The study of culture has been central inthe field of anthropology since the found-ing of the discipline. The question of howthe human cultural capacity evolved is oneof the few questions to which all four sub-fields of anthropology can make significantand complementary contributions. The pur-pose of this review is to describe the progressthat cultural primatology has made in defin-ing how cultural processes operate and howculture evolved, and to relate this progressto research efforts in other branches ofanthropology.

One of the problems that has hamperedanthropology’s progress as a cohesive disci-pline in studying culture is the lack of a com-monly shared definition of what culture is.There are hundreds of definitions, but mostfall into two classes: those focusing on ob-servable behavioral variation, and those fo-cusing on the mental processes or constructsresponsible for producing cultural variants.Because of the difficulty in directly accessingthe contents of nonhuman primates’ minds,cultural primatologists have adopted the for-mer type of definition. Here I define culture asbehavioral variation that owes its existence atleast in part to social learning processes, sociallearning being defined as changes in behav-ior that result from attending to the behavioror behavioral products of another individual.This definition of culture will be viewed asoverly simplified by some scholars, but it hasthe advantage of being easily operationalizedand relevant to both humans and nonhumans.Also, it captures the core element of all pre-vious definitions in cultural primatology andvirtually all previous definitions within cul-tural anthropology: the idea that information,skills, practices, or beliefs are transmitted viasocial inheritance rather than through geneticinheritance [see Laland & Hoppitt (2003),Durham (1991), and Kroeber & Kluckhohn(1952) for further discussion of definitions ofculture].

CULTURAL PRIMATOLOGY:HISTORY OF APPROACHES

Japanese Foundations

The field of cultural primatology was foundedby the Japanese researcher Kinji Imanishi,who in his 1952 paper on “The Evolu-tion of Human Nature” inspired Japaneseprimatologists to search for between-groupdifferences in behavior in nonhuman pri-mates. In the decades that followed, a groupof dedicated researchers produced a steadystream of papers documenting interestingwithin- and between-site variability in thediet, food-processing techniques, courtshipsignals, bathing habits, grooming techniques,and solitary play habits of Japanese macaques,Macaca fuscata (reviewed in Itani & Nishimura1973, McGrew 1998, Perry & Manson 2003).These papers are noteworthy for their rich-ness of description of the transmission pro-cess; many report the precise order in whichindividuals acquired novel traits and discussthis finding in the context of the social or-ganization of the group, reporting groupmembers’ kinship, rank, and spatial organi-zation. In these early papers, researchers as-sumed that behavior was determined eitherby species-universal “instinct” or by “culture,”i.e., that variability in behavior was necessarilya consequence of culture. They did not seri-ously consider the possibility that subtle varia-tions in ecology or individual experience couldinduce individuals to behave differently fromconspecifics at other places owing to asociallearning processes.

“Cultural Panthropology”and its Offshoots

With rare exceptions (e.g., Green 1975,Stephenson 1973, McGrew & Tutin 1978,Hannah & McGrew 1987), Western scien-tists ignored the developing field of culturalprimatology until the 1990s, when the com-munity of chimpanzee (Pan troglodytes) fieldresearchers began reporting striking intersite

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behavioral variation, particularly with regardto tool use (e.g., Boesch 1996, Nishida 1987).This body of research received far more at-tention from the scientific community gener-ally, in part because more of it was written inEnglish and in part perhaps because anthro-pologists found it more plausible to accept thenotion of culture in the closest living relativeto humans than they did in a monkey species.The goal of many of these early chimp culturepapers was apparently to convince cultural an-thropologists that humans are not alone inexhibiting culture, that there is evolutionarycontinuity between humans and other pri-mates regarding cultural diversity and relianceon social learning. These publications (partic-ularly Boesch 1996, McGrew 1992, Whitenet al. 1999) focused on documenting the ge-ographic distribution of behavioral variantsacross study sites, creating a sort of “chim-panzee relations area file.” One strength ofthis body of work is the acknowledgment thatnot all between-site behavioral variation isnecessarily cultural in origin; some variationmay be due to intersite genetic variation or toecological differences that lead individuals atdifferent sites to arrive at different behaviorpatterns because of asocial learning processes(i.e., trial-and-error learning) (Nishida 1987,Whiten et al. 1999). However, in contrastwith the earlier Japanese macaque research,the chimpanzee field researchers rarely de-scribed in their “culture” papers the relevantaspects of within-community social dynamicsthat may have produced the observed patternsof behavioral variation. Perhaps in part forthis reason, nonprimatologists reading theselists of putative cultural differences (divorcedfrom descriptions of the complex social dy-namics of chimpanzees that are presumablyinvolved in creating and maintaining thesevariations) perceived chimpanzee “culture” asrather sterile in comparison with their con-ceptions of human culture. It is not entirelyclear why the chimpanzee researchers aban-doned the Japanese researchers’ tradition ofsupplementing discussions of social transmis-sion sequences with richly detailed “simian so-

ciology.” Perhaps it is because transmissionchains are harder to document accurately ina fission-fusion species like chimpanzees, inwhich all members of the same social groupare rarely found in the same place at the sametime, than in stable social groupings like thoseof macaques. Also, behavioral analyses that arerelatively easy to accomplish for stable socialgroups can be methodologically more difficultin fission-fusion species, in which most groupmembers are out of view of one another mostof the time, and not all dyads have equal op-portunities to interact with one another. An-other factor may be that the two most pres-tigious scientific journals (Science and Nature)limit contributions to a few pages, making itimpossible to report the contextual detail thatwas typical of early Japanese cultural prima-tology. Certainly the chimpanzee researcherswriting about culture were well aware of thecomplexities of chimpanzee social structureand relationship negotiation; in fact, some ofthem had written extensively about these top-ics in other venues. They simply had not inte-grated this information into their articles onculture to the extent that Japanese researchershad.

