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• W2972718624 abstract "The cultural repertoires of apes have been charted by identifying cultural differences between populations. A new approach focused on great apes’ intense ‘peering’ during social learning suggests that they may possess many more cultural elements than currently thought. The cultural repertoires of apes have been charted by identifying cultural differences between populations. A new approach focused on great apes’ intense ‘peering’ during social learning suggests that they may possess many more cultural elements than currently thought. A veritable deluge of studies has documented the cultural transmission of information from individual to individual across a range of vertebrate and invertebrate species [1Whiten A. A second inheritance system: the extension of biology through culture. Interf.Focus. 2017; 7: 20160142Google Scholar]. Some of the earliest roots of this current research bonanza were in primatology, first in the famous mid-twentieth-century discovery of the spread of sweet-potato washing and other traditions among Japanese macaques [2McGrew W.C. Chimpanzee Material Culture: Implications for Human Evolution. Cambridge University Press, Cambridge1992Crossref Google Scholar], and later in field studies of the great apes [2McGrew W.C. Chimpanzee Material Culture: Implications for Human Evolution. Cambridge University Press, Cambridge1992Crossref Google Scholar]. Just a few years after her first reports of tool use in wild chimpanzees, Jane Goodall boldly wrote of ‘cultural elements in a chimpanzee community’ [3van Lawick-Goodall J. Cultural elements in a chimpanzee community.in: Menzel E. Precultural Primate Behaviour. Karger, Basel1973: 144-184Google Scholar], fuelled both by observations of juveniles’ attention avidly focused on the technical skills of their elders (Figure 1) and early indications that chimpanzees behaved in different ways across African study sites, in ways not apparently explicable by ecological factors. By 1999, it was possible to document these differences in a systematic survey across research sites, revealing an unprecedent array of 39 such putative cultural variations in wild chimpanzees [4Whiten A. Goodall J. McGrew W.C. Nishida T. Reynolds V. Sugiyama Y. Tutin C.E.G. Wrangham R.W. Boesch C. Cultures in chimpanzees.Nature. 1999; 399: 682-685Crossref PubMed Scopus (1568) Google Scholar]. This discovery was soon followed by reports of similar arrays of 19–24 cultural variants in orangutans [5van Schaik C.P. Ancrenaz M. Borgen G. Galdikas B. Knott C.D. Singleton I. Suzuki A. Utami S.S. Merrill M. Orangutan cultures and the evolution of material culture.Science. 2003; 299: 102-105Crossref PubMed Scopus (696) Google Scholar] and more recently by a further convergent report of 22 such putative traditions in gorillas [6Robbins M.M. Ando C. Fawcett K.A. Gruiter C.C. Hedwig D. Iwata Y. Lodwick J.L. Masi S. Salmi R. Stoinski T.S. et al.Behavioural variation in gorillas: evidence of potential cultural traits.PLoS One. 2016; 11 (e0160483)Crossref Scopus (56) Google Scholar]. The impressive scope of great ape cultures inferred by these studies was undreamt of half a century ago, as illustrated in a statement by Peter Medawar that “human beings owe their biological supremacy to the possession of a form of inheritance quite unlike that of other animals: … the apparatus of culture” [7Medawar P. Unnatural science.The New York Review of Books. 1977; (Feb. 3rd, 13–18)PubMed Google Scholar]. Nevertheless, in a new study Caroline Schuppli and Carel van Schaik [8Schuppli C. van Schaik C.P. Animal cultures: how we’ve only seen the tip of the iceberg.Evol. Hum. Sci. 2019; 1: e2Crossref Scopus (35) Google Scholar] now suggest that the scope of ape culture has been radically underestimated by the focus on behavioural differences between communities. They advocate a new methodological approach that additionally accommodates cultural adaptations to local ecological contexts, and even cultural universals. Schuppli and van Schaik [8Schuppli C. van Schaik C.P. Animal cultures: how we’ve only seen the tip of the iceberg.Evol. Hum. Sci. 2019; 1: e2Crossref Scopus (35) Google Scholar] concentrate on avid visual attention (Figure 1), labelling it ‘peering’. In an earlier paper that constitutes a crucial foundation for the present one, Schuppli and colleagues [9Schuppli C. Meulman E.J.M. Forss S.I.M. Aprilinayati F. van Noordwijk M.A. van Schaik C.P. Observational learning and socially induced practice of routine skills in immature orangutans.Anim. Behav. 2016; 119: 87-98Crossref Scopus (63) Google Scholar] systematically tested the validity of peering as an index of observational learning in young wild orangutans. This is important because peering itself need not necessarily implicate learning: for example, it might instead reflect intent to scrounge food items [10Corp N. Byrne R.W. The ontogeny of manual skill in wild chimpanzees: evidence from feeding on the fruit of Saba florida.Behav. 