Ecology,Biology and Social Life: Explaining the Origins of Primate Sociality

Usually, these are to be found in attempts to review or to popularize sociobiology and its various modern incarnations. See, for example, Thornhill R. Palmer C. , A natural history of rape: Biological bases of sexual coercion (Cambridge, MA, 2000); Barash D. , Sociobiology: The whisperings within (London, 1979); Blackmore S. , The meme machine (Oxford, 1999); and Wright R. , The moral animal: Why we are the way we are (New York, 1994). Perhaps as a result of the popularity of evolutionary accounts of human nature, those who have tried to combat them in print — For example, Rose H. Rose S. (eds), Alas, poor Darwin: Arguments against evolutionary psychology (London, 2000) — Have concentrated on detailed criticism of these accounts, rather than on the production of alternative stories.

This is by no means the point at which the ‘nature/nurture’ debate began, but it did become particularly intense during this period. For example, consider the case of Cyril Burt. The first psychologist to be honoured by the British government, his work on the heritability of IQ had a powerful influence over the development of governmental education policy: His twin studies had demonstrated that a child's intelligence was innate. Although his work did not go unchallenged during his lifetime, it was not until the 1970s — At the point where the controversy surrounding sociobiology was in full flower — That he was directly accused of having committed fraud in order to advance his belief that intelligence was inborn. Kamin Leon (The science and politics of IQ (London, 1974)) had argued that his work was flawed, but it was only in 1976 that Oliver Gillie claimed in the Sunday Times that he had actively falsified data, and Leslie Hearnshaw's biography, Cyril Burt: Psychologist (New York, 1979) regretfully confirmed this. More recent assessments of Burt's work — For example, Joynson R. , The Burt affair (London, 1989), or Mackintosh N. , Cyril Burt: Fraud or framed? (Oxford, 1995) — Have arrived at less damning conclusions, suggesting that Burt was guilty of nothing more than poor scientific practice. The IQ debate continued to simmer, and occasionally seethe, during the latter half of the twentieth century, as the work of Herrenstein R. Murray C. , The bell curve: Intelligence and class structure in American life (New York, 1994), or Jacoby R. Glauberman N. (eds), The bell curve debate (New York, 1995) illustrates, and often in a context that seemed directly related to the rise of neo-conservatism in US politics.

This predisposition has in recent years finally begun to receive the scholarly attention it deserves, as part of the vast expansion of the human/animal literature. For examples, see Burt J. , Animals in film (London, 2003), Daston L. Mitman G. (eds), Thinking with animals: New perspectives on anthropomorphism (New York, 2006), Franklin A. , Animals and modern cultures: A sociology of human-animal relations in modernity (London, 1999), Ham J. Senior M. , Animal acts: Configuring the human in Western history (New York, 1997), Haraway D. , The companion species manifesto: Dogs, people and significant others (Chicago, 2003), Manning A. Serpell J. , (eds), Animals and human society: Changing perspectives (London, 1994), Ritvo H. , The animal estate: The English and other creatures in the Victorian Age (Cambridge, MA, 1987), Mitman G. , Reel nature: America's romance with nature on film (Cambridge, MA, 1999), Mitchell R. Thompson N. Miles L. (eds), Anthropomorphism, anecdotes and animals (New York, 1997), and Crist E. , Images of animals: Anthropomorphism and animal mind (Philadelphia, 1999).

Mitman G. , The state of nature: Ecology, community and American social thought, 1900–1950 (Chicago, 1992).

Scott J. P. , “Introduction”, Annals of the New York Academy of Sciences, li (1950), 1003–5.

Anon, “Preface”, Folia primatologica, i (1963), 1.

Washburn S. , “The new physical anthropology”, Transactions of the New York Academy of Sciences, xiii (1951), 258–304, outlines his vision for this new orientation, which was arguably his most influential paper ( Marks J. , “Sherwood Washburn, 1911–2000”, Evolutionary anthropology, ix (2000), 225–6). The origins of this framework and its links with the emergence of the ‘man the hunter’ paradigm are discussed in Haraway D. , Primate visions: Gender, race and nature in the world of modern science (London, 1989). Its specific impact on anthropology is assessed in Strum S. Lindberg D. (eds), The new physical anthropology (New York, 1998), an edited collection intended to illustrate the immense influence that Washburn had had over twentieth-century American anthropological practice.