The landmark publication of the “culturalpanthropologists,” to use Whiten’s (2003)memorable phrase, appeared in Nature andincluded an impressive list of 39 behav-ior patterns (a) that were present in somechimpanzee communities but not othersand (b) for which ecological explanations ofthe geographic variation seemed improbable(Whiten et al. 1999). This paper set chim-panzees apart from other nonhuman primatesas the species having the most elaborate cul-tural diversity. However, similar collaborativestudies had never been performed on otherspecies; it is quite likely that the “behavioraldiversity gap” between chimpanzees and otherprimates will shrink once equivalent stud-ies are performed, just as the Whiten et al.(1999) study shrank the gap between chim-panzees and humans. One other study, onorangutans, of similar design has been pub-lished since Whiten’s. Orangutans seem like

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unlikely candidates for cultural diversity be-cause they lead a more solitary existence thanpractically any other anthropoid primate, yetvan Schaik and his colleagues (2003) docu-mented 24 behavioral variants suspected to becultural in a study comparing 6 study sites inSumatra and Borneo. The orangutan’s virtu-ally solitary life makes it, in some ways, anideal candidate for documenting the acquisi-tion of behavioral traits from particular socialpartners because each individual has extensivecontact with only one or two other associates.Thus, Russon (2003) has the best data set doc-umenting social influence on development offoraging techniques for any wild ape, and herdata make more plausible the claim that theintersite variation seen in orangutans is trulycultural in nature. Recent work by Whitenet al. (2005) on captive chimpanzees also lendscredence to the idea that observed patternsof variation in the field are possibly sociallylearned by demonstrating that (a) individu-als can acquire new techniques for gainingfood by watching a group member who hasbeen trained by an experimenter, and (b) chim-panzees who know multiple techniques tendto conform to the technique predominantlyused by their companions.

Many theoretical reasons show that tradi-tions are as common in many monkey speciesas they are in apes and that they simply havenot been discovered owing to lack of researcheffort by researchers interested in such ques-tions (Perry & Manson 2003). In general,monkeys live in social settings in which theyhave a larger number of social contacts fromwhom they could acquire behavioral practices,relative to those apes (chimps and orangutans)in which cultural diversity has been doc-umented best. Also, many monkeys areomnivorous (which should make social cuesuseful in making food choice decisions) andengage in extractive foraging (which wouldmake tool use and other complex forag-ing techniques useful). Likewise, many mon-keys exhibit cooperative social relationshipsand live in complex social environments ofthe type that might necessitate extensive

communication about their relationships andpromote the development of flexible bond-testing rituals (Perry et al. 2003a).

Mechanisms of Social Learning

While field researchers in the 1990s were busydocumenting behavioral variations in chim-panzees, laboratory researchers were hard atwork documenting the cognitive mechanismsunderlying cultural transmission. Early workon the cognitive psychology of social learningwas guided by two conflicting anthropomor-phic assumptions, both of which turned outto be false to some degree. Because imitation(copying of a model’s motor actions) comes soeasily to human children, and because folkloreabout primates is riddled with “monkey see,monkey do” stories, some researchers (e.g.,the early Japanese primatologists) started withthe assumption that imitation must be easy,a cheap trick for learning many new skillsquickly. Other researchers, noting that ex-treme cultural elaboration in humans greatlyexceeded cultural capacities in nonhumans,assumed that some extremely complex cog-nitive machinery must be necessary to pro-duce culture; therefore, they speculated thatthere would be sharp differences between hu-mans and apes, and likewise between apes andmonkeys, in their capacities to learn socially,innovate, and form traditions. Tomasello &Call (1997), in their review of the literature,found considerable evidence for simple formsof social learning such as local and stimulusenhancement (i.e., being attracted to a par-ticular object or location with which a modelis engaged) and emulation learning (observa-tional learning of the cause-and-effect rela-tionships created by a model’s manipulations)in both monkeys and apes. The apes were bet-ter than monkeys at emulation, and human-raised apes were far better at imitation thaneither ape-reared apes or monkeys. The gen-eral conclusion by the early 1990s was thatcopying motor actions was easy for humans,difficult for apes (although present at least inhuman-reared, i.e., “enculturated,” apes) and

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essentially absent in monkeys (Byrne 1995,Tomasello 1996).

Some of the most detailed and careful workon social learning was performed on captivebrown capuchin monkeys (Cebus apella) byFragaszy’s and Visalberghi’s labs (see Fragaszyet al. 2004 for a review of this body of work).The tasks that they gave their animals wereall centered around food choice and food-processing techniques (some requiring merelydexterity, such as foraging on pumpkin seeds,and others requiring tool use such as nutcracking, obtaining peanuts from inside glasstubes with sticks, and operating a juice dis-penser). The general conclusions from theseexperiments were that social influence is im-portant in the development of particular skillsand preferences but that relatively simplesocial-learning processes such as social facili-tation (an increase in a behavioral element al-ready present in the repertoire that is contin-gent on concurrent performance of the samebehavior by others in the vicinity), stimulusenhancement, and local enhancement, ratherthan imitation, are typically employed by themonkeys. For example, in the food choice ex-periments, the subject monkeys were stimu-lated to increase their consumption of bothnovel and familiar foods when the demonstra-tor monkey was also eating, but they did notprefer the particular food eaten by the demon-strator (Visalberghi & Addessi 2003). Theyargued that capuchins’ tendencies to coordi-nate their activities in space and time chan-nel their behavior such that they are likely,in the wild, to end up trying the foods eatenby the more knowledgeable members of theirsocial group, even if they do not attend verywell to the specific properties of those foods.Capuchins who were given ample opportu-nity to observe experienced monkeys pokingsticks through tubes to remove the peanuts in-side them failed to show that they understoodthe relevant aspects of the tube, implying thatthey were not imitating the model. Again, itseems that capuchin monkeys do gain an en-thusiasm for particular tasks and objects bywatching others engage in them, even though

they do not learn fine-grained details aboutthe way in which objects are to be manipu-lated. Visalberghi (1993) and Fragaszy et al.(2004) concluded that the role of social in-fluence is to channel the monkeys’ attentiontoward particular objects and general activi-ties, after which they gain knowledge of thetask via trial-and-error learning.

Puzzlingly, however, results from wildCebus capucinus in the field indicate that youngcapuchins do visually attend to the specificproperties of foods being processed by group-mates: They show significant preferences forobserving groupmates who are foraging onfood items that are rarer in the diet, as well asthose items that require more complex pro-cessing, which suggests that they are seekingout rather specific information about what toeat and how to process food (Perry & Ordonez2006). Data from wild C. capucinus also showthat the more time pairs of monkeys spend inclose association with one another, the morelikely they are to share a particular foragingtechnique (Panger et al. 2002; S. Perry, un-published observations); such results are diffi-cult to reconcile with a social-learning modelthat excludes imitation entirely. Although im-itation is difficult for most nonhuman pri-mates, and it is not nearly so often employedin their daily lives as are the cognitively sim-pler social-learning mechanisms, imitation isnonetheless possible for many nonhuman pri-mates under a narrow range of circumstances.This interpretation is consistent with one cap-tive study that produced some suggestive ev-idence that capuchins can imitate under cer-tain conditions, although they appear to do itquite rarely (Custance et al. 1999).

