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Evolution and the social sciences

Robin I.M. Dunbar History of the Human Sciences 2007 20: 29 DOI: 10.1177/0952695107076197 The online version of this article can be found at: http://hhs.sagepub.com/content/20/2/29

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H I S T O RY O F T H E H U M A N S C I E N C E S
[20:2; 2950; DOI: 10.1177/0952695107076197]

Vo l . 2 0 N o . 2
pp. 2950

2007 SAGE Publications (Los Angeles, London, New Delhi and Singapore)

Evolution and the social sciences

ROBIN I. M. DUNBAR

ABSTRACT When the social sciences parted company from evolutionary biology almost exactly a century ago, they did so at a time when evolutionary biology was still very much in its infancy and many key issues were unresolved. As a result, the social sciences took away with them an understanding of evolution that was in fact based on 18th- rather than 19th-century biology. I argue that contemporary evolutionary thinking has much more to offer the social sciences than most people have assumed. Contemporary evolutionary research on human behaviour focuses on two main issues at the micro-social scale: understanding the trade-offs in individual decision-making and understanding the cognitive constraints that limit exibility of decisions. I offer examples of both of these approaches. Finally, I consider the broader question of the macro-social scale. Key words behavioural decisions, cognitive constraints, Darwinism, evolution, naturenurture

It would probably be an understatement to say that evolution has not had a particularly good press within the social sciences: it has not done so for the better part of a century. Yet, in some ways, this is surprising. The founding fathers of modern sociology were much inspired by evolutionary ideas, and scenarios for the evolution of society were a major component of both anthropology and sociology during the closing decades of the 19th century.

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Indeed, Herbert Spencer, historically one of the more inuential founders of modern sociology, was a committed advocate of Darwinian ideas. However, during the early years of the 20th century, a radical move against biology began within the social sciences. Origin stories were gradually banished as topics of serious discussion and, under the inuence of Durkheim and Boas, the assumption that any aspect of human behaviour (and, in particular, social behaviour) could have a biological basis was dismissed as untenable. In my view, this was unfortunate, albeit understandable when seen from the perspective of the times. It was unfortunate for two reasons. First, it predisposed the social sciences to ignore potentially valuable ideas that were later to emerge within evolutionary biology. Second, it was particularly unfortunate because of its timing. In this respect, it is important to appreciate that evolutionary ideas within biology were still in great ux at the end of the 19th century when the rift between the social and biological sciences rst opened up. The second half of the 19th century had witnessed the grand battle between the two major schools of evolutionary thinking, those associated with Lamarck (largely reecting an 18th-century perspective) and with Darwin (reecting radically new ideas that emerged in the middle of the 19th century). While both agreed that evolution occurred, they differed fundamentally about the mechanisms involved, and the implications that followed. Most of this history (which I summarize below) is well known and a detailed account with citations would be inappropriate and out of place in the present context. Rather, my purpose here is simply to sketch in this background so as to provide a better understanding of how and why the contemporary situation came to be. For those requiring an authoritative account of the recent history of biology, the best source is undoubtedly Mayr (1982). The older school, followers of the great French biologist Jean-Baptiste Lamarck, viewed the processes of evolution as the natural outcome of the capacities inherent in the organism. They generally adhered to the venerable Aristotelian concept of the Great Chain of Being whereby species gradually and inevitably evolved from one form into another along a continuum of increasing complexity. Lamarck famously added to this his concept of the evolution of acquired characters a form of biological tinkering whereby modest but nonetheless striking changes in physical form might be introduced along the way through the unusually heavy use of some part of the body (giraffes necks and blacksmiths arms being the conventional examples). Excess use leading to the overemphasis of some body-part would be transmitted down the generations because of resulting changes in the biological nature that underpinned inheritance. Nonetheless, acquired characters notwithstanding, the fundamental processes of evolution remained the natural progression along the Great Chain. Humans, as the most complex beings known, were naturally higher up the chain (whose pinnacle traditionally lay with God), and that, by denition, meant they must be one of the oldest:

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implicitly or explicitly, the processes of evolution could not be speeded up, so a species age was necessarily correlated with its level of advance along the Great Chain. Evolution was, by its nature, progressive. Darwin upset this particular apple cart by arguing that evolution was a random process with no xed direction. Novel characters emerged by chance, and were promoted or removed by natural selection acting as a kind of environmental lter. He had come to these views on two major grounds. One was the discovery that life forms on different continents did seem not to follow the same Lamarckian pattern up the Great Chain; indeed, the life forms on different continents could be radically different. The Beagle voyage played a seminal role in this, of course, but Darwin was by no means the only biologist to notice this as a consequence of the great age of Victorian exploration: as is well known, Alfred Russel Wallace had come to similar conclusions, and he was far from being alone in this. The second piece of formative evidence underpinning Darwins view was pigeon-breeding. Darwins interactions with professional breeders, and his own experiments, taught him that careful selective breeding could result in dramatic changes in an animals form in just a few generations. The concept of natural selection was conceived as an analogy to the breeders art. Once this was understood, two important consequences followed. First, evolution could not be directional: there was no one-directional progressive tramline on which every species was forced to run. Second, as a direct consequence, an organisms complexity tells us nothing about its age, merely about the intensity of the selection pressures that it has been under. Unfortunately for the social sciences, the battlelines between these two opposing conceptions of how evolution worked remained rmly drawn until well into the early decades of the 20th century. Only with the development of the so-called synthetic theory of evolution (sometimes also known as NeoDarwinism) in the 1930s was the Lamarckian ghost nally laid to rest. At the time that the social sciences parted company from biology, the Lamarckian view still held wide sway, not only within biology but especially outside the discipline. Indeed, many people were unable to distinguish between the two. One of those was Spencer himself, whose enthusiastic promotion of Darwinian ideas (Spencer was in fact responsible for coining the Darwinian catchphrase survival of the ttest) involved an amalgam of Darwins and Lamarcks theories. Social Darwinism (as Spencers main contribution came to be known) was not, in actual fact, Darwinian at all, but rather Lamarckian (which is largely why Darwin himself distanced himself from it, much to Spencers eternal disappointment). This was no great fault on Spencers part, because many people Darwin included successfully managed to hold ideas from both traditions comfortably at the same time. In Darwins case, this was the idea of acquired characters: he never managed to provide a convincing mechanism of inheritance for his theory, and was, in the later editions of the