The number of books and articles that use primate behaviour to explain human activities is far too large to reference here. Familiar to many people will be the work of Desmond Morris — The naked ape (London, 1967), The human zoo (London, 1969) — Or that of Robert Ardrey — African genesis: A personal investigation into the animal origins and nature of man (London, 1961) — While more modern examples include Wrangham R. Peterson D. , Demonic males: Apes and the origins of human violence (London, 1997), Small M. , Female choices: Sexual behaviour of female primates (New York, 1995), and Power M. , The egalitarians – human and chimpanzee: An anthropological view of social organisation (Cambridge, 1991). The last three books in particular illustrate the vast possibilities that the huge variety of different primate behaviours and social structures provides for those who wish to ‘naturalize’ human activities. While Wrangham and Peterson emphasize the long history of male aggression in the evolution of the ape line (with the caveat that bonobos are rather different), Small focuses her attention on the evolution of female behaviour, especially the evolution of sexual independence in the female primate line. In turn, Power argues that rather than demonstrating the inevitability of hierarchical behaviour, an examination of chimp society reveals the roots of egalitarianism. See Rees A. , “Higamous, hogamous, woman monogamous”, Feminist theory, i (2000), 365–70, for a discussion of these ‘naturalized’ accounts of human activities. 9. See, for example, Haraway , op. cit. (ref. 7), Radick G. , “Primate language and the playback experiment, in 1890 and 1980”, Journal of the history of biology, xxxviii (2005), 461–93, Strum S. Fedigan L. , (eds), Primate encounters: Models of science, gender and society (Chicago, 2000), and Asquith P. “Anthropomorphism and the Japanese and Western traditions in primatology”, in Else J. G. Lee P. C. (eds), Primate ontogeny, cognition, and social behaviour (Cambridge, 1986), 61–71.

Yerkes Robert was a comparative psychologist. He was responsible for the creation of the Yerkes Primate Laboratories in Monkey Park, Florida, and believed that an understanding of non-human primate mind was essential to an appreciation of the workings of that of the human primate. For his own account of his approach, see Yerkes R. M. , Chimpanzees: A laboratory colony (New Haven, 1943), or Yerkes R. M. Yerkes A. , The great apes: A study of anthropoid life (New Haven, 1929). For a commentary on the development of Yerkes's methodological approach, see Thomas M. , “Yerkes, Hamilton and the experimental study of ape mind: From evolutionary psychiatry to eugenic politics”, Studies in the history and philosophy of biological and biomedical sciences, xxxvii (2000), 273–94. Solly Zuckerman's first appointment was as an anatomist with the Zoological Society of London. He was involved in the establishment of the baboon colony at London Zoo, and spent a short period working with Robert Yerkes in Florida, before returning to Britain. Throughout his life, he maintained his interests in primate behaviour and evolution, even as he became one of the key influences over British scientific policy in the post-war years. See Peyton J. , Solly Zuckerman: A scientist out of the ordinary (London, 2001), or Burt J. , “Solly Zuckerman: The making of a primatological career in Britain, 1925–1945”, Studies in the history and philosophy of biological and biomedical sciences, xxxvii (2006), 295–310.

Yerkes Yerkes , op. cit. (ref. 10), 582.

Zuckerman S. , The social life of monkeys and apes (London, 1932), 11.

Zuckerman , op. cit. (ref. 12), 13.

Carpenter Clarence Ray , Henry Nissen and Harold Bingham were sent respectively to study howler monkeys, chimpanzees and gorillas. For details, see Nissen H. , “A field study of the chimpanzee”, Comparative psychology monographs, viii (1931), 1–122; Bingham H. , “Gorillas in a native habitat”, Carnegie Institution publications, cdxxvi (1932); and Carpenter C. , “A field study of the behaviour and social relations of howling monkeys (Alouatta palliata)”, which was first published in Comparative psychology monographs, x (1934), 1–186, then reprinted in Carpenter C. , Naturalistic behaviour of nonhuman primates (Pennsylvania, 1964), 3–92. Of the three, Carpenter's was undoubtedly the most successful, and his influence over the ‘second-generation’ of American field primatologists can perhaps be gauged from the fact that in 1964, his early papers and monographs had to be collected and reprinted, since they had become part of the basic literature of the new discipline.