Numerous studies of imitation have beendone in a variety of taxa in the past decade, andnow the distinctions drawn previously are lesswell defined (see Whiten et al. 2004 for a re-view of social learning in apes). Researchershave documented imitative capacities notonly in chimpanzees and orangutans, buteven in callitrichids (Voelkl & Huber 2000)and in some nonprimates (e.g., birds: Zentall2004).


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To summarize the state of the literature onsocial-learning mechanisms, the picture is stillmurky as to which mechanisms are employedby which species and under which circum-stances, and much work is still needed to sortout this mess (see for example Whiten et al.2004). There are, however, some clear take-home messages. All primate species that havebeen thoroughly investigated rely on a num-ber of social-learning mechanisms, some ofthem quite simple (e.g., attraction to locationsoccupied by groupmates or to objects beinghandled by groupmates) and others perhapsmore cognitively challenging (such as associ-ation of an object with a particular behavioralgoal; or imitation of fine motor details). Thesimpler mechanisms are more commonly em-ployed than is true imitation in monkeys, apes,and even humans (Tomasello 1999). True im-itation is not an exclusively human cogni-tive trick, but it is certainly more commonlyused by humans than by other species (Boesch& Tomasello 1998, Tomasello 1999). One ofthe main challenges for psychologists workingin this area will be to determine which fac-tors trigger the use of certain types of social-learning mechanisms as opposed to others.Which features of the task, of the demonstra-tor, and of the social context stimulate the an-imal to pay attention to details such as thephysical properties of the objects being ma-nipulated (e.g., color, smell, size, shape), themotor movements of the demonstrator, theapparent behavioral goals of the demonstrator(as determined by the consequences of theiractions), or the movements of manipulatedobjects relative to other objects?

Cultural Transmission Theory

The study of culture in nonhumans becamefar more interesting theoretically when be-havioral ecologists began to focus on cul-ture. Theoreticians such as Boyd & Richerson(1985), Cavalli-Sforza & Feldman (1981), andLaland (Laland & Kendal 2003) began tomodel gene-culture coevolution and to pro-mote cultural transmission as a topic that had

relevance to biology (see Richerson & Boyd2004 for an accessible account of the ideas ex-pressed by these formal models, and Durham1991 for empirical examples of gene-culturecoevolution). For the first time, culture wasviewed as a topic of serious scientific inquiryin biology, and behavioral biologists were cap-tivated by the idea that certain behaviors couldbe inherited socially in a manner roughly anal-ogous to the way in which genetic inheritanceoccurs. Laland and his colleagues (Laland& Kendal 2003, Odling-Smee et al. 2003)were particularly influential as ambassadorsof gene-culture coevolution theory to prima-tologists, and they emphasized that sociallylearned behaviors could alter the environmentin such a way that the modified environmentexerted selective pressure on genetically in-herited traits. This process by which animalscan, by their own behavior, alter the selectivepressures on future generations was dubbedniche construction (Odling-Smee et al. 2003).Although I know of only one empirical studyin primatology that directly incorporates thisidea (Flack et al. 2006), it was nonetheless aneffective attention getter in that it brandedsocial learning as a topic of evolutionary im-portance and thereby diverted research efforttoward social learning in general.

The Socioecology of Animal SocialLearning and Traditions: What, Why,When, and Who?

Once more behavioral ecologists joined thecultural primatology bandwagon, the researchemphasis started to shift away from the ques-tion “Does Species X have culture?” to moretheoretically interesting questions such as“Under which conditions should individualsengage in social learning?,” “Who should anindividual copy?,” “How do various socioeco-logical variables affect the dynamics of socialtransmission?,” and “What adaptive benefitsaccrue to animals who engage in social learn-ing?” (see Laland 2004 for a cogent discus-sion of the first two issues). Several researchprograms are actively engaged in answering

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the first three questions [see for example pa-pers in Box & Gibson 1999, Fragaszy & Perry2003a, Heyes & Galef 1996, as well as thespecial issue of Learning and Behavior (2004;vol. 32, number 1) devoted to social learning].The latter question (“How is social learningadaptive?”) remains difficult to answer in anempirically rigorous way, although it is thetopic of frequent speculation. Clearly the so-cial learning of new foraging strategies im-proves the efficiency of foraging, and perhapsopens up new resources that would other-wise be unavailable (e.g., Boesch & Boesch1990). Some scholars have proposed that so-cial learning may aid primates in identifyingpredators and medicinal plants, guide choicesof where to look for food, determine appropri-ate travel routes, define group membership,determine appropriate mating or social part-ners, assess rivals, and communicate about thequality of social relationships (see review inPerry 2003). Table 1 shows a few examples ofputative traditions documented (with varyinglevels of certainty about the role of social in-fluence) in the four best-studied primate gen-era. This is by no means an exhaustive reviewof wild primate traditions, but the review ar-ticles cited in the table will direct the readerto most well-documented primate traditionsdiscovered prior to 2003.

Some researchers focused primarily onhow the speed and extent of cultural trans-mission is affected by various sorts of demo-graphic conditions or behavior types. For ex-ample, Huffman & Hirata (2003) found noconsistent effect of group size on diffusionrates, although they did find that innovationsregarding food type and experimental tasksspread more slowly than do play and food-processing innovations. Similarly, van Schaik& Pradhan (2003) created a model that sug-gests that group size has very little impact onthe likelihood of social transmission. Otherresearchers emphasized how factors such asthe rate of environmental change, the costof acquiring a trait by trial-and-error learn-ing, and the relative reliability of social ver-sus asocial cues would affect the tendency of

animals to engage in social learning (see re-views by Dewar 2003, Laland 2004, Laland &Kendal 2003, Richerson & Boyd 2004). Thesetopics were also pursued by empirical re-searchers of birds, rodents, and other species(see for example Fragaszy & Perry 2003a). Butthe most quintessentially primatological topicto emerge during this time period was thestudy of how personality factors, emotions,and the quality of social relationships influ-ence the probability that social learning willoccur. Nonhuman primates, like humans, ex-hibit considerable interindividual variation inpersonality types and emotional expression,and they live in complex societies in whichsocial relationships are often sharply differ-entiated (Aureli & Schaffner 2002, Capitanio2004). For these reasons, it seems logical toassume that not all pairs of individuals areequally likely to learn from one another, anda good understanding of the social dynamicsof a society will enhance our understanding ofthe dynamics of cultural transmission.

