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      Naive, captive long-tailed macaques ( Macaca fascicularis fascicularis) fail to individually and socially learn pound-hammering, a tool-use behaviour

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          Abstract

          A subspecies of long-tailed macaques ( Macaca fascicularis aurea; Mfa) has been reported to use stone tools and a specific technique to process nuts in Southeast Asia, a behaviour known as ‘pound-hammering’. The aim of this study was to examine the development of pound-hammering in long-tailed macaques: whether this behavioural form can be individually learnt or whether it has to rely on some forms of social learning. Given the absence of Mfa from captivity, long-tailed macaques of a highly related subspecies ( Macaca fascicularis fascicularis; Mff) were experimentally tested by providing them with the ecological materials necessary to show pound-hammering. A baseline was first carried out to observe whether pound-hammering would emerge spontaneously without social information. As this was not the case, different degrees of social information, culminating in a full demonstration of the behaviour, were provided. None of the subjects ( n = 31) showed pound-hammering in any of the individual or social learning conditions. Although these data do not support the hypothesis that individual learning underlies this behaviour, no evidence was found that (at least) Mff learn pound-hammering socially either. We propose that other—potentially interacting—factors may determine whether this behaviour emerges in the various subspecies of long-tailed macaques, and provide a novel methodology to test the role of social and individual learning in the development of animal tool-use.

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          Ratcheting up the ratchet: on the evolution of cumulative culture.

          Some researchers have claimed that chimpanzee and human culture rest on homologous cognitive and learning mechanisms. While clearly there are some homologous mechanisms, we argue here that there are some different mechanisms at work as well. Chimpanzee cultural traditions represent behavioural biases of different populations, all within the species' existing cognitive repertoire (what we call the 'zone of latent solutions') that are generated by founder effects, individual learning and mostly product-oriented (rather than process-oriented) copying. Human culture, in contrast, has the distinctive characteristic that it accumulates modifications over time (what we call the 'ratchet effect'). This difference results from the facts that (i) human social learning is more oriented towards process than product and (ii) unique forms of human cooperation lead to active teaching, social motivations for conformity and normative sanctions against non-conformity. Together, these unique processes of social learning and cooperation lead to humans' unique form of cumulative cultural evolution.
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            Social learning in animals: categories and mechanisms.

            There has been relatively little research on the psychological mechanisms of social learning. This may be due, in part, to the practice of distinguishing categories of social learning in relation to ill-defined mechanisms (Davis, 1973; Galef, 1988). This practice both makes it difficult to identify empirically examples of different types of social learning, and gives the false impression that the mechanisms responsible for social learning are clearly understood. It has been proposed that social learning phenomena be subsumed within the categorization scheme currently used by investigators of asocial learning. This scheme distinguishes categories of learning according to observable conditions, namely, the type of experience that gives rise to a change in an animal (single stimulus vs. stimulus-stimulus relationship vs. response-reinforcer relationship), and the type of behaviour in which this change is detected (response evocation vs. learnability) (Rescorla, 1988). Specifically, three alignments have been proposed: (i) stimulus enhancement with single stimulus learning, (ii) observational conditioning with stimulus-stimulus learning, or Pavlovian conditioning, and (iii) observational learning with response-reinforcer learning, or instrumental conditioning. If, as the proposed alignments suggest, the conditions of social and asocial learning are the same, there is some reason to believe that the mechanisms underlying the two sets of phenomena are also the same. This is so if one makes the relatively uncontroversial assumption that phenomena which occur under similar conditions tend to be controlled by similar mechanisms. However, the proposed alignments are intended to be a set of hypotheses, rather than conclusions, about the mechanisms of social learning; as a basis for further research in which animal learning theory is applied to social learning. A concerted attempt to apply animal learning theory to social learning, to find out whether the same mechanisms are responsible for social and asocial learning, could lead both to refinements of the general theory, and to a better understanding of the mechanisms of social learning. There are precedents for these positive developments in research applying animal learning theory to food aversion learning (e.g. Domjan, 1983; Rozin & Schull, 1988) and imprinting (e.g. Bolhuis, de Vox & Kruit, 1990; Hollis, ten Cate & Bateson, 1991). Like social learning, these phenomena almost certainly play distinctive roles in the antogeny of adaptive behaviour, and they are customarily regarded as 'special kinds' of learning (Shettleworth, 1993).(ABSTRACT TRUNCATED AT 400 WORDS)
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              Cultural innovation and transmission of tool use in wild chimpanzees: evidence from field experiments.

              Chimpanzees (Pan troglodytes) are the most proficient and versatile users of tools in the wild. How such skills become integrated into the behavioural repertoire of wild chimpanzee communities is investigated here by drawing together evidence from three complementary approaches in a group of oil-palm nut- ( Elaeis guineensis) cracking chimpanzees at Bossou, Guinea. First, extensive surveys of communities adjacent to Bossou have shown that population-specific details of tool use, such as the selection of species of nuts as targets for cracking, cannot be explained purely on the basis of ecological differences. Second, a 16-year longitudinal record tracing the development of nut-cracking in individual chimpanzees has highlighted the importance of a critical period for learning (3-5 years of age), while the similar learning contexts experienced by siblings have been found to result in near-perfect (13 out of 14 dyads) inter-sibling correspondence in laterality. Third, novel data from field experiments involving the introduction of unfamiliar species of nuts to the Bossou group illuminates key aspects of both cultural innovation and transmission. We show that responses of individuals toward the novel items differ markedly with age, with juveniles being the most likely to explore. Furthermore, subjects are highly specific in their selection of conspecifics as models for observation, attending to the nut-cracking activities of individuals in the same age group or older, but not younger than themselves. Together with the phenomenon of inter-community migration, these results demonstrate a mechanism for the emergence of culture in wild chimpanzees.
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                Author and article information

                Journal
                R Soc Open Sci
                R Soc Open Sci
                RSOS
                royopensci
                Royal Society Open Science
                The Royal Society Publishing
                2054-5703
                May 2018
                9 May 2018
                9 May 2018
                : 5
                : 5
                : 171826
                Affiliations
                [1 ]School of Psychology, University of Birmingham , Birmingham B15 2TT, UK
                [2 ]Department for Early Prehistory and Quaternary Ecology, University of Tübingen , Tübingen 72070, Germany
                Author notes
                Author for correspondence: Elisa Bandini e-mail: elisa-bandini@ 123456hotmail.it

                Electronic supplementary material is available online at https://doi.org/10.6084/m9.figshare.c.4077707.

                Author information
                http://orcid.org/0000-0002-8293-9171
                Article
                rsos171826
                10.1098/rsos.171826
                5990768
                29892375
                83ca0fe3-5371-48e2-b234-0db961ee8377
                © 2018 The Authors.

                Published by the Royal Society under the terms of the Creative Commons Attribution License http://creativecommons.org/licenses/by/4.0/, which permits unrestricted use, provided the original author and source are credited.

                History
                : 6 November 2017
                : 3 April 2018
                Funding
                Funded by: Economic and Social Research Council, http://dx.doi.org/10.13039/501100000269;
                Award ID: ES/K008625/1
                Categories
                1001
                42
                14
                70
                Biology (Whole Organism)
                Research Article
                Custom metadata
                May, 2018

                zone of latent solutions,individual learning,social learning,pound-hammering,macaca fascicularis aurea,macaca fascicularis fascicularis

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