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Origin, forced back onto Lamarckian mechanisms1 though not onto Lamarcks theories of evolutionary progress. For Spencer, it was the idea of progressive evolution: his view was very much one of a natural unfolding of social complexity as a part of a natural developmental sequence and shared much in common with the ideas of the German biologist Ernst Haekel who interpreted Darwin in exactly this way. Unfortunately, by the time the biologists had nally sorted out what was happening to their own satisfaction, the rift with the social sciences was well entrenched, and the message never got across. Progressive evolution continued to be a major foundation for understanding sociality within the social sciences for the better part of half a century after it had been discredited within biology. More importantly, when evolutionary ideas were eventually rmly ejected from the social sciences once and for all, it continued to colour sociologists perceptions and understanding of evolutionary biology. There was, of course, a second issue separating the social and biological sciences during the latter half of the 20th century that derived in part from these earlier discussions, and this was the issue of genetic determinism. For biologists, the discovery of genes (and later DNA) provided a fundamental platform for understanding evolutionary processes. Evolution occurred only because genes were inherited by offspring from their parents, thereby maintaining the integrity of the so-called germ line that was necessary to allow evolution to occur in the Darwinian manner. Rare mutations provided the grist for the mill of natural selection, but what later became known as Weizmanns Doctrine or biologys Fundamental Theorem (that genes inuence the body, but the body [i.e. use in the Lamarckian sense] cannot change the genes themselves2) remained the cornerstone that made evolution possible. It is easy enough to understand why genes came to play such an important role in biological thinking. Most biologists were, and to a large extent still are, concerned with the physical attributes of the organism: how did its bodyparts come to be, how did species (dened mainly by differences in these body-parts) come to be? And for these, genes play an important and substantive role (even if, as has been recognized in the so-called interactionist stance since the 1960s, the genes are not independent of the environment in which the organism develops). The discovery of genes, and later DNA, inevitably came to play an important role as the focus of biologists interest. However, it is easy to forget that genes as such played no role at all in either Darwins theory of evolution or, indeed, Mendels theory of inheritance. To be sure, both argued for a mechanism that allowed traits to be passed down the generations in a reasonably (that is to say, statistically) reliable way, but both remained wholly ignorant of the biochemical properties of genes as we know them today: these came to be understood, at rst rather dimly, only with the opening decades of the 20th century getting on for a quarter of a century after these two particular giants had died.

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In fact, Mendels theory of inheritance (and hence Neo-Darwinism itself) rests only on what Mendel referred to as the delity of copying between parents and offspring. Mendels entire theory breathed not a word about genes, but merely asserted that something (which he called a factor) allowed offspring to resemble their parents in respect of individual traits or characters. It was Mendels genius to gure out a mathematical model of how this might work at the genetic level that has more than stood the test of time. In the excitement of the later discovery of DNA, the signicance of all this was often lost sight of. It was not until the 1980s that evolutionary biologists began to point out that, in actual fact, learning itself is a perfectly good Darwinian process, and whatever mechanisms might underpin this (the way animals learn and what they may be predisposed to learn) would make just as good a basis for Darwinian evolution as anything else. Thus was born the notion of cultural evolution as a topic of study for evolutionary biologists, and, more notoriously perhaps, Dawkinss concept of a meme.3 Having set the broad historical context, let me now briey summarize the three main evolutionary approaches to the study of human behaviour, and then provide some examples of these approaches.

EVOLUTIONARY APPROACHES It is important to be clear at the outset that evolutionary approaches to the study of behaviour do not normally concern themselves with evolutionary history that is to say, the evolutionary origins of behaviour and the sequence of changes by which behaviour evolves sequentially from one species to another. Rather, the focus lies mainly on the micro-evolutionary processes underpinning the mechanisms of selection and, in the case of behaviour, the cognitive mechanisms involved. By the same token, there has been very limited interest in the historical aspects of the evolution of societies after the 1960s (although this may now be changing). By convention, there are three broad approaches to the evolutionary study of behaviour. The oldest of these is that of the behavioural ecologists; this approach has its origins in the study of animal behaviour (usually now referred to as behavioural ecology) and is largely the remit of evolutionarily minded social anthropologists whose main focus of study is traditional societies (who typically refer to themselves as evolutionary anthropologists). The second approach is that associated with what was previously referred to as Darwinian Psychology, but is now more commonly referred to as Evolutionary Psychology in the narrow sense;4 its focus of interest is principally on cognitive mechanisms (the design of the mind) and is largely the province of scientists whose intellectual background lies within psychology (and in particular cognitive and social psychology). The third, and much the smallest,