The Japanese primatologists recognizedearly on that there must be some link be-tween factors such as personality, social dy-namics, and the effectiveness of social trans-mission (Imanishi 1957). This early work wasinspired by a Freudian theoretical frameworkrather than a socioecological one (the field ofsocioecology had yet to be created) and wassupported more by case studies and anecdotes(as is typical in psychoanalytic research) ratherthan by more systematic quantitative tests ofhypotheses. Imanishi’s writings were some-what vague on the mechanics of how iden-tification (the “mechanism for introducingculture into personality”) worked, and his as-sumptions regarding young animals’ choicesof role models (i.e., that cultural traits werelearned almost exclusively from the motherand from the “leader male”) were not rig-orously tested. Kawamura (1959) also em-phasized the importance of troop-specific at-titudes as key factors determining the easeof social transmission and the importance ofrelationship quality in predicting the prob-ability of transmission. He noted that the


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monkeys of some troops were far more re-laxed around humans and also more curiousabout novel items being held by their group-mates, and those groups exhibiting more openattitudes were more likely to acquire andtransmit new behaviors such as the adop-tion of novel dietary items. Kawamura, likeImanishi, thought that the social organizationcontributed to the acquisition of a basic atti-tude that was then adopted (somehow by so-cial transmission) by all group members (Itani& Nishimura 1973) and that this attitude facil-itated or inhibited social-learning processes.Subsequent work by researchers working withJapanese macaques were attentive to factorssuch as kinship, rank, class, and centrality inthe group’s spatial structure when describingpatterns of social transmission [see Itani &Nishimura (1973) for an English review ofthis literature]. Itani emphasized the impor-tance of knowing the history of experiencesfor the members of each troop: For exam-ple, history of provisioning can have a pro-found effect on the openness of monkeys totrying new foods (Itani & Nishimura 1973).The patterns that emerged were fairly com-plicated, such that it was not possible to makestrong generalizations about how particularsocial and demographic factors influenced so-cial transmission; rather, the type of culturaltrait (e.g., food choice, food processing, play-related behavior) had a strong impact on thetransmission speed, the type of innovator, andthe pathway by which social transmission oc-curred (e.g., by mother to offspring, frompeers to peers, etc.) (Huffman 1996, Huffman& Hirata 2003). This makes sense because noone individual is likely to be the most knowl-edgeable and skilled performer of all types ofbehaviors, nor do all group members have thesame needs regarding the sorts of behaviorsthat are appropriate for them to learn at aparticular time in their lives. A more sophis-ticated and flexible social-learning strategy isexpected that takes into account the observer’scurrent knowledge and skills, the availabilityof asocial cues for learning the necessary skills,the skill level and tolerance levels of avail-

able models, and the specifics of the task itself(Laland 2004).

After a long gap in which there was verylittle further discussion of the role of person-ality and social dynamics in social transmis-sion processes, an important theoretical pa-per was published emphasizing that “moreextensive and more frequent behavioral co-ordination in time and space will be achievedamong groups exhibiting an egalitarian or tol-erant style of social dynamics” (Coussi-Korbel& Fragaszy 1995, p. 1446) and that such be-havioral coordination is presumed to enhancethe effectiveness of social learning. If two in-dividuals can relax in one another’s presencewithout one fearing domination or food theftby the other, then there is more opportunityfor the animals to associate frequently and toconcentrate on learning a new task in one an-other’s presence. The idea that the quality ofsocial dynamics affects the probability of so-cial transmission began to receive more publicrecognition in anthropology when Boesch &Tomasello (1998) published their theories re-garding the relationship between social struc-ture and social transmission processes. Forexample, they emphasized that the extent towhich social relationships are differentiated(i.e., extreme variation in the quality of rela-tionships between dyads) is expected to affectthe homogeneity of cultural traits. Also, theextent to which the social structure is egalitar-ian is predicted to affect the degree to whichsome group members can impose social normson others, which in turn will affect the homo-geneity and patterning of cultural transmis-sion. In 1999, van Schaik et al. published theirmodel of the evolution of material culture.One of the central tenets of the van Schaiket al. model was that both gregariousness andinterindividual tolerance during foraging ac-tivities are necessary for a tool use innovationto be propagated. Thus far, the van Schaiket al. model has been tested only in a broad-strokes sort of way, for example, by doingcross-site comparisons in which mean partysize is used to predict the number of feed-ing traditions at a particular site (van Schaik


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2003, van Schaik et al. 2003). No attempt hasbeen made to do the far more time-consumingexercise of investigating within-group diver-sity of relationship quality, with direct mea-sures of tolerance, and its effects on socialtransmission. However, this task is currentlybeing undertaken at my field site for white-faced capuchins, as well as by experimentalresearchers working on social learning in cap-tive brown capuchins (K. Bonnie and F.B.M.de Waal, personal communication).

CULTURAL PRIMATOLOGYAND CULTURALANTHROPOLOGY: PROSPECTSFOR MUTUAL INFLUENCE

One of the virtues of the theoretical orienta-tion of cultural primatology that could prof-itably be imported into cultural anthropologyis the tendency to examine multiple levels ofexplanation (following Tinbergen 1963) forthe existence of culture (Fragaszy & Perry2003b, McGrew 1998). An ethologist study-ing culture from Tinbergen’s approach wouldask such questions as the following:

1. Proximate level of analysis: By whichmechanisms are cultural traits transmit-ted from one individual? Which factorsenhance or inhibit the social transmis-sion of information or skills?

2. Developmental level of analysis: Overthe lifespan of an individual animal, howare new behavioral traits acquired? (e.g.,To what extent is social learning in-volved in the acquisition of new be-havioral traits, and from whom is theanimal learning these traits?) Whichfactors speed the acquisition of sociallylearned traits?

3. Phylogenetic level of analysis: What isthe evolutionary history of culture? Towhat extent was the human form ofculture present in our ancestors andour closest living relatives (i.e., thegreat apes)? Looking across a broadtaxonomic range, in what taxa do wesee human-like social-learning abilities,

and what can this taxonomic distri-bution tell us about the evolution ofculture?