See, for example, the discussions in Washburn S. , (ed.), Social life of early man (London, 1962); all of the contributions to the special edition of Symposia of the Zoological Society of London, x (1963), 1–128, especially Zuckerman's concluding remarks (pp. 119–23); Southwick C. (ed.), Primate social behaviour (New Jersey, 1963); Howell F. C. (ed.), African ecology and human evolution (London, 1964); DeVore I. (ed.), Primate behaviour: Field studies of monkeys and apes (New York, 1965); Altmann S. (ed.), Social communication among primates (Chicago, 1967); and Jay P. (ed.), Primates: Studies in adaptation and variability (New York, 1968).

Further discussion of primatological methodological debates can be found in Radick G. , “What's in a name? The vervet predator calls and the limits of the Washburnian synthesis”, Studies in the history and philosophy of biological and biomedical sciences, xxxvii (2006), 334–62; Rees A. , “A place that answers questions: Primatological field sites and the making of authentic observations”, Studies in the history and philosophy of biological and biomedical sciences, xxxvii (2006), 311–33; Montgomery G. , “Place, practice and primatology: Clarence Ray Carpenter, primate communication and the development of field methodology”, Journal of the history of biology, xxxviii (2005), 495–533; Strum Fedigan , op. cit. (ref. 9); and Haraway , op. cit. (ref. 7).

Primates, unlike most other mammals, can be found in relatively permanent bisexual groups.

Zuckerman , op. cit. (ref. 12), 31.

Carpenter, for example, despite paying rather more attention to ecology and environment, still concurred with the basic premise that it was sex that drew male and female primates together — See, for example, Carpenter, op. cit. (ref. 14, 1964), 66, or Carpenter, “Societies of monkeys and apes”, in Carpenter, op. cit. (ref. 12, 1964), 342–57, p. 353. Marshall Sahlins produced an equally influential account of the close relationship between human and non-human primate social structure that placed the integrative power of sex at the heart of his model. See Sahlins M. , “The social life of monkeys, apes and primitive man”, Human biology, xxxi (1959), 54–73, p. 56.

See Jolly A. , “Breeding synchrony in wild Lemur catta”, in Altmann (ed.), op. cit. (ref. 15), 3–14; Imanishi K. , “Social organisation of subhuman primates in their natural habitat”, Current anthropology, i (1960), 393–407; and Connaway C. Koford C. , “Estrous cycles and mating behaviour in a free-ranging band of rhesus monkeys”, Journal of mammalogy, xlv (1965), 577–88.

Lancaster J. Lee R. , “The annual reproductive cycle in monkeys and apes”, in DeVore (ed.), op. cit. (ref. 15), 486–513.

So, for example, in the postscript to the second edition of his monograph Social lives, which appeared in 1982, Zuckerman dealt with the controversy that had grown up around seasonality as follows: Washburn's “belief that sexual interest plays an insignificant part, if indeed any, in holding together the social units of societies of monkeys and apes seems to be based mainly on a compilation of reports about the times of birth of apes and monkeys made by two of his pupils, Lancaster and Lee, who, apparently without any direct experience of this subject, set about their task with a preconception of what they wanted to reveal” ( Zuckerman S. , The social life of monkeys and apes (London, 1982), 346). He went on to criticise the work that had been completed on primates since his day, especially the emphasis on the amount of time spent in the field (it is impossible to know “whether the observations of the man who was there the longest were the most reliable or valuable”, p. 364), and the turn to quantification of data (where fieldworkers appeared to be making use of “figures which do little more than either confirm the obvious or add to the inventory of trivial observation. The statistical analysis of inexact measures may add a spurious air of exactitude, but it also soon evokes the suspicion that one is being ‘blinded’ by science”, p. 371). It is probably safe to conclude that Zuckerman disapproved of the turn primatology had taken.

See, for example, Washburn S. DeVore I. , “The social life of baboons”, Scientific American, cciv (1961), 62–71, or Washburn S. DeVore I. , “Social behaviour of baboons and early man”, in Washburn (ed.), op. cit. (ref. 15), 91–105.

For examples, see Altmann S. , “Field observations on a howling monkey society”, Journal of mammalogy, xl (1959), 317–30; Chance M. , “The sociability of monkeys”, Man, lv (1955), 162–5; Chance M. , “The nature and special features of the instinctive social bond of primates”, in Washburn (ed.), op. cit. (ref. 15), 17–33; and Imanishi , op. cit. (ref. 20).

See ref. 7 for an account of Washburn's anthropological influences. DeVore also conducted major field research projects in anthropology, such as the Harvard Kalahari Project (1963–80), of which he was co-director. See DeVore I. Lee R. , Kalahari hunter-gatherers (Cambridge, MA, 1976).

