Publisher's copyright statement:The nal publication is available at Springer via http://dx.doi.org/10.3758/LB.38.3.220.Additional information: Use policyThe full-text may be used and/or reproduced, and given to third parties in any format or medium, without prior permission or charge, for personal research or study, educational, or not-for-prot purposes provided that:• a full bibliographic reference is made to the original source • a link is made to the metadata record in DRO • the full-text is not changed in any way The full-text must not be sold in any format or medium without the formal permission of the copyright holders.Please consult the full DRO policy for further details.

A vital prerequisite for cumulative culture, a phenomenon often asserted to be unique to humans, is the ability to modify behaviour and flexibly switch to more productive or efficient alternatives. Here, we first established an inefficient solution to a foraging task in five captive chimpanzee groups (N = 19). Three groups subsequently witnessed a conspecific using an alternative, more efficient, solution. When participants could successfully forage with their established behaviours, most individuals did not switch to this more efficient technique; however, when their foraging method became substantially less efficient, nine chimpanzees with socially-acquired information (four of whom witnessed additional human demonstrations) relinquished their old behaviour in favour of the more efficient one. Only a single chimpanzee in control groups, who had not witnessed a knowledgeable model, discovered this. Individuals who switched were later able to combine components of their two learned techniques to produce a more efficient solution than their extensively used, original foraging method. These results suggest that, although chimpanzees show a considerable degree of conservatism, they also have an ability to combine independent behaviours to produce efficient compound action sequences; one of the foundational abilities (or candidate mechanisms) for human cumulative culture.

Cumulative culture underpins humanity’s enormous success as a species. Claims that other animals are incapable of cultural ratcheting are prevalent, but are founded on just a handful of empirical studies. Whether cumulative culture is unique to humans thus remains a controversial and understudied question that has far-reaching implications for our understanding of the evolution of this phenomenon. We investigated whether one of human’s two closest living primate relatives, chimpanzees, are capable of a degree of cultural ratcheting by exposing captive populations to a novel juice extraction task. We found that groups (N = 3) seeded with a model trained to perform a tool modification that built upon simpler, unmodified tool use developed the seeded tool method that allowed greater juice returns than achieved by groups not exposed to a trained model (non-seeded controls; N = 3). One non-seeded group also discovered the behavioral sequence, either by coupling asocial and social learning or by repeated invention. This behavioral sequence was found to be beyond what an additional control sample of chimpanzees (N = 1 group) could discover for themselves without a competent model and lacking experience with simpler, unmodified tool behaviors. Five chimpanzees tested individually with no social information, but with experience of simple unmodified tool use, invented part, but not all, of the behavioral sequence. Our findings indicate that (i) social learning facilitated the propagation of the model-demonstrated tool modification technique, (ii) experience with simple tool behaviors may facilitate individual discovery of more complex tool manipulations, and (iii) a subset of individuals were capable of learning relatively complex behaviors either by learning asocially and socially or by repeated invention over time. That chimpanzees learn increasingly complex behaviors through social and asocial learning suggests that humans’ extraordinary ability to do so was built on such prior foundations.

a.whiten@st-andrews.ac.uk 2 The experimental study of cumulative culture and the innovations essential to it is a young science, with child studies so rare that the scope of cumulative cultural capacities in childhood remains largely unknown. Here we report a new experimental approach to the inherent complexity of these phenomena. Groups of 3-4 year old children were presented with an elaborate array of challenges affording the potential cumulative development of a variety of techniques to gain increasingly attractive rewards. In contrast to a prior study, we found evidence for elementary forms of cumulative cultural progress, with inventions of solutions at lower levels spreading to become shared innovations, and some children then building on these to create more advanced but more rewarding innovations. This contrasted with markedly more constrained progress

Chimpanzees (Pan troglodytes) demonstrate much cultural diversity in the wild, yet a majority of novel behaviours do not become group-wide traditions. Since many such novel behaviours are introduced by low-ranking individuals, a bias toward copying dominant individuals (‘rank-bias’) has been proposed as an explanation for their limited diffusion. Previous experimental work showed that chimpanzees (Pan troglodytes) preferentially copy dominant over low-rank models. We investigated whether low ranking individuals may nevertheless successfully seed a beneficial behaviour as a tradition if there are no ‘competing’ models. In each of four captive groups, either a single high-rank (HR, n=2) or a low-rank (LR, n=2) chimpanzee model was trained on one method of opening a two-action puzzle-box, before demonstrating the trained method in a group context. This was followed by eight hours of group-wide, open-access to the puzzle-box. Successful manipulations and observers of each manipulation were recorded. Barnard’s exact tests showed that individuals in the LR groups used the seeded method as their first-choice option at significantly above chance levels, whereas those in the HR groups did not. Furthermore, individuals in the LR condition used the seeded method on their first attempt significantly more often than those in the HR condition. A network-based diffusion analysis revealed that the best supported statistical models were those in which social transmission occurred only in groups with subordinate models. Finally, we report an innovation by a subordinate individual that built cumulatively on existing methods of opening the puzzle-box and was subsequently copied by a dominant observer. These findings illustrate that chimpanzees are motivated to copy rewarding novel behaviours that are demonstrated by subordinate individuals and that, in some cases, social transmission may be constrained by high-rank demonstrators.