4. Ultimate or adaptive level of analysis(i.e., the typical genetic consequencesof engaging in a particular behavior):What is culture good for? What is theadaptive value of having and employ-ing social-learning mechanisms? Underwhich circumstances is it adaptive torely on social cues as opposed to asocialcues for making behavioral decisions?

Cultural primatologists and other ethol-ogists have addressed all these questions,whereas cultural anthropologists have tended,throughout the history of the discipline, tofocus on a different set of questions. Culturalanthropologists have focused more on the pat-terning of cultural elements across culturegroups (a topic rarely addressed by culturalprimatologists, although see Whiten et al.1999 and van Schaik et al. 2003 for somequite preliminary explorations in this direc-tion). The early diffusionists were interestedin how cultural elements were patterned ge-ographically (e.g., Ratzel 1896–1898) and inthe way core cultural elements cohered into“worldviews” (Frobenius 1933) or trait com-plexes (Herskovits 1926). Other scholars, in-cluding many archaeologists (e.g., Flannery1972, 1973) and linguists (e.g., Croft 2000;Labov 1994, 2001; Thomason & Kaufman1988) examined cultural macroevolution—e.g., the phylogeny of languages and hori-zontal diffusion of linguistic traits. Ratzel, forexample, discusses how migrations and con-quests by one ethnic group or another in-fluence cultural diffusion. In this early work,the emphasis was on cultures or culturaltraits as the unit of analysis, and the issueof how individual actors come to adopt thecustoms of their culture was ignored. Lateranthropologists such as Kroeber (1963) andGluckman (1955) emphasized culture changeas a product of internal sociopolitical changesrather being primarily a product of contactbetween culture groups. The emphasis on

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cultural traits as the unit of analysis is sharedby biologically inspired research on memet-ics (see papers in Aunger 2000 for a criticalevaluation of the “meme” concept). Althoughthere is much to be gained from such an ap-proach, it is important to realize that culturaltraits do not have a life of their own; they areexpressed and transmitted by individual ac-tors, who vary in interesting ways accordingto their psychological characteristics, devel-opmental histories, and positions in a socialnetwork. Both the properties of the “memes”(or cultural traits) and the properties of theindividual actors are important when consid-ering the probability of cultural transmission.

Only relatively recently in the history ofanthropology have anthropologists begun tolocalize cultural concepts in the minds of in-dividuals and to consider which factors mightmake some concepts and practices more sta-ble and transmissible than others. Cogni-tive anthropologists [e.g., Atran (1990), Boyer(1999), Sperber & Hirschfeld (2004)] discussthis issue at length, as do many linguistic an-thropologists [see, for example, Blevins (2004)for proximate and developmental factors af-fecting the transmission quality of phono-logical traits, and also references cited inRicherson & Boyd (2004)]. In addressing thisissue of the interface between the psychol-ogy of learning and the characteristics of cul-tural traits, cultural primatology is of mostuse to anthropology generally. Researcherssuch as Whiten (2002), Tomasello (1999), andGreenfield (Greenfield et al. 2000) have donecomparative work with both humans and non-human primates examining the cognitive pro-cesses involved in social learning. Althoughour understanding of the specific mechanismsinvolved in learning is still far from perfect(see above), much headway has been made indetermining the similarities and differencesbetween species in their social-learning ca-pacities and also in determining the factorsaffecting within-species variation in the con-texts in which individuals depend on particu-lar types of social cues. New research on theneurobiology of social learning (e.g., work on

mirror neurons, which fire both when an ac-tion is observed and when it is performed)will no doubt enhance our understanding ofthe proximate mechanisms underlying socialtransmission (Williams et al. 2001).

The second important contribution of cul-tural primatology is the focus on how so-cial and ecological variables (such as those tobe described in the final section of this re-view) affect individuals’ tendencies to adoptparticular sorts of cultural traits and the fo-cus on how social dynamics affect diffusionof cultural traits. Scholars presume that re-lationships between these variables will holdin humans as well as in nonhuman primates.Very little work has been done on the pre-cise way in which different relationship qual-ities and interaction styles (such as toleranceof the teacher or model, history of coopera-tion and hence trust between the model andlearner, dominance style, and mutual emo-tional engagement/joint attention during theactivity) affect the ease of social transmissionin humans. However, evidence from studies ofItalian schoolchildren demonstrates that earlyprosocial behavior in humans (whether ratedby self, peers, or teachers) is a better predictorof academic performance than is prior aca-demic performance or aggressive tendencies(Caprara et al. 2000). It is not clear whichcomponents of prosocial behavior cause thisenhancement of sociocognitive abilities. It isextraordinarily difficult to obtain longitudinaldata on the details of social dynamics, person-ality factors, and learning opportunities (notjust of academic skills but also of the skills im-portant for daily life) over the course of anindividual’s development. The fission-fusionnature of human society and the strong ob-jection that privacy-valuing humans exhibitto being followed constantly by a researchermake these sorts of data hard to collect. Al-though such data are time consuming to col-lect in any species, it is easier to collect themin nonhuman primates than in humans. Be-cause they share many of those human char-acteristics that are critical for the emergenceof culture, nonhuman primates can provide a


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useful model for explaining which factors pro-mote social transmission.

Cultural primatologists and cultural an-thropologists alike would do well to thinkmore about the role of emotions in facili-tating cultural transmission. The subfield ofcultural anthropology known as psychologi-cal anthropology has addressed this issue inways that are roughly analogous to the wayJapanese primatologists looked at personalityand culture. However, psychological anthro-pologists have used a variety of approachesto study the relationship between personal-ity and culture, some focusing more on in-dividual decision making and others focus-ing more on broader cultural patterns andculture-specific personalities (see Bock 1988for a review of the early history of this field).Clearly the types of emotions displayed bymodels influence the amount and quality ofattention they receive from observers, whichaffects what they learn. Emotions are likely tobe particularly important in establishing so-cial norms, in which case the type of emotionaldisplay affects the way the two participants inan interaction respond to one another. For ex-ample, conformist transmission in humans islikely to be emotionally mediated via shame(Fessler 1999): People are ashamed to be dif-ferent or to be ridiculed by others, and thissimple mechanism is important in giving riseto all sorts of traditions that are seemingly ar-bitrary, such as which hand to hold your forkin. Shame, pride, and the self-righteousnessassociated with punishing people for not con-forming to social norms may be essential forcreating the cultural complexity we see in hu-mans. These emotions are thought to be lack-ing in nonhuman primates (Fessler 1999), andthis lack may explain (proximately and phylo-genetically) some of the differences betweenhuman cultures and simpler nonhuman cul-tures. Another factor that may be important indifferentiating nonhuman cultures from hu-man cultures is the prevalence of prestige-biased transmission in humans (Henrich &Gil-White 2001); prestige (freely conferreddeference based on respect, as opposed to

dominance, which is deference due to fearof aggression) is likely a phenomenon that israre or nonexistent in nonhuman primates, al-though this is a relatively unexplored area ofresearch in primatology (but see Ottoni et al.2005). The range of emotions possible in aspecies may be correlated with the types ofcultural transmission, social complexity, andcultural complexity that are possible.