The emergence of the baboon model is briefly discussed in Haraway , op. cit. (ref. 7), but is covered in far more detail, particularly in relation to the consequences of the model's development, in Strum Fedigan , op. cit. (ref. 9). Sperling S. , “Baboons with briefcases vs. langurs in lipstick”, in di Leonardo M. (ed.), Gender at the crossroads of knowledge: Feminist anthropology in the postmodern era (Berkeley, 1991), 204–34, also develops an analysis of the baboon model that contrasts it with the understanding of langur behaviour over the same period.

Washburn DeVore , op. cit. (ref. 23), 63–64.

DeVore I. Washburn S. , “Baboon ecology and human evolution”, in Howell (ed.), op. cit. (ref. 15), 335–67.

Partly, this originated in the work of Charles Otis Whitman, but is more famously associated with ethology's primary parents, Karl Lorenz and Niko Tinbergen. See Burkhardt R. , Patterns of behaviour: Konrad Lorenz, Niko Tinbergen, and the founding of ethology (Chicago, 2005).

Washburn S. DeVore I. , “Baboon behaviour”, 16mm film, Motion Picture Production Department, Berkeley, 1960.

See the discussions in Strum Fedigan , op. cit. (ref. 9), especially the contributions of T. Rowell, “A few peculiar primates”, pp. 57–70, and Jolly A. , “The bad old days of primatology?”, pp. 71–84.

Jay P. , “The Indian langur monkey”, in Southwick (ed.), op. cit. (ref. 15), 114–23; Hall K. , “Behaviour and ecology of the wild patas monkeys, Erythrocebus patas, in Uganda”, in Jay P. (ed.), op. cit. (ref. 15), 32–119.

Koyama N. , “On dominance rank and kinship of a wild Japanese monkey troop in Arashiyama”, Primates, xi (1967), 335–90.

Rowell T. , “Forest living baboons in Uganda”, Journal of the Zoological Society of London, cxlix (1966), 344–64, p. 362.

Crook J. Gartlan S. , “Evolution of primate societies”, Nature, ccx (1966), 1200–3, p. 1200.

Particularly, researchers were concerned about the potential impact on behaviour of feeding the animals in order to encourage them to remain close enough to be seen. Goodall's Gombe site was especially criticised, to the point where the suggestion was made that her observations of chimpanzee ‘warfare’ were the direct result of her decisions to feed the chimps with bananas, thus artificially raising levels of tension and aggression. See Reynolds V. , “How wild are the Gombe chimpanzees?”, Man, x (1975), 123–5, for a discussion of the case Gombe Rowell T. , “Variability in the social organisation of primates”, in Morris D. (ed.), Primate ethology (London, 1967), 219–35, for a more general review of the possible ‘artefactual’ causes of primate variation.

Jolly A. , The evolution of primate behaviour (London, 1972).

Crook Gartlan , op. cit. (ref. 35).

Crook Gartlan , op. cit. (ref. 35), 1200.

Although they do note that the “[p]ossession of weapons would have substituted for gross sexual dimorphism” (Crook and Gartlan, op. cit. (ref. 35), 1202) in the case of humans.

This account may appear dominated by the activities of large males, with the result that the behaviour of females and juveniles is sidelined. In part, this reflects what some researchers have, somewhat ironically, called the “bad old days of primatology”, where male primatologists went out to watch male primates fighting each other, and ignored equally complex, but far less obvious female behaviour: Not until women entered the field in large numbers was the importance of female relationships and decision making realized. This picture of primatological life has, however, been challenged by practitioners such as Rowell , op. cit. (ref. 31), and Jolly , op. cit. (ref. 31), who have argued that this represents an over-simplification of the situation in the 1960s, and one that is limited to the Western primatological tradition. Space does not permit further exploration of these debates here.

Janson C. , “Primate socioecology: The end of a golden age”, Evolutionary anthropology, ix (2000), 73–86, p. 74.

Hladik C. , “Ecology, diet and social patterning in Old and New World primates”, in Tuttle R. (ed.), Socioecology and psychology of primates (The Hague, 1975), 3–35; and Clutton-Brock T. Harvey P. , “Primate ecology and social organisation”, Journal of zoology, clxxxiii (1977), 1–39.