Various non-human animal species have been shown to exhibit behavioural traditions. Importantly, this research has been guided by what we know of human culture, and the question of whether animal cultures may be homologous or analogous to our own culture. In this paper, we assess whether models of human cultural transmission are relevant to understanding biological fundamentals by investigating whether accounts of human payoff-biased social learning are relevant to chimpanzees (Pan troglodytes). We submitted 4- and 5-year-old children (N = 90) and captive chimpanzees (N = 69) to a token–reward exchange task. The results revealed different forms of payoff-biased learning across species and contexts. Specifically, following personal and social exposure to different tokens, children's exchange behaviour was consistent with proportional imitation, where choice is affected by both prior personally acquired and socially demonstrated token–reward information. However, when the socially derived information regarding token value was novel, children's behaviour was consistent with proportional observation; paying attention to socially derived information and ignoring their prior personal experience. By contrast, chimpanzees' token choice was governed by their own prior experience only, with no effect of social demonstration on token choice, conforming to proportional reservation. We also find evidence for individual- and group-level differences in behaviour in both species. Despite the difference in payoff strategies used, both chimpanzees and children adopted beneficial traits when available. However, the strategies of the children are expected to be the most beneficial in promoting flexible behaviour by enabling existing behaviours to be updated or replaced with new and often superior ones.

Conformity to the behavioural preferences of others can have powerful effects on intragroup behavioural homogeneity in humans, but evidence in animals remains minimal. In this study, we took advantage of circumstances in which individuals or pairs of captive chimpanzees, Pan troglodytes, were 'migrated' between groups, to investigate whether immigrants would conform to a new dietary population preference experienced in the group they entered, an effect suggested by recent fieldwork. Such 'migratory-minority' chimpanzees were trained to avoid one of two differently coloured foods made unpalatable, before 'migrating' to, and then observing, a 'localmajority' group consume a different food colour. Both migratory-minority and local-majority chimpanzees displayed social learning, spending significantly more time consuming the previously unpalatable, but instead now edible, food, than did control chimpanzees who did not see immigrants eat this food, nor emigrate themselves. However, following the migration of migratory-minority chimpanzees, these control individuals and the local-majority chimpanzees tended to rely primarily upon personal information, consuming first the food they had earlier learned was palatable before sampling the alternative. Thus, chimpanzees did not engage in conformity in the context we tested; instead seeing others eat a previously unpalatable food led to socially learned and adaptive re-exploration of this now-safe option in both minority and majority participants. Author ContributionsG.L.V. collected the data and carried out the statistical analyses; G.L.V. and A.W. drafted the manuscript with all authors contributing to the final manuscript; the original conception of the methodology was the work of A.W. and E.W.; the original methodology was adapted for captive chimpanzees by G.L.V. and A.W.; S.D. conducted inter-rater reliability, S.P.L. conducted and monitored all chimpanzee 'migrations' and provided on-site logistical support along with S.J.S. Many of the daily choices faced by animals require decisions about whether to engage in personal exploration of the environment (asocial learning) or instead to exploit the existing knowledge of others by learning socially (Kendal, Coolen, van Bergen, & Laland, 2005;Kendal, Coolen, & Laland, 2009;Laland, 2004). Evolutionary theory predicts that if appropriate decision-making rules can be economically employed, social learning will itself be selective. Such selectivity may be pursued through heuristics termed social learning strategies (Laland, 2004), or transmission biases (Boyd & Richerson, 1985;Henrich, 2001), which dictate who, what, when or even how to copy. The identification of such heuristics has proved instructive in understanding how cultures evolve in humans and other species (Kendal et al., 2015;Rendell et al., 2011). A variety of social learning strategies have recently been identified in diverse animal taxa (Kendal et al., 2009;Laland, 2004;Rendell et al., 2011), such as preferentially copying 'dominant' or 'knowledgeable' in...

No abstract