And what can cultural primatologists learnfrom cultural anthropologists? Cultural an-thropologists have always emphasized the im-portance of historical perspective. Culturalprimatology is such a new field that very fewcurrently ongoing studies can claim to havea historical perspective on culture change [al-though Huffman’s (1996) study of stone han-dling in macaques and the study by Perryet al. (2003a) of the appearance and disso-lution of social conventions in capuchins area step in the right direction]. Primatologistswho have studied culture should, research re-sources permitting, maintain long-term datacollection on these topics in the same popula-tions to add a temporal component to theirstudies. Another important contribution bycultural anthropologists is the emphasis onthe patterning of cultural elements [i.e., thestudy of how certain cultural elements seemto co-occur consistently owing to an under-lying symbolically based ideology (Frobenius1933) or for functional reasons (Harris 1979)].For example, a particular subsistence style(e.g., horticulture or pastoralism) and its as-sociated technologies tend to co-occur witha particular descent system (e.g., matrilin-eality or patrilineality) and a particular re-ligion that contains symbolic elements thathelp organize and reinforce that descent sys-tem. Among the most striking differences be-tween human culture and nonhuman primatetraditions, given our current understandingof these issues, are (a) the difference in thedegree to which symbolic communication isinherent in the cultural traits and degree towhich cultural traits are clustered around aconceptual core, (b) the degree to which ac-tive teaching is part of the social transmission

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process in humans (although it is importantto consider the possibility that active teach-ing plays a more important role in Westerncultures than in other human populations),and (c) the extent to which human cultureis cumulative—i.e., that new cultural vari-ants are modifications of past cultural variantsrather than brand new innovations (Richerson& Boyd 2004, Tomasello 1996, Whiten et al.2003). Although these differences are impor-tant, it nonetheless seems that humans andnonhuman primates have much in commonregarding their social transmission processesand that further comparative work will be cru-cial to understand the evolution of humancultural capacities.

A WORKING MODEL OF HOWCULTURAL TRAITS ARISE

Here I present a model (Figure 1) of howmost (or at least many) cultural primatolo-gists conceptualize the process of how cul-tural traits are formed and maintained in non-human primates and humans. Although thismodel has not been presented in its entiretyin any other place, work by other authors hasdeveloped particular links in this model, as ar-ticulated earlier in this paper. Of course, manyother aspects of the cultural process have beenomitted from this model—for example, I havenot explicitly discussed gene-culture coevolu-tion here. The reason is because this topic hasnot yet been addressed empirically in nonhu-man primate studies; nor do I know of any re-searcher who has imminent plans to do so. Acomplete model of cultural processes would,of course, have to integrate genetic feedback(Richerson & Boyd 2004).

Basically, cultural primatologists are inter-ested in how external variables (i.e., aspects ofthe physical and social world) affect particularpsychological attributes of animals (i.e., atti-tudes toward particular tasks and individuals,and also mental representations of particularactions), which in turn affect the tendency forthem to exhibit behavior patterns that are ei-ther similar to, or different from, the behav-

Psychological attributes(e.g., attitudes toward environment

and toward conspecifics)

Behavior patterns(e.g., traditions and social norms,

or lack thereof)

External variables(e.g., demographic factors,

food abundance and distribution,

risk level)

Figure 1Model of factors affecting the propensity to develop traditions and socialnorms.

iors of their groupmates in similar contexts.The external variables can exert an effect onpsychological attributes and behavior on anevolutionary time scale (via natural selection)and/or on an ontogenetic time scale. Let mefirst describe in more detail what I mean byeach of these three categories and then givesome specific examples of how particular sortsof traditions and social norms could arise.

The sorts of external variables I have inmind consist primarily of those variables thathave long interested primate socioecologists:aspects of the demography and ecology of theanimals. For example, variables such as sex ra-tio, group size, birth cohort size, and the ratioof adults to immatures in a group will affectthe number and type of models from whichany particular individual can learn. Availabil-ity of models is likely to affect the prob-ability and speed of social transmission oftraits, as well as the patterning of the spread


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of the trait through the group. Ecologicalvariables of interest include food abundance,distribution, and diversity. Variables such asfood abundance and distribution influence as-pects of primates’ social relationships, such astolerance, dominance style, and propensity tocooperate (van Schaik 1989, 1996). The de-gree of risk in the environment (e.g., preva-lence of exposure to toxins, predators, anddangerous conspecifics) is likely to affect indi-viduals’ levels of caution in testing new foodsor interacting with unknown animals, whichin turn could affect their abilities and propen-sities to learn about novel foods or animalsvia asocial means (trial-and-error learning).Another potentially important influence ongroup dynamics that has not so often beenconsidered is the role of individual personali-ties, and particularly the personalities of long-term alpha males and females, who set thetone for how tolerant group dynamics can be,in general. To give an example, the succes-sive alpha males of one of my capuchin groupsconsistently engage in separating interven-tions, punishing attempts by other males toassociate with one another. In contrast, themales of some of my other groups are toler-ant of other males’ affiliations, such that thesemales are free to associate and cooperate fre-quently, and the group has a more relaxed so-cial atmosphere overall. Such influences couldpersist even after the death or emigration ofthe animal(s) whose personality created thenorm, as in Sapolsky & Share’s (2004) docu-mentation of a founder’s effect on a culture ofpeacefulness in baboons.