Eisenberg J. F. Muckenhirn N. A. Rudran R. , “The relation between ecology and social structure in primates”. Science, clxxvi (1972), 863–74.

See ref. 41, but also it is worth realizing that Eisenberg et at. do devote serious attention to the question of female behaviour — Why do they live in matrilineal groups, for example, or why they might tolerate the disruptive and potentially competitive presence of more than one male in the group? As primatological researchers have themselves emphasized, a close examination of primatological theory and practice shows that it was not the case that females were entirely excluded from accounts of social life during this period, although they did tend to be found on the theoretical margins. Again, space precludes a thorough examination of this topic.

Lorenz , for example, argued that lethal aggression between members of the same species was normally ‘inhibited’ by certain behavioural patterns. So, for example, dogs would not normally fight to the death; instead the fight would end when one animal ‘submitted’ to the other, using characteristic postures and vocalisations that inhibited further aggression from the dominant animal. In the case of humanity, however, such inhibitions had been by-passed by cultural ingenuity: The invention of weapons that acted at a distance. If a soldier cannot see his rival cry “pax”, he is unlikely to show mercy. See Lorenz K. , On aggression (London, 1966).

So, for example, while it was well known that animals of the same species were known to behave differently in different habitats, it was also the case that animals of different species found in the same habitat would also show variable patterns of social organization. Ecology did not entirely determine social structure, but it did represent one of the most fundamental influences over its formation.

Eisenberg , op. cit. (ref. 44), 867.

Washburn S. Jay P. Lancaster J. , “Field studies of old world monkeys and apes”, Science, cl (1965), 1541–7.

Wynne-Edwards V. C. , Animal dispersal in relation to social behaviour (London, 1962); and Williams G. C. , Adaptation and natural selection: A critique of some current evolutionary thought (Princeton, 1966).

Hamilton W. D. , “The genetical evolution of social behaviour I”, Journal of theoretical biology, vii (1964), 1–16; Hamilton W. D. , “The genetical evolution of social behaviour II”, Journal of theoretical biology, vii (1964), 17–32; Trivers R. , “Parental investment and sexual selection”, in Campbell B. (ed.), Sexual selection and the descent of man (Chicago, 1972), 136–79; Trivers R. , “Parent-offspring conflict”, American zoologist, xiv (1974), 249–64; and Maynard-Smith J. , “Evolution and the theory of games”, American scientist, lxiv (1976), 41–45.

Wilson E. O. , Sociobiology: The new synthesis (Cambridge, MA, 1975). There is not space to discuss the detailed progress of the controversy here. Segestrale U. , Defenders of the truth: The sociobiology debate (Oxford, 2001) provides a useful history.

‘Proximate’ cause refers to the immediate event prompting the occurrence of behaviour. ‘Ultimate’ cause relates to the selective pressures that may have caused the behaviour to evolve in the first place.

See Richard A. Schulman S. , “Sociobiology: Primate field studies”, Annual review of anthropology, xi (1982), 231–55, for an extended discussion of the immediate impact that sociobiology had on field studies, and on the difficulties of operationalizing sociobiological concepts in the field.

Rowell T. , “Long term changes in a population of Ugandan baboons”, Folia primatologica, xi (1969), 241–54.

In the United States, historically the discipline is divided into four ‘sub-fields’, namely cultural anthropology, physical anthropology, linguistics and archaeology. It was not just the development of sociobiology that caused problems for this interdisciplinary approach, but also the rise of post-modernism, with its emphasis on the contingent and constructed nature of the world, an attitude that researchers with backgrounds in the natural sciences found hard either to apprehend or to accept. Ironically, however, primatologists themselves found themselves actively participating with historians and sociologists of science in the construction of the history of their discipline: No mean feat in the aftermath of the “Science Wars”. See for example, Strum Fedigan , op. cit. (ref. 9); and Latour B. Strum S. , “Human social origins: Please tell us another story”, Journal of social and biological structures, ix (1986), 169–87.

It is essential to emphasise this point. The hostility was explicitly directed to sociobiological accounts of the origins of behaviour. Opposition to sociobiology did not imply opposition to evolutionary theory: Many scientists who criticized sociobiological thinking did so on the grounds that sociobiology represented a misapprehension of the nature of Darwinian evolution.