2002; 139: 137-168Crossref Scopus (53) Google Scholar]. Young wild capuchins have been shown to preferentially watch the best adult nutcrackers [11Ottoni E.B. de Resende B.D. Izar P. Watching the best nutcrackers: what capuchin monkeys (Cebus apella) know about others’ tool-using skills.Anim. Cogn. 2005; 8: 215-219Crossref PubMed Scopus (112) Google Scholar], but this benefits their scrounging as well as any potential for observational learning. Accordingly, Schuppli and colleagues [9Schuppli C. Meulman E.J.M. Forss S.I.M. Aprilinayati F. van Noordwijk M.A. van Schaik C.P. Observational learning and socially induced practice of routine skills in immature orangutans.Anim. Behav. 2016; 119: 87-98Crossref Scopus (63) Google Scholar] applied multiple tests of the hypothesis that young orangutans’ peering serves primarily a learning function. All yielded confirmatory evidence. For example, in different contexts, such as stick-tool use and nest building, episodes of peering were followed by heightened rates of exploration of the items concerned. The frequency of peering was predicted by the complexity and difficulty of the procedures peered at and rose along with the learning of new skills and diminished as competence was manifested. Having assured themselves of the validity of peering as in index of social learning, the authors went on in the new paper [8Schuppli C. van Schaik C.P. Animal cultures: how we’ve only seen the tip of the iceberg.Evol. Hum. Sci. 2019; 1: e2Crossref Scopus (35) Google Scholar] to document all the contexts in which peering occurs in the everyday lives of juvenile orangutans at two field sites, Suaq in Sumatra and Tuanan in Borneo. From these records the authors estimate that over the course of their development, orangutans may engage in as many as 38,000 episodes of peering. Peering occurred in as many as 124–195 different contexts across the two sites, spanning a diverse range of activities including tool use, foraging choices and nest building. As the authors remark, these figures vastly exceed the fewer than 30 cultural variants earlier documented at these sites through the ‘traditional’ approach [12van Schaik C.P. Ancrenaz M. Djojoasmoro R. Knott C.D. Morrogh-Bernard H.C. Nuzuar K.O. Atmoko S.S.U. van Noordwijk M.A. Orangutan cultures re-visited. In Orangutans: Geographic Variation in Behavioral Ecology and Conservation.in: Wich S.A. Atmoko S.S.U. Setia T.M. van Schaik C.P. Oxford University Press, Oxford2009: 299-309Google Scholar]. Thus, they conclude, “when looking closely at great ape skill acquisition, it seems that immatures learn virtually all of their skills socially” [8Schuppli C. van Schaik C.P. Animal cultures: how we’ve only seen the tip of the iceberg.Evol. Hum. Sci. 2019; 1: e2Crossref Scopus (35) Google Scholar]. In 1976 Dawkins [13Dawkins R. The Selfish Gene. Oxford University Press, Oxford1976Google Scholar] proposed a cultural analogy of the gene — the meme — conceptualized as a “unit of cultural transmission” [13Dawkins R. The Selfish Gene. Oxford University Press, Oxford1976Google Scholar]. The term has since been assimilated into everyday discourse. Memes were suggested to include human catch-phrases and ways of making pots, but in principle any culture should be decomposable into a vast list of its constituent memes. Is this what we now have in Schuppli and van Schaik’s list of elements for the two orangutan sites? These appear to be offered as putative ‘units of cultural transmission’, picked out by the peering focused on them by juveniles. The difficulty here, and indeed in all attempts to describe a community’s cultural repertoire in terms of a discrete number of cultural units or elements, lies in what is to count as a unit. In humans, is Catholicism a meme, or is it constituted of numerous memes at lower levels, like angels, altars and prayers? Likewise, in orangutans, is using a tool to poke seeds out of a fruit a meme, or are constituent parts like holding the tool in one’s mouth the memes? This fundamental question of ‘unit granularity’ is not explicitly addressed by the authors. Dawkins suggested that a gene be defined as any bit of DNA that “lasts for enough generations to serve as a unit of selection” [13Dawkins R. The Selfish Gene. Oxford University Press, Oxford1976Google Scholar]. This criterion can be directly applied to what is to count as a meme, transmitted across multiple cultural generations. Whether the elements defined by Schuppli and van Schaik [8Schuppli C. van Schaik C.P. Animal cultures: how we’ve only seen the tip of the iceberg.Evol. Hum. Sci. 2019; 1: e2Crossref Scopus (35) Google Scholar] elements are indeed units of (cultural) selection is a challenge yet to be addressed. Some exciting beginnings have been made in identifying the operation of selection and cultural evolution in non-human species more generally, for example in birdsong [14Aplin L.M. Culture and cultural evolution in birds: a review of the evidence.Anim. Behav. 