Psychological attributes include at leasttwo types of phenomena: (a) attitudes (e.g.,emotional responses to particular types ofsituations) and (b) mental representations oftasks. The first, attitudes, is of primary interestto socioecologists interested in determininghow external variables such as those discussedabove translate into patterns of behavioral re-sponses because some attitudes are more con-ducive to social learning than others. Exam-ples of attitudes are tolerance, neophobia, andemotional engagement (high interest level in

an object, task, or individual animal). Toler-ance has been hypothesized to enhance theprobability of social learning (Coussi-Korbel& Fragaszy 1995, van Schaik et al. 1999).Neophobia decreases the odds that an ani-mal will learn about a new food by trial-and-error learning. Strong expression of emotionby models toward the objects with which theyare engaged increases the salience of objectsto the observer such that observers are morelikely to explore the objects (Fragaszy et al.2004). And high emotional engagement be-tween participants seems to be a critical fea-ture of traditional bond-testing rituals in ca-puchins (Perry et al. 2003a) and interactionrituals in humans (Collins 1993). It seemsclear that emotions add salience to learningsituations, focusing the observer’s attention onparticular aspects of its environment. Beingsensitive to the emotional states and gaze di-rections of others enables joint attention ofthe model and the observer on the same as-pects of an activity pattern, thereby enhancingthe probability that some aspect of the model’sknowledge and/or behavior will be acquiredby the observer.

The second type of psychological at-tribute, mental representations, refers to theanimal’s conceptualization of (a) a particu-lar object’s properties, (b) a particular courseof action, (c) a cause-and-effect relationshipbetween two objects or individuals, or (d ) aparticular type of social interaction. For oneindividual to learn socially from observing ademonstrator’s actions, the observer will haveto produce a mental representation that is atleast partially overlapping with the demon-strator’s mental representation of the task, al-though the degree of overlap may be veryslight (Richerson & Boyd 2004). It is this par-ticular black box that most concerns thosecultural primatologists who do experimentalwork to discern the precise social learningmechanisms employed by primate species inspecific contexts. Of course, it is quite diffi-cult to know what mental representations arepresent in animals’ minds because the onlyclues we have are their behavioral outputs.

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Typically when one primate observes anotherperforming some task and learns socially fromthese observations, it does not exhibit a per-fect replication of what it has observed, whichimplies that the primate is not conceptualiz-ing the task in exactly the same way as themodel. The aspects of a behavioral sequencethat seem salient to one observer may not bethe same aspects that are salient to another.To give a hypothetical example, if three ani-mals were watching a fourth conspecific crackopen nuts with a stone and eat them, onemight learn from this observation that nutsare good to eat, the second might learn thatcombining uncracked nuts and stones pro-duces cracked nuts (without appreciating theimportance of the specific motor actions in-volved), and the third might learn that thespecific act of pounding yields food (but mayfail to understand why a stone is a partic-ularly good item to pound with). All threeof these observers might take the differentbits of information they acquired from watch-ing the model and then modify those men-tal representations with further informationacquired from subsequent trial-and-errorlearning, until they devised their own sub-tly different techniques for opening nuts. It isdevilishly difficult to know exactly how an ani-mal conceptualizes its understanding of a par-ticular task and the exact extent to which oneanimal’s mental representation overlaps withanother animal’s. But it is important to under-stand that even if two animals have only partialoverlap in their mental representations, theycan converge on similar or identical behaviorpatterns such that a tradition is formed.

Finally let us consider the behavioral pat-terns that cultural primatologists are trying toexplain: traditions and social norms. I define atradition as a relatively long-lasting behaviorpattern shared by multiple practitioners ow-ing to some form of social learning. Most ofthe traditions that have been studied by cul-tural primatologists consist of readily identi-fiable, discrete behaviors (e.g., quirky forag-ing techniques such as nut cracking or potatowashing) that can be reliably coded as present

or absent in a particular individual. Foragingtechniques, and material culture in particu-lar, have been a focus of study both in hu-mans and nonhumans because of the ease withwhich such traits can be identified. But thereis another whole type of behavioral patterns,which consists of group-typical responses toparticular social situations, that can perhapsbe defined best as social norms. Such behav-ior patterns would technically fit my defini-tion of tradition because of the role of sociallearning in their establishment, but I distin-guish them from traditions merely becausethey are harder to identify and operational-ize. Social norms are patterns of social be-havior that typify particular groups and arevariable between groups, but which each in-dividual within a group can follow to varyingdegrees, such that it is difficult to classify anyparticular individual as possessing or lackingthe trait. The acquisition of social norms isdirectly influenced by social interaction withothers but not necessarily by copying what hasbeen observed: The process of internalizing anorm could come about by taking a comple-mentary role in an interaction rather than byadopting the same role as the model. Althoughthe term social norm would imply punishmentfor violation of a social rule to many scholars,that condition is not a necessary part of mydefinition for the purposes of this discussion.

A well-documented human example of asocial norm is the culture of honor in thesouthern United States, in which southernersare far more polite than northerners in gen-eral but also are far more likely to respondto insults with extreme physiological stressresponses and impulsive violence (Nisbett &Cohen 1996). Some nonhuman primate ex-amples of social norms would be peaceful-ness in the forest baboon troop of Masai Mara(Sapolsky & Share 2004) and matrilineal in-heritance of dominance rank in macaques,in which a monkey socially inherits the rankjust below her mother and just above theolder sister that is closest to her in domi-nance rank (Chapais 1988). Some primatolo-gists have produced experimental evidence for


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social norms. For example, rhesus macaques(Macaca mulatta) rarely reconcile, yet when deWaal & Johanowicz (1993) housed rhesus ju-veniles with juvenile stumptail macaques (M.arctoides), which reconcile far more frequently,the young rhesus developed a stumptail-likereconciliation rate that persisted after the rhe-sus were placed back in all-rhesus groups. Yetthe rhesus juveniles did not acquire species-typical stumptail reconciliation gestures, sug-gesting that social attitudes rather than motorpatterns were transmitted between individu-als of the two species. Also, Kummer (1996)found that female hamadryas baboons (whichare accustomed to exclusive relationships witha single male) can quickly adapt to the dif-ferent consort style of savanna baboons (inwhich male-female sexual relationships aremore fluid and transient and males are lesscontrolling) and vice versa when females fromone species are transplanted into groups of theother species.