The first primatologist to observe such behaviour in langur monkeys, Yukimaru Sugiyama, argued that it was a means through which the formation of social bonds between males and females was facilitated ( Sugiyama Y. , “Group composition, population density and some sociological observations of Hanuman langurs (Presbytis entellus)”, Primates, v (1964), 7–38). Other researchers felt it might be a means of dissipating intense aggression, or a strategy for population regulation and the maintenance of troop structure, or a means of avoiding incest. See respectively, Mohnot S. M. , “Some aspects of social changes and infant killing in the hanuman langur Presbytis entellus (Primates: Cercopithecidae) in Western India”, Mammalia, xxxv (1971), 175–98; Rudran R. , “Adult male replacement in one male troops of purple faced langurs (Presbytis senex senex) and its effect on population structure”, Folia primatologica, xix (1973), 166–92; and Itani J. , “A preliminary essay on the relationship between social organisation and incest avoidance in nonhuman primates”, in Poirer F. (ed.), Primate socialisation (New York, 1972), 165–71.

Hrdy S. B. , “Male-male competition and infanticide among the langurs (Presbytis entellus) of Abu, Rajasthan”, Folia primatologica, xxii (1974), 19–58.

See for example Jay P. , “Aspects of maternal behaviour among langurs”, Annals of the New York Academy of Sciences, cii (1962), 468–76; Jay P. , “The common langur of North India”, in DeVore I. (ed.), op. cit. (ref. 15), 197–249.

Dolhinow P. , “Normal monkeys?”, American scientist, lxv (1977), 266. This theme of ‘abnormality’ was key to the debate, but of course it was in practice very hard to define what an ‘unnatural’ primate group might be. Frequently the default definition was ‘affected by human influence’, but since most primate field sites could be characterized as in some way subject to this, subjective judgements had to be made with regard to when that influence became unmanageable. Hrdy S. B. , “More monkey business?”, American scientist, lxvi (1978), 667–8, made this point very forcefully, and tried to work with colleagues to develop a series of indices against which the level of human influence could be measured objectively. See Bishop N. Hrdy S. Teas J. Moore J. , “Measures of human disturbance in the habitats of South Asian monkeys”, International journal of primatology, ii (1981), 153–67.

Curtin R. Dolhinow P. , “Primate social behaviour in a changing world”, American scientist, lxvi (1978), 468–75.

The initial confrontation between Hrdy and Dolhinow was played out in the pages of American scientist, but soon proliferated. Explicit discussion of the controversy, and attempts to seek strategies for resolution, can be found in Hrdy S. B. , “Infanticide among animals: A review, classification and examination of the implications for the reproductive strategies of females”, Ethology and sociobiology, i (1979), 13–40; Boggess J. , “Troop male membership changes and infant killing in langurs (Presbytis entellus)”, Folia primatologica, xxxii (1979), 65–107; Ripley S. , “Infanticide in langurs and man: Adaptive advantage, or social pathology?”, in Cohen M. N. (eds), Biosocial mechanisms of population regulation (New Haven, 1980); and Hausfater G. Vogel C. , “Infanticide in langur monkeys (genus Presbytis): Recent research and a review of hypotheses”, in Chiarelli A. B. Corruccini R. S. (eds), Advanced views in primate behaviour (Berlin, 1982). This first stage of the controversy culminated in a conference held at Cornell University in 1982 under the auspices of the Wenner Glen Foundation, the proceedings of which appear as Hausfater G. Hrdy S. (eds), Infanticide: Comparative and evolutionary perspectives (New York, 1984). An examination of the contents of these edited conference proceedings makes it clear that the only area where sexually selected infanticide remained controversial was behavioural primatology.

See the reports in Hausfater Hrdy , op. cit. (ref. 63), and also Parmigiani S. vom Saal F. (eds), Infanticide and parental care (London, 1994).

This can be seen in two early reviews of the infanticide literature, namely Hrdy , op. cit. (ref. 63), and Sherman P. , “Reproductive competition and infanticide in Belding's ground squirrels and other animals”, in Alexander R. D. Tinkle D. W. (eds), Natural selection and social behaviour: Recent research and new theory (New York, 1981), 311–31. Hrdy While , whose review focuses on the primate examples, found it necessary to spend a substantial portion of her article discussing the controversy, Sherman, working on Belding's ground squirrels and addressing the animal literature more generally, mentions it only twice, and then only to indicate that it exists in relation to the primatological literature.

Hausfater G. , “Infanticide in nonhuman primates: An introduction and perspective”, in Hausfater Hrdy , op. cit. (ref. 63), 145–50.

