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Different operationalizations of culture. The cultural behaviors captured by the MoE (C ME ) are socially learned behaviors with a patchy geographic distribution but without ecological correlates (mostly conspicuous and/or high complexity behaviors such as tool use). The cultural behaviors with ecological correlates (C Ecol ) are socially learned behaviors that vary between populations because they are influenced by a population's local ecology (e.g. feeding skills). The sum of C ME and C Ecol are all socially learned behaviors that vary across populations (C Var ). Cultural universals (C U ) are socially learned behaviors and knowledge that we find consistently across populations (e.g. basic subsistence and social skills). The sum of all socially learned behaviors represents an individual's cultural knowledge (C 1 = C Var + C U ). See supplementary Table S3 descriptions of the behaviors depicted on the pictures.
Source publication
For humans we implicitly assume that the way we do things is the product of social learning and thus cultural. For animals, this conclusion requires proof. Here, we first review the most commonly used procedure for documenting animal culture: the method of exclusion, which charts geographic behavioral variation between populations as evidence for c...
Contexts in source publication
Context 1
... to the minimal definition of culture ( Boyd and Richerson, 1985), an individual's cultural repertoire is the sum of all socially learned skills and knowledge (C 1 =∑SLS; Figure 3). According to the population-level concept of culture (culture as all behaviors that show interpopulation geographic variation), it is the sum of all SLS minus the sum of all cultural behaviors shared between the compared populations (C Var = ∑C 1 − ∑C U , Figure 4). The difference between these two measures of cultural repertoire sizes is, among other factors, dependent on the innovation rate in relation to transmission efficiency. ...
Context 2
... strength of ecological influences on culture is a positive function of the difference we get from subtracting the cultural repertoires captured by the MoE (C ME ) from the sum of all socially learned behaviors that show geographic variation (C Var ), i.e. C Var -C ME = ∑C Ecol (Figure 4). A large difference implies that a lot of cultural variation is ecologically induced. ...
Context 3
... base of the great ape culture iceberg The orangutan example suggests that by relying on the MoE to assess cultural repertoires we have so far only discovered the tip of the great ape culture iceberg (i.e. C 1 >> C ME ; Figure 4). The MoE produces a biased sample of highly complex and conspicuous behaviors and dismisses a vast array of socially learned behaviors that covary with ecological factors. ...
Citations
... However, even if ecological factors are causing the behavioral variation between UNP and SCaNP, we cannot rule out the possibility that this technique is a cultural trait in UNP. Cultural traits are acquired by socially biased learning mechanisms, and environmental factors can affect the emergence or maintenance of these cultural traits [26][27][28] . At UNP, the immatures closely observed proficient individuals, manipulated stones, sticks, and the operculum, and explored spider burrows. ...
Primates employ different tools and techniques to overcome the challenges of obtaining underground food resources. Humans and chimpanzees are known to tackle this problem with stick tools and one population of capuchin monkeys habitually uses stone tools. Although early hominids could have used stones as digging tools, we know little about when and how these could be useful. Here, we report a second primate population observed using stone tools and the first capuchin monkey population to habitually use the ‘stick-probing’ technique for obtaining underground resources. The bearded capuchin monkeys (Sapajus libidinosus) from Ubajara National Park, Brazil, use ‘hands-only’ and ‘stone-digging’ techniques for extracting underground storage organs and trapdoor spiders. Males also use ‘stick-probing’ and ‘stone-stick’ techniques for capturing trapdoor spiders. Tool use does not increase success in obtaining these resources. Stone-digging is less frequent in this population than in the only other known population that uses this technique. Females use stones in a lower proportion of their digging episodes than males in both populations. Ecological and cultural factors potentially influence technique choice and sex differences within and between populations. This population has a different pattern of underground food exploration using tools. Comparing this population with others and exploring the ecological and cultural factors under which capuchin monkeys employ different tools and techniques will allow us to better understand the pressures that may have shaped the evolution of those behaviors in primates.
... Culture, or shared socially learned traits persisting across generations, is observed in various taxa from birds to cetaceans and primates (Rendell and Whitehead 2001;White-head et al. 2004;Aplin 2019;Schuppli and van Schaik 2019;Whiten 2019Whiten , 2021. Cultural traits are inherited through social transmission and may compete with each other (Galef 1995;O'Brien and Lyman 2000;Mesoudi et al. 2004), either for expression when traits are not mutually exclusive or for hosts when traits are mutually exclusive. ...
A change to a population’s social network is a change to the substrate of cultural transmission, affecting behavioral diversity and adaptive cultural evolution. While features of network structure such as population size and density have been well studied, less is understood about the influence of social processes such as population turnover—or the repeated replacement of individuals by naive individuals. Experimental data have led to the hypothesis that naive learners can drive cultural evolution by better assessing the relative value of behaviors, although this hypothesis has been expressed only verbally. We conducted a formal exploration of this hypothesis using a generative model that concurrently simulated its two key ingredients: social transmission and reinforcement learning. We simulated competition between high- and low-reward behaviors while varying turnover magnitude and tempo. Variation in turnover influenced changes in the distributions of cultural behaviors, irrespective of initial knowledge-state conditions. We found optimal turnover regimes that amplified the production of higher reward behaviors through two key mechanisms: repertoire composition and enhanced valuation by agents that knew both behaviors. These effects depended on network and learning parameters. Our model provides formal theoretical support for, and predictions about, the hypothesis that naive learners can shape cultural change through their enhanced sampling ability. By moving from experimental data to theory, we illuminate an underdiscussed generative process that can lead to changes in cultural behavior, arising from an interaction between social dynamics and learning.
... In this paper, we therefore refer to these aspects of nest building as 'multistep nest features'. During the nest-building process, individuals may also regularly produce vocalizations (so called 'nest sounds'), which are more likely to be produced during night nest construction (Hardus et al., 2009;Paul, 2007;Prasetyo et al., 2009;Schuppli & van Schaik, 2019;Wich et al., 2012). Night nest construction is arguably more critical than day nest construction because night nests are used for a longer Table 1 Definitions of orang-utan behaviours and nest elements ...
Keywords: needing to learn hypothesis nest-building behaviour ontogeny selectivity skill learning Sumatran orang-utan trial-and-error practice Nest building is an important subsistence behaviour that young great apes must learn to become competent adults. Orang-utans show a remarkable degree of variability and selectivity for a broad range of features in their nest building. However, the details of when different aspects of nest-building skills emerge remain unclear. We used data on 27 immature Sumatran orang-utans and 20 mothers collected over a decade at Suaq Balimbing, Sumatra to investigate when immatures develop their nest-building skills and examine when nest tree species preferences emerge. We found