2019; 147: 179-187Crossref Scopus (73) Google Scholar]. Schuppli and van Schaik [8Schuppli C. van Schaik C.P. Animal cultures: how we’ve only seen the tip of the iceberg.Evol. Hum. Sci. 2019; 1: e2Crossref Scopus (35) Google Scholar] interpret their results through the analogy of an iceberg, representing the scope of ape culture (Figure 2). So far, they argue, we have glimpsed only its tip visible above the surface, which is made up of the most noticeable elements documented by the earlier cross-cultural studies, such as forms of tool use. Beneath these is a large set of behaviour patterns missed by the cross-cultural approaches because of their reliance on excluding elements explicable by local ecological circumstances. This weakness of the cross-cultural approach [4Whiten A. Goodall J. McGrew W.C. Nishida T. Reynolds V. Sugiyama Y. Tutin C.E.G. Wrangham R.W. Boesch C. Cultures in chimpanzees.Nature. 1999; 399: 682-685Crossref PubMed Scopus (1568) Google Scholar, 5van Schaik C.P. Ancrenaz M. Borgen G. Galdikas B. Knott C.D. Singleton I. Suzuki A. Utami S.S. Merrill M. Orangutan cultures and the evolution of material culture.Science. 2003; 299: 102-105Crossref PubMed Scopus (696) Google Scholar, 6Robbins M.M. Ando C. Fawcett K.A. Gruiter C.C. Hedwig D. Iwata Y. Lodwick J.L. Masi S. Salmi R. Stoinski T.S. et al.Behavioural variation in gorillas: evidence of potential cultural traits.PLoS One. 2016; 11 (e0160483)Crossref Scopus (56) Google Scholar] was recognized by its proponents, seeing it as a first, broad-brush attempt to start to grasp the extent and scope of ape culture. The new study of Schuppli and van Schaik [8Schuppli C. van Schaik C.P. Animal cultures: how we’ve only seen the tip of the iceberg.Evol. Hum. Sci. 2019; 1: e2Crossref Scopus (35) Google Scholar] is the first to provide an evidence-based means of delineating a much broader range of cultural adaptions to an ape community’s environment. The base layer of the pyramid includes behaviours that may even be universal in the species, yet still culturally acquired (‘knife use’ might be an equivalent in our own species, for example). These middle and lower parts of the iceberg include many basic and simple behavioural elements, such as selective food choice. It is far from implausible that cultural inheritance is important at this level, given the large size of apes' dietary repertoire and the vastly greater array of potential items never ingested. Over 200–300 different items (varied parts of numerous species, including fruits, seeds, leaves, stems, pith, flowers, bark, and roots, not to mention vertebrate and invertebrate prey) may be eaten by chimpanzees [15Inskipp T. Chimpanzee (Pan troglodytes).in: Caldecott J. Miles L. World Atlas of Great Apes and Their Conservation. Univ California Press, Berkeley2005: 53-81Google Scholar], gorillas [16Ferriss S. Western gorilla (Gorilla gorilla).in: Caldecott J. Miles L. World Atlas of Great Apes and Their Conservation. Univ California Press, Berkeley2005: 105-127Google Scholar] and orangutans [17McConkey K. Bornean orangutan (Pongo pygmaeus).in: Caldecott J. Miles L. World Atlas of Great Apes and Their Conservation. Univ California Press, Berkeley2005: 161-183Google Scholar], including a highly selective sub-set of all the potential parts of the many hundreds of species that make up tropical forests. This means there may be well over 1,000 options that could in principle be explored individually, so the value of peering at and learning from the accumulated wisdom underlying the skills and choices of one’s mother, and later others, could indeed be very significant. Field experiments, so far conducted only with other primate species, confirm that infants’ diet selection can be highly dependent on the choices of their mother and her community [18van de Waal E. Borgeaud C. Whiten A. Potent social learning and conformity shape a wild primate’s foraging decisions.Science. 2013; 340: 483-485Crossref PubMed Scopus (283) Google Scholar]. In the new ‘peering’ based study of Schuppli and van Schaik [8Schuppli C. van Schaik C.P. Animal cultures: how we’ve only seen the tip of the iceberg.Evol. Hum. Sci. 2019; 1: e2Crossref Scopus (35) Google Scholar], and the challenge of its conclusions, we may be seeing a significant leap in our efforts to capture the true scope of ape culture. The underlying approach may significantly extend the methods applied to study social learning research in the future, in a variety of species other than apes." @default.
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• W2972718624 title "Social Learning: Peering Deeper into Ape Culture" @default.
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