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branch is the study of cultural evolution, sometimes (but by no means always) associated with the concept of the meme. This is mainly the province of evolutionary biologists interested in the population-level dynamics of geneculture co-evolution; as a result, it is very heavy on mathematical modelling, but is often also concerned with the psychological processes of learning that underpin cultural transmission. Because it is a rather arcane area, I shall say no more about this third approach. However, broad summaries of these positions and their articulation can be found in Barrett et al. (2002), Laland and Brown (2002), Dunbar et al. (2005) and in the various chapters in Dunbar and Barrett (2007). The two main approaches, those associated with behavioural ecology and with Evolutionary Psychology sensu stricto, differ both in the foci of their interests and in philosophy. Behavioural ecologists tend to be interested mainly in the extent to which an individuals behaviour maximizes its genetic tness. Genetic tness is a technical measure of an individuals relative contribution to its species gene pool. Since it is a relative, and not an absolute, measure such studies always involve comparisons between individuals to determine whether those individuals that possess a trait (or act in a particular way) have proportionately higher tness than those that do not. Since tness is technically measured several generations into the future, it is conventional within evolutionary studies to use one of a range of proxies. These include lifetime reproductive success (i.e. total number of offspring born or surviving to a given age), number of matings within a more limited time period (perhaps, a single breeding season), or some suitable index associated with the particular trait in question (e.g. energy returns for foraging strategies). The choice of metric is determined by the nature of the problem in hand. The link between the chosen metric and tness is simply covered by what is usually known as the phenotypic gambit (Grafen, 1984), which assumes that, all else equal, there will be a direct one-to-one translation between these proxy indices and actual tness on the average. The broad assumption is that natural selection will hone the cognitive machinery and basic physiology and anatomy so that the organism will choose those behavioural options available to it in such a way as to maximize tness. This is a strictly empirical question, but the assumption that individuals naturally act so as to maximize tness provides a powerful heuristic tool for the study of behaviour because it makes very specic predictions that can be tested against empirical data. Evolutionary theory has used the mathematics of economics (but without the conceptual baggage of economics5) to develop a very sophisticated body of quantitative theory. The precision of many of the predictions that can be made from theory means that, if the data do not match the predictions of theory, then we can ask whether this is because we have overlooked some crucial component in the organisms biology or because the hypothesis (or the theory on which it is based) is wrong. Often the form

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of the difference between observed and expected patterns will cue us in to what we have overlooked. It is important to appreciate that costing out the tness consequences of behaviour involves assessing the net benet (i.e. the benets minus the costs) since all actions incur costs. Biological and behaviour phenomena often exhibit an inverted-U-shaped pattern of net benets: doing something that is generally speaking good for you too often incurs costs that increase exponentially such that the costs rapidly wipe out whatever benets are gained. A widely conrmed principle in behavioural ecology known as Lacks Principle states that the optimal number of offspring (or, in its original formulation for birds, eggs laid) will invariably be well below the physiological maximum: if the costs of rearing additional offspring increase with their number, large broods will suffer disproportionately high mortality and the net number of surviving offspring will often be lower than those obtained by more restrained parents (Lack, 1968). More is not always better in evolution. This is because, to borrow a phrase from the eminent evolutionary biologist John Maynard Smith, evolution is not interested in babies as such, but rather in (great-)grandchildren (i.e. tness). A second important lesson is that everything in evolutionary biology is context-dependent: there are no absolutes in biology, no traits or behavioural strategies that are absolutely better in all circumstances (which we might therefore expect to occur universally). Indeed, behaviour in particular is so context-dependent that high- and low-ranking individuals in the same social group may actually have radically different optimal strategies because their positions in the social hierarchy within the group inuence the costs or benets, or both, in different ways (for this specic example, see Cheney, 1983). In short, behavioural ecologists typically assume that behaviour is currently adaptive (i.e. that organisms are sufciently exible to ne-tune their behaviour quite subtly to even small changes in the costs and benets that they encounter) and thus interest themselves in the differences between individuals (or, more generally, between species). The Evolutionary Psychology approach, in contrast, tends to focus more explicitly on human universals. Its underlying assumption is that there has not been signicant evolution in the human brain within the last 100,000 years or so, but cultural evolution since the Agricultural Revolution 10,000 years ago has radically changed the environment within which humans make their behavioural choices. Since it is assumed that the human mind (i.e. brain) has been adapted to conditions that existed at some time in the past, this may lead to a signicant disjunct between actual behaviour and what is now optimal in the novel circumstances in which we nd ourselves. Our contemporary behaviour need not, and often may not, be adaptive. In the limit, they would argue that the behavioural ecologists attempt to determine whether or not behaviour is adaptive is misguided: contemporary behaviour can tell us nothing about how or why behaviour (or the cognition that underpins it)