Further possibilities for cognitive rules inprimates in which the role of social learningmay prove to be influential are the acquisitionof decision rules determining which animal toside with in a coalition (Perry et al. 2004) orthe tendency to punish certain behaviors. So-cial norms are slowly established over time, asmaturing animals or immigrants adjust theirbehaviors in accordance with the behaviorpatterns exhibited by the majority of animals(or, perhaps, with the behavior patterns exhib-ited by particular influential animals). Becausean individual is unlikely to conform to thenorm in 100% of its opportunities to do so, it

is difficult to say with certainty when any par-ticular animal has internalized a social norm.Likewise, it is hard to say at a group-wide levelwhen a social norm has been clearly estab-lished. For these methodological reasons, andfor the reason that vast amounts of behavioraldata are necessary to document changes in so-cial norms over the history of a group or inthe course of an individual’s ontogeny, veryfew primatologists have devoted research ef-fort to studying social norms. Nonetheless,they are a fascinating topic of inquiry andone that is likely to shed light on the originsof human culture (de Waal 2004). Althoughmost of the empirical anthropological workon the dynamics of human culture changehas also focused on material culture andsubsistence technology (Richerson & Boyd2004), perhaps for the same methodologi-cal reasons that have created this researchbias in cultural primatology, the most pro-found and subtle aspects of cultural variationclearly center around differences in status re-lations and appropriate means of negotiatingsocial relationships in a complex society. Al-though much human relationship negotia-tion employs symbolic communication in theform of language, many aspects of humansocial norms consist primarily of gestures,postures, and appropriate behavioral coordi-nation in space and time (for example, culture-specific ways of greeting a socially dominantindividual or of resolving a conflict). Socialnorms of these types may prove not to beso different from social norms in nonhumanprimates.

ACKNOWLEDGMENTS

Thanks are given to the Max Planck Institute for Evolutionary Anthropology for funding mewhile I wrote this review. I also thank Filippo Aureli, Josep Call, Richard Lesure, and MikeTomasello for helpful discussions, and Joseph Manson for comments on the manuscript.

LITERATURE CITED

Atran S. 1990. Cognitive Foundations of Natural History: Towards an Anthropology of Science.Cambridge, UK: Cambridge Univ. Press. 360 pp.

Aunger R. 2000. Darwinizing Culture: The Status of Memetics as a Science. Oxford, UK: OxfordUniv. Press. 242 pp.

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Contents ARI 13 August 2006 13:30

Annual Review ofAnthropology

Volume 35, 2006

Contents

Prefatory Chapter

On the Resilience of Anthropological ArchaeologyKent V. Flannery � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � 1

Archaeology

Archaeology of Overshoot and CollapseJoseph A. Tainter � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � �59

Archaeology and Texts: Subservience or EnlightenmentJohn Moreland � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � 135

Alcohol: Anthropological/Archaeological PerspectivesMichael Dietler � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � 229

Early Mainland Southeast Asian Landscapes in the FirstMillennium a.d.

Miriam T. Stark � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � 407

The Maya CodicesGabrielle Vail � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � 497

Biological Anthropology

What Cultural Primatology Can Tell Anthropologists about theEvolution of CultureSusan E. Perry � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � 171

Diet in Early Homo: A Review of the Evidence and a New Model ofAdaptive VersatilityPeter S. Ungar, Frederick E. Grine, and Mark F. Teaford � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � 209

Obesity in Biocultural PerspectiveStanley J. Ulijaszek and Hayley Lofink � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � 337

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Evolution of the Size and Functional Areas of the Human BrainP. Thomas Schoenemann � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � 379

Linguistics and Communicative Practices

Mayan Historical Linguistics and Epigraphy: A New SynthesisSøren Wichmann � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � 279

Environmental DiscoursesPeter Muhlhausler and Adrian Peace � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � 457

Old Wine, New Ethnographic LexicographyMichael Silverstein � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � 481

International Anthropology and Regional Studies

The Ethnography of FinlandJukka Siikala � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � 153

Sociocultural Anthropology

The Anthropology of MoneyBill Maurer � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � �15

Food and GlobalizationLynne Phillips � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � �37

The Research Program of Historical EcologyWilliam Balée � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � �75

Anthropology and International LawSally Engle Merry � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � �99

Institutional Failure in Resource ManagementJames M. Acheson � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � 117

Indigenous People and Environmental PoliticsMichael R. Dove � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � 191

Parks and Peoples: The Social Impact of Protected AreasPaige West, James Igoe, and Dan Brockington � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � 251

Sovereignty RevisitedThomas Blom Hansen and Finn Stepputat � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � 295

Local Knowledge and Memory in Biodiversity ConservationVirginia D. Nazarea � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � 317

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Food and MemoryJon D. Holtzman � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � 361

Creolization and Its DiscontentsStephan Palmié � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � 433

Persistent Hunger: Perspectives on Vulnerability, Famine, and FoodSecurity in Sub-Saharan AfricaMamadou Baro and Tara F. Deubel � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � 521

Theme 1: Environmental Conservation

Archaeology of Overshoot and CollapseJoseph A. Tainter � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � �59

The Research Program of Historical EcologyWilliam Balée � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � �75

Institutional Failure in Resource ManagementJames M. Acheson � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � 117

Indigenous People and Environmental PoliticsMichael R. Dove � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � 191

Parks and Peoples: The Social Impact of Protected AreasPaige West, James Igoe, and Dan Brockington � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � 251

Local Knowledge and Memory in Biodiversity ConservationVirginia D. Nazarea � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � 317

Environmental DiscoursesPeter Mühlhäusler and Adrian Peace � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � 457

Theme 2: Food

Food and GlobalizationLynne Phillips � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � �37

Diet in Early Homo: A Review of the Evidence and a New Model ofAdaptive VersatilityPeter S. Ungar, Frederick E. Grine, and Mark F. Teaford � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � 209

Alcohol: Anthropological/Archaeological PerspectivesMichael Dietler � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � 229

Obesity in Biocultural PerspectiveStanley J. Ulijaszek and Hayley Lofink � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � 337

Food and MemoryJon D. Holtzman � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � 361

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Old Wine, New Ethnographic LexicographyMichael Silverstein � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � 481

Persistent Hunger: Perspectives on Vulnerability, Famine, and FoodSecurity in Sub-Saharan AfricaMamadou Baro and Tara F. Deubel � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � 521

Indexes

Subject Index � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � 539

Cumulative Index of Contributing Authors, Volumes 27–35 � � � � � � � � � � � � � � � � � � � � � � � � � � � 553

Cumulative Index of Chapter Titles, Volumes 27–35 � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � � 556

Errata

An online log of corrections to Annual Review of Anthropology chapters (if any, 1997 tothe present) may be found at http://anthro.annualreviews.org/errata.shtml

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Documents

What Cultural Primatology Can Tell Anthropologists about ...users.clas.ufl.edu/...CulturalPrimatology_ARA_2006.pdf · that cultural primatology has made in deﬁn-ing how cultural