The fact that ecology remained vitally important to understanding the behaviour of individual primates and the nature of social structure can be seen from work such as that produced by Wrangham R. , “An ecological model of female-bonded primate groups”, Behaviour, lxxv (1980), 262–300. See Janson , op. cit. (ref. 42), for a detailed account of the importance of socioecology in the development of primatological research in the 1980s and '90s.

Janson , op. cit. (ref. 42).

van Schaik C. Dunbar R. , “The evolution of monogamy in large primates: A new hypothesis and some crucial tests”, Behaviour, cxv (1990), 30–62.

Bartlett T. Sussman R. Cheverud J. , “Infant killing in primates: A review of observed cases with specific reference to the sexual selection hypothesis”, American anthropologist, xcv (1990), 958–90.

The adaptive hypothesis made a number of predictions: That the attacking male should not be related to the infant, that he should then go on to mate with the mother, that the mother should then produce her next infant more quickly than she would have done had the dead infant survived. Sussman and his colleagues required each reported infanticide to meet each of these conditions, before accepting it as a potential act of sexually selected infanticide. Because of the chaotic conditions surrounding male replacements in the field and the difficulty of continuous observation, it was often hard for fieldworkers to complete all of the required detail: In particular, it was often hard to be certain that a particular male had without doubt been responsible for the death of a particular infant. Unsurprisingly then, Sussman and his co-workers found that the infanticide ‘database’ was far smaller than had been assumed, and as indicated, most infanticides had occurred at one site. The fact that most cases of infanticide had been seen at Jodhpur, however, could well be explained by the excellent observation conditions there and the point that field studies had been carried out there long-term. You would, one could argue, expect most reports to come from the place where animals had been studied the longest and under the best conditions.

Sussman R. Cheverud J. Bartlett T. , “Infant killing as evolutionary strategy: Reality or myth?”, Evolutionary anthropology, iii (1995), 149–51.

See Sommer V. , “The holy wars about infanticide: Which side are you on? And why?”, in van Schaik C. Janson C. (eds), Infanticide by males and its implications (Cambridge, 2000), 9–26; and Hrdy S. Janson C. van Schaik C. , “Infanticide: Let's not throw out the baby with the bathwater”, Evolutionary anthropology, iii (1995), 151–4.

This may be explained by the fact that Sussman was editor of American anthropologist, in which the first attack on the sexual selection hypothesis had appeared, and which was later to publish Anne Dagg's attempt to destabilize the consensus that infanticide in lions was the result of sexual selection (“Infanticide by male lions hypothesis: A fallacy influencing research into human behaviour”, American anthropologist, c (1995), 940–50), as well as Craig Packer's robust rebuttal of her claims (“Infanticide is no fallacy”, American anthropologist, cii (2000), 829–31). Further exchanges appeared in Evolutionary anthropology and in Anthropology news ( Silk J. Stanford C. , “Infanticide article disputed”, Anthropology news , September 1999, 27–28; Dagg A. , “Sexual selection is debatable”. Anthropology news , December 1999, 20). It is perhaps ironic that, according to some accounts ( Shea C. , “Motive for murder?”, Inside publishing, ix (1999), 23–25), Sussman had been given the editorship of American anthropologist, the flagship journal of the American Anthropological Association, in the hope that he would be able to heal some of the wounds that the ‘post-modernist’ turn had inflicted on the anthropological body.

There have, naturally, been prominent scientific critics of, for example, sociobiology and evolutionary psychology — See, for example, Lewontin R. Rose S. Kamin L. , Not in our genes: Biology, ideology and human nature (London, 1984); Eldredge N. , Reinventing Darwin: The great debate at the high table of evolutionary theory (London, 1995); and Gould S. J. , Mismeasure of man (London, 1984) — But this paper has been primarily concerned not with the sociobiology debate in particular, but with the way in which the balance between biological and environmental explanations of animal activity shifted over the course of the development of a particular area of behavioural biology.

Sussman R. Chapman A. , The origins and nature of sociality (New York, 2004), 7; and Sober E. Wilson D. Sloan , Unto others: The evolution and psychology of unselfish behaviour (Cambridge, MA, 1998).

Skinner D. , “Racialised futures: Biologism and the changing politics of identity”, Social studies of science, xxxvi (2006), 459–88; and Campbell N. , “Suspect technologies: Scrutinising the intersection of science, technology and policy”, Science, technology and human values, xxx (2005), 374–402.