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is adaptive. Although such a hardline conception of the issue is probably an increasingly minority view, it nonetheless reminds us that the principal focus of the Evolutionary Psychologists lies in understanding the extent to which human behaviour (in particular) is guided by the particular design of the mind that we have inherited. Underlying this is, however, a view of human behaviour that is rather more deterministic than the behavioural ecologists would allow: the human mind is designed to facilitate certain kinds of behaviour, because exactly those kinds of behaviour (or attitudes) were adaptive in the environment in which humans evolved (some time in the late Pleistocene prior to the Agricultural Revolution). In contrast to the tness-matching methodology of the behavioural ecologists, the Evolutionary Psychologists methodology relies rather more heavily in demonstrating tness for purpose based on the design characteristics of either specic cognitive mechanisms or the behaviour produced by those mechanisms. In this respect, the two sub-disciplines focus on alternative denitions that evolutionary biologists conventionally use to determine the adaptiveness of biological phenomena. Biologists have always insisted that evidence for adaptation can be provided either by demonstrating tness consequences (a relativistic measure) or by demonstrating engineering design for purpose (an absolute measure) (Williams, 1966). The eye, for example, provides a classic example of design for purpose (an adaptation): we can examine its properties and show that they do indeed produce the supposed effect (a procedure sometimes known in evolutionary biology as reverse engineering). Antler size in deer would provide an example of the alternative approach: we would correlate antler size with number of matings achieved during a breeding season, and infer a selective advantage (and hence adaptedness) from the presence of a correlation. Having sketched out the basics of the philosophical assumptions in the two main evolutionary approaches to the study of (human) behaviour, I want to emphasize that, despite the acrimony that has sometimes existed between them (for a recent summary, see Laland and Brown, 2002), these are in fact two ends of a continuum (or, strictly speaking, maybe it should be three corners of a triangle). They both represent perfectly good ways of doing evolutionary biology. Their difference lies simply in the questions to which they give priority. The behavioural ecologists are interested primarily in the processes that drive evolutionary change and, once such change has occurred, keep it in place; they are interested in individual differences between and within populations of individuals. In contrast, the Evolutionary Psychologists sensu stricto focus on the nature of the adaptation itself; they are interested in the universal features that characterize groups of organisms (usually species). They are asking, and seeking to answer, questions at different levels of the explanatory framework. However, the lines can become blurred, and many of those who study human behaviour from an evolutionary point of view

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mix both perspectives, depending on the questions they happen to be asking. In the following section, I give some examples of these two approaches, but I choose these examples specically to point out that the same researchers can work at both levels.

CASE STUDIES FROM THE BEHAVIOURAL ECOLOGY APPROACH There have been studies in the behavioural ecology tradition of both traditional societies and contemporary modern societies. A range of such studies can be found in Barrett et al. (2002). I will describe just two examples here from my own work. One concerns mate choice strategies, based on analyses of personal ads. The second concerns parental investment decisions, with a particular focus on a Gypsy (or Roma) population which has a more ethnographic avour. Choosing the right mate can have signicant consequences for an individuals tness in a number of different respects. As a general principle, organisms can choose between investing in matings (in order to maximize the number of conceptions achieved in a lifetime) or in parental care (in order to maximize the number of offspring successfully reared to adulthood). Since evolution is ultimately interested in the number of grandchildren, or better still great-grandchildren (offspring that do not in their turn reproduce do not contribute to ones genetic tness), there is an inevitable trade-off between these two options, and hence, in principle at least, a more or less innite number of ways in which the two can be balanced against each other (this is, essentially, Lacks Principle again). The extent to which this is possible for any given species will ultimately depend on the balance of the costs and benets of these two strategic options. Because the costs of rearing offspring are both heavy and prolonged in humans (mainly as a consequence of our very large brains), the demands of parental care have to be met if children are to be reared successfully. However, within that constraint, it is possible still to opt for a strategy that maximizes matings at the expense of rearing, providing the mate is prepared to carry a disproportionate share of the rearing burden. Because of the way mammalian reproductive biology is organized (an unavoidable given for mammals), there is, in the general mammalian case, an inevitable asymmetry in the way the two sexes experience these effects. All else equal, male mammals can make no or only a very limited direct contribution to the business of rearing (through parental investment), at least in the early phases (i.e. up to the point of weaning), and maximizing the number of matings provides them with the most effective means of maximizing the number of offspring produced. In contrast, because females are committed willy-nilly to a signicant investment, they will gain less by maximizing the number of matings per se and

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proportionately more by ensuring that (1) the male(s) who father their offspring provide the best genetic contribution to their offspring and (2) the circumstances in which they have to rear the subsequent offspring are optimal (in terms of the provision of the resources required for successful rearing and that does not necessarily involve any contributions from the father). Thus, there is in principle a natural contrast in the two sexes reproductive strategies. While this contrast is naturally moderated in the case of humans by the fact that the costs of rearing are very heavy (thanks mainly to the need to grow a very large brain), humans nonetheless remain exposed to the same dichotomy and we can expect this to be reected in the mate choice strategies of the two sexes: males can be expected to attempt to maximize matings whenever possible, and will be less discriminating as a result in their choice of partners (other than having a preference for fertile partners), whereas women will be more discriminating (i.e. choosy) and will place greater weight either on the genetic quality of males or the males abilities to contribute (directly or indirectly) to the business of rearing, or on both. The latter dichotomy itself reminds us that there is an opportunity to trade one dimension off against another, and we may expect, again on general evolutionary principles, that womens mate choice preferences will have created a selection environment in which males might come in two general types (sometimes referred to in the evolutionary literature as cads and dads: Draper and Harpending, 1982; Cashdan, 1996). However, since evolutionary processes both are dynamic and depend on the contextual contingencies of the balance between costs and benets, we can expect that, while these general principles hold good, individuals may nonetheless trade off their opportunities on the different criteria. This will be especially true of males, because female mate choice is more complex and involves more dimensions, making it harder for women to identify an optimal (i.e. perfect) mate. That will inevitably leave males the option of ne-tuning their behaviour to target one dimension at the expense of another. All this is standard behavioural ecology theory, and can be found in any appropriate textbook. Personal ads provide a very convenient way of studying mate choice strategies in humans, not least because they provide a succinct summary both of the advertisers expectations and of what they have to offer. More importantly, they do so in a form that can be easily quantied, either by counting the number of advertisements that exhibit a particular trait or by counting the number of words an advertiser uses within a particular category (thereby giving a simple way of quantifying the advertisers inherent interest or emphasis on that category). We have been able to show (Waynforth and Dunbar, 1995; Pawowski and Dunbar, 1999a, 1999b, 2001) that (1) women are much more choosy than men in what they demand in a prospective partner (a nding conrmed by a wide range of other psychological and

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sociological research on mate choice: for a summary, see Grammer, 1989), (2) both sexes target their demands on those traits that have most impact on their tness (fertility in the case of men, resources/status and parental care capacities in the case of women) and (3) both sexes adjust their demands in the light of what they themselves have to offer (i.e. those with weaker bargaining hands, as dened by the interests of the other sex, tone down their own demands in order to remain competitive). More particularly, Pawowski and Dunbar (1999b) were able to demonstrate that advertisers perception of where they stand in the mating market was, in general, extremely sensitive (by and large, they really did understand how strong a bargaining hand they held) and directly inuenced how demanding they were: those with less to offer demanded less. Men who lacked economic resources to bring into a relationship, for example, were less demanding in what they expected of a partner or emphasized alternative attributes (e.g. willingness to tolerate and invest in children from a previous relationship) that might help to offset their initial disadvantage. Once a mate has been chosen, of course, the business of reproduction is the core to the evolutionary process, although a caveat needs to be added as a reminder: it is not necessary to reproduce oneself in order to maximize ones tness. What many consider to be the fundamental theorem of evolutionary biology (conventionally known as Hamiltons Rule: see Barrett et al., 2002) reminds us that, when the issue is the replication of genes in future generations, there are in fact two routes by which this can be achieved: one is by direct personal reproduction and the other is through the reproduction of relatives who share the same gene(s). Hamiltons Rule formally provides one explanation as to how altruistic behaviour6 could evolve in a Darwinian world. However, it stands as a general reminder of two important points. First, everything we do has implications for our tness, not least because any action may affect the interests of our relatives as well as our own individual interests. Second, whenever we decide to do something, we are always choosing between at least two alternative courses of action: even in the most trivial case of my deciding to give money to a beggar in the street, my doing so means both (1) that money will not be available later for me to use for my own purposes and (2) it will not be available for me to invest in my relatives (which would allow me to benet through kin selection). In short, there are always consequences for every action, although in some cases those consequences may be quite trivial. It is the balance between the net benets (benets minus costs) of these alternatives that lies at the root of the evolutionary process. In this particular context, the issue is the choices that parents make between investing in their various offspring. One implication of Hamiltons Rule is that different individuals (including different offspring) will inevitably vary in their tness value (for a good if technically difcult summary of this issue

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and its implications for human social structure, see Hughes, 1988). Not only does their relatedness to me vary, but their tness value (to me) will also vary as a function of their age (because age affects the number of future offspring they can produce), their sex (because sex affects their reproductive riskiness), and any number of other variables. All these sum together to dene what biologists refer to as reproductive value (Fisher, 1930). This led Trivers and Willard (1973), in a seminal paper, to point out that, in order to maximize their own tness, parents may choose to invest differentially in their various offspring. This may include preferring one sex of offspring over another, or discriminating for and against offspring of different birth order. The key issue here is how investment in an individual offspring inuences that offsprings future reproductive opportunities. Trivers and Willard pointed out that, when the two sexes differ in the variance in lifetime reproductive output, then parents in good condition who can afford to invest heavily should prefer the more risky sex, whereas those in poor condition who cannot afford to invest so heavily should prefer the less risky sex. In mammals, the nature of the reproductive machinery will inevitably mean that the variance in male reproduction will usually exceed that of females, even though both must have the same mean (because that is set by the rate at which the reproductively more limited sex can reproduce). In mammals, females are limited in their reproductive turnover time, whereas males can compete to mate with many females (as a result of which some males will do very much better than average, whereas others may fail to reproduce at all). Translated into human terms, this leads to the expectation that, if investment in children directly inuences their reproductive success and does so differentially for the two sexes, wealthier parents should favour sons, whereas poorer parents should favour daughters. One consequence of this would be the expectation that inheritance of land and other forms of wealth will most often be male-biased, but only when sons can put such resources to better use than daughters by using them to acquire more wives (by which I mean to include not only formal polygamy, but also the acquisition of mistresses in formally monogamous societies). The Trivers-Willard Effect (as this is known) has been widely documented in animals, and a number of studies have attempted to explore its relevance to humans. Of these, the most complete7 has been that by Bereczkei and Dunbar (1997). They compared parental investment behaviour in two subgroups within contemporary Hungarian society, namely Gypsies (Roma) and ethnic Hungarians living in the same locations. As members of an economic underclass, the Gypsies represent a group whose opportunities are severely limited even in comparison with the lower-class ethnic Hungarians among whom they live. However, Gypsy women that married into the relatively wealthier Hungarian community had signicantly higher lifetime reproductive outputs than those who married into their own ethnic community, mainly because of the economic resources available in the Hungarian community. This classic

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form of hypergamy provides exactly the conditions for a Trivers-Willard Effect. Bereczkei and Dunbar (1997) were able to show, from analyses of detailed interviews with a large sample of women from both communities, that Gypsy parents invested more heavily in their daughters, whereas ethnic Hungarian parents invested more in their sons. This included differences in the length of the interbirth interval following a son versus a daughter (a measure of direct biological investment), abortion rates after sons versus daughters (higher after sons than after daughters among the Gypsies, but reversed in the ethnic Hungarians), the duration of lactation (it was up to 6 months longer for Gypsy girls than Gypsy boys, but the reverse was the case among the Hungarians) and even the duration of secondary schooling (which incurred a direct nancial cost for the parents, even under the Communist regime). More importantly, the strength of this bias varied across four communities (two Gypsy and two ethnic Hungarian) as a direct function of their capacity to benet from hypergamy. In other words, these investment decisions were not xed in some simpleminded sense, but were being titrated very carefully against the anticipated tness pay-offs. That this was so was demonstrated by showing a linearly and exactly proportional relationship between the ratio of investment in the two sexes across the four communities (rural and urban Gypsy8 communities, and rural and urban ethnic Hungarian communities) and the ratio of tness pay-offs indexed by the numbers of grandchildren produced through sons versus daughters.

CASE STUDIES FROM THE EVOLUTIONARY PSYCHOLOGY APPROACH My choice of examples to illustrate the Evolutionary Psychology approach is slightly unusual, in that most Evolutionary Psychologists would focus their interests on the way cognition limits the kinds of strategic decisions associated with mate choice or other everyday behaviours. There is, for example, now a very large literature on the way that evolution acting on the human mind inuences behaviours as diverse as disgust, leadership, mate choice, friendship patterns and altruistic behaviour, among others (for recent overviews, see Schaller et al., 2006; Forgas et al., in press). Instead, I will take a slightly larger perspective and use, as my core example of this general approach, the work we have done on the size and structure of human social networks. I do so mainly to show that the Evolutionary Psychology and behavioural ecology (evolutionary anthropology) approaches are not intellectually mutually exclusive, but rather reect an underlying unity of conceptual structure. We have been able to demonstrate that neocortex volume (essentially the thinking part of the brain) correlates directly with social group size across a

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wide range of primate species, and that this relationship predicts extremely closely the size of human social groups. The predicted size for human social groups is approximately 150 individuals, and this value turns out to be extremely common in a wide range of human social contexts. These range from the size of villages in traditional societies to the size of clans (or regional groups) among hunter-gatherers, the size at which formal hierarchical structures (line management, police forces) become necessary in both contemporary communities and business organization,9 the optimal size for church congregations and the size of personal social networks (as determined both by soliciting lists of friends and from small world experiments) (Dunbar, 1993, 2004). In addition, we have been able to show that human social networks are highly structured, such that they have the form of a series of concentric circles (the circles of acquaintanceship): each successive circle of acquaintanceship includes more individuals, but involves relationships of markedly reduced intimacy and frequency of contact (Hill and Dunbar, 2003; Zhou et al., 2005). More surprisingly, the size of these successive circles has a very characteristic ratio: each grouping is exactly three times larger than that contained within it. We are not sure why this pattern should be so consistent, but there are grounds for believing that there may be a cognitive limit on the number of relationships that can be held at a particular level of intimacy. Stiller and Dunbar (2007), for example, showed that the size of the innermost circle of acquaintanceship (on average 5) was a more or less linear function of individuals capacities to handle mind-reading tasks (i.e. the kinds of social cognition that is involved in theory of mind, or the ability to understand anothers mental states). The fact that humans are embedded in complex multi-level social systems has important implications for the more conventional Evolutionary Psychology approach, because it reminds us that forces other than simple self-interest may be at play when humans choose how to behave. We are embedded in social networks that exist for a purpose to enable us to achieve those goals of biological survival and reproduction as efciently as possible. Such socialities are, in effect, implicit social contracts (which may, of course, sometimes be made explicit with verbal agreements): they allow us to trade some loss of self-interest (we have to give up some personal interests to enable the group to survive as a cooperating community) in anticipation of much greater future gains via community-level processes. This recognition of the importance of these multi-level processes represents a new and exciting development in evolutionary biology (Plotkin and Odling-Smee, 1981; Laland et al., 2000) precisely because it allows us to draw attention once again to the community and societal levels at which individuals interact with each other. This, of course, is simply to remind ourselves of Hamiltons Rule. As social creatures, we are dependent on the effectiveness with which our local social

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structures work. Indeed, the evolutionary perspective would argue that societies exist precisely to facilitate our capacity to solve the everyday problems of survival and reproduction cooperatively. This is a generic primate characteristic: all primate societies are implicit social contracts of this form, and they involve a ne-tuned balancing of the demands of system stability against our own personal self-interests. Cheating on the social contract risks destabilizing the social system, thereby losing us its benets. This is the familiar freerider (or freeloader) phenomenon (see Dunbar, 1999, 2004). The pressure to cheat is always there, because it will always be in our individual shortterm interests to try to gain more from the social contract than we put in. This ne-tuned balancing of selsh versus group interests has turned up repeatedly in a range of experimental studies inspired by the evolutionary approach (Henrich et al., 2005; Madsen et al., in press). In all these studies, the social and economic context has a strong inuence on exactly how subjects respond to a particular challenge, just as the behavioural ecology perspective embedded as it is within evolutionary biology insists should be the case.

CONCLUDING REMARKS These issues become of particular interest from the macro-sociological scale because they emphasize the fact that humans are embedded in complex, multilevel social structures. The evolutionary perspective differs only insofar as it reminds us that, no matter how much we are socialized (in the Durkheimian sense) into the particular society that we belong to, nonetheless we do take personal decisions about how to behave towards each other. As a result, there is often a natural tension between what is in our short-term personal interest and what is in our longer-term interest via the communal life. The need to balance those two essentially incompatible demands creates the imperfections of social life with which we perennially have to grapple. Indeed, it might be argued that the evolutionary perspective uniquely provides us with an explanation as to why society should be such an imperfect phenomenon why, despite its obvious advantages, societies are always riven by conicts and bedevilled by individuals who willfully insist on cheating the system. Note that, by the same token, the evolutionary approach does not thereby provide a justication for freeriding and other more explicit forms of theft and exploitation; rather, it simply provides an explanation as to why we should constantly have to struggle with them. It is also important to appreciate that the evolutionary approach is a statistical one. It does not assume that every individual makes exactly the right decision, but, rather, only that on average a population of individuals will do so. The fact that some individuals get it wrong does not disprove the

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hypothesis. That is only true if a majority of individuals do so. The evolutionary perspective always requires some individuals to get it wrong, otherwise there would be no variation on which natural selection can act; indeed, when variation ceases to exist, evolution grinds to a halt. Perhaps the most frequent example raised in this context is voluntary childlessness (or, in the more general case, the reduction in family size associated with the demographic transition). The evolutionary bottom line is that voluntary childlessness is a choice that we can make, and it is one that we may take up as a result of some form of cultural transmission (i.e. the third arm of the evolutionary approach to behaviour). It may be that it genuinely is maladaptive in a biological sense, but if so we might expect the enthusiasm for it to wane in due course (and it is probably far too early to make any judgement on that yet). However, it is important not to take too nave a view of complex biological processes: because individuals are embedded in societies that are essential to their success, it is sometimes necessary to act in ways that are, in immediate terms, maladaptive (in the sense of not being in ones shortterm self-interest). One of those may be to persuade people to adhere to norms of behaviour that, in the long term, are in everyones interests. Moreover, we need to remember that personal reproduction is not the only way of contributing to ones own tness: investing in the reproduction of relatives may be just as effective. We would need to show that the voluntarily childless do not invest proportionately more often in relatives before we can conclude that they are behaving in a strictly maladaptive way. There is now ample evidence from a number of studies that voluntarily or involuntarily childless couples invest more heavily in their nieces and nephews (Pollett et al., in press; see also Silk, 1980, 1990). Doing so allows individuals to gain tness through the genes that they share with their close relatives, in line with Hamiltons theory of kin selection. One of the problems that social scientists often have when engaging with evolutionary biology is that they tend to drastically underestimate the complexity of the biological processes (and hence the arguments) involved. One example is provided by Moore (1990)10 in an attempted rebuttal of Chagnons (1988) imputed claim that humans are naturally warlike. Moore, in what seems like a perfectly sensible analysis, used historical demographic data from the Cheyenne to show that peace chiefs (hereditary chiefs who took no part in active war) had larger family sizes than war chiefs (nonhereditary chiefs responsible for actively leading the tribe in war), not least because many war chiefs died on the battleeld before even having the chance to marry, let alone reproduce. Moore argued that if there was such a thing as a gene for warlike behaviour (presumably ultra-aggressiveness), then such a gene would very rapidly be bred out of the gene pool by the intense selection pressure acting against it from the high mortality rates incurred by aggressive war chiefs. Unfortunately, while such an analysis would sufce for

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many simple anatomical traits (eye colour, leg length), it fails to take note of the complexities that enter into the calculations where behaviour in multilevel societies is concerned. In effect, it attacks the problem at the wrong end. The actual choices faced by Cheyenne were not between being a peace chief and being a war chief as two equal options (whether these are genetic traits, socially inherited traits or personal choices), but rather between the costs and benets of being either a peace chief or a war chief versus being a humble brave at the bottom (relatively speaking) of the social hierarchy for two completely separate groups of men (sons of peace chiefs versus the orphaned sons of braves,11 respectively) (Dunbar, 1991). Only the sons of peace chiefs could be peace chiefs (with the benets that arise therefrom), whereas the orphaned sons of ordinary members of Cheyenne society faced an invidious choice between taking the very risky (but, if they survived, ultimately very successful12) strategy of trying to become a war chief or being outcasts on the fringes of society (where their chances of marriage were all but zero). In fact, the demographic data show very nicely that the relative frequencies of these two types of chief exactly balanced the risks they ran and benets they gained. In effect, being a war chief is, for orphans, what evolutionary biologists would refer to as a best of a bad job strategy (Krebs and Davies, 1993): they do not have the option of being a peace chief, so they actually only have two options to choose between whereas the sons of peace chiefs have three and wisely decline to consider the war chief route. Moores mistake is to have assumed that he is dealing with a situation involving two alleles for a gene for warlike behaviour (e.g. aggressiveness versus peacefulness), when in fact he is dealing with a single gene (the gene for aggressiveness possessed by every individual in the population) which can be expressed to different extents in different contexts. Indeed, this is not even necessarily a genetic issue, but rather one of how individuals respond (often in a conscious and calculating manner) to the circumstances in which they nd themselves. By the same token, the demographic transition becomes understandable when reproduction is seen in the context of the need for increasingly expensive parental investment as societies shift from agricultural to knowledgebased economies, with the resulting need to invest in costly education in the latter (Mace, 1998). As Mace points out, we have often tended to assume that societies are homogenous, when in fact they are not: as in the Cheyenne case, each of us operates within a limited socio-economic context which not only limits whom we compete with but also dictates the options we have available to us and the costs and benets of pursuing those options. It seems that the Registrar Generals socio-economic classes have an evolutionarily intrusive reality because they affect what you can afford to do when the strategies available to you are economically costly. In short, the evolutionary perspective is invariably much more complex than it often appears at rst sight

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especially if rst sight equates the evolutionary approach with the genetic determinism of behaviour. If I have a plea on which to end, it is that social scientists refrain from the kinds of knee-jerk abhorrence of the evolutionary approach, and ask instead how this approach might benet what they do. The key message is that the evolutionary approach is not the same thing as genetic determinism; rather, it is about individuals strategic decision-making under the constraints of circumstance. More importantly, perhaps, adopting an evolutionary approach does not ipso facto necessitate throwing all of sociology, anthropology, economics, or even history, into the dustbin. These disciplines remain fundamentally important to the evolutionary endeavour because they provide the essential details of both the context and the costs and benets that are required by the evolutionary approach. The evolutionary approach is not an alternative to the various social science disciplines, but rather a complement. And, importantly, it is a complement that offers the opportunity of integrating the disparate social sciences into a single intellectual framework.

NOTES
1 The problem was not resolved until the rediscovery of Mendels laws of inheritance in the rst decade of the 20th century, long after both Lamarck and Darwin were dead. It was the synthesizing of Mendels mechanism of inheritance with Darwins theory of natural selection that produced the grand synthesis of NeoDarwinism during the 1920s and 1930s (Mayr, 1982). The key distinction here is the role that use (or natural selection) plays. Weizmanns point was that use cannot alter the germline (i.e. the actual genetic material itself); rather, it merely alters the frequencies with which different versions (mutants, alleles) get passed into the next generation. I should hasten to emphasize that memes per se are not an integral part of the biological study of cultural evolution and its underlying processes. They are simply a convenient metaphor for thinking about the processes of cultural evolution so as to avoid unnecessary cross-slippage from the biological concept of the gene. Many who study cultural evolutionary processes do not even bother to use the term. The term evolutionary psychology (often lower-case) is also used to refer to all three of these sub-elds combined. This distinction is not widely appreciated, but it is important to understand that many of those who label themselves as evolutionary psychologists (in the broad sense) would not sign up to some of the more extreme views of the Evolutionary Psychologists (in the narrow sense) (for a lucid overview, see Laland and Brown, 2002). The critical distinction lies in the choice of the utility function (that which is maximized). Economists assume that this is monetary prot (or, at least more recently, occasionally psychological variables such as happiness). For behavioural ecologists, these are merely proxy indices for the real utility function,

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EVOLUTION AND THE SOCIAL SCIENCES namely tness (which is usually better approximated by real biological outcomes such as numbers of offspring). In the biologists view, the complexities of the real world mean that proximate economic goals like happiness or prot are not always well correlated with tness: maximizing the numbers of surviving offspring produced (i.e. tness) does not always involve maximizing the numbers of offspring born (or conceived), because producing too many may dilute the parents ability to invest sufciently in each to ensure their survival. The important issue is that the mathematics of economics is neutral as to the exact nature of the utility function, so the same mathematics can be used in both cases. Formally, Hamiltons Rule states that a gene for altruism could evolve within a population whenever the benets to the recipient, when devalued by the coefcient of relatedness between altruist and recipient, exceed the cost to the altruist, when both are measured in terms of the numbers of additional future offspring gained or lost, respectively. This, in a nutshell, is Hamiltons Theory of Kin Selection. Altruism is formally dened as behaviour that results in a recipient gaining additional future reproductive opportunities which result in the actor losing future reproductive opportunities as a direct result. In the limiting case, the actor loses all future reproductive opportunities if he or she actually gives up his or her life in the process. Altruism in the everyday sense (e.g. small donations to beggars) will rarely come under this rubric because the costs will not usually be sufcient to affect the altruists tness; evolutionary biologists sometimes refer to altruism of this kind as weak altruism and do not consider that it needs an explanation. However, it is worth emphasizing that kin selection is just one of at least four biological explanations for the evolution of altruistic behaviour (for more details, see Barrett et al., 2002 or any other standard textbook). Complete in that, uniquely among studies of the Trivers-Willard effect in humans, they show that the assumptions of the Trivers-Willard Hypothesis hold in their study population, that the effects arise as a within-population phenomenon, and that the tness consequences of the alternative behavioural strategies follow. It is important to appreciate that the Hungarian Gypsy community was settled in villages under imperial edict in the mid-19th century, and has not been nomadic for a century and half. The urban Gypsy and ethnic Hungarian communities lived in the same towns, the rural ones in separate but nearby ethnically homogenous villages. The GoreTex company provides a particularly nice example of this because they have insisted on subdividing their factories into separate buildings whenever the number of employees in a factory reaches 150 precisely so as to avoid the need to have hierarchical management structures while at the same time ensuring the efcient smooth running of each factory unit (Gladwell, 2000). It may be no accident that they have been a particularly successful company. I cite this example without intending to be personally critical of Moore. My point is simply that, given his disciplinary background, Moores analysis was a perfectly natural one. Many other examples could be chosen: this is simply one that has been worked out in detail. Moore (1990) reminds us that war chiefs were, almost without exception, the orphaned sons of ordinary members of society.

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11

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12 Although war chiefs could not marry and habitually pursued risky strategies like taking oaths never to leave the eld of battle alive unless they won, those that did survive did in fact revert to being normal members of society (i.e. they were relieved of their battleeld oaths and allowed to marry). These individuals were often very attractive husbands (and, in all likelihood, illicit lovers) and often produced more than the average number of children (even without taking into account any illegitimate offspring they may have had).

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BIOGRAPHICAL NOTE is British Academy Research Professor in the School of Biological Sciences, University of Liverpool, and co-director of the British Academys centenary research project Lucy to Language. His research focuses on the evolution of sociality, and combines eldwork on animals and humans in their natural setting, experimental studies of cognition and comparative analysis of brain evolution. Address: British Academy Centenary Research Project, School of Biological Sciences, University of Liverpool, Crown Street, Liverpool L69 7ZB, UK. [email: rimd@liv.ac.uk]
ROBIN DUNBAR, FRAI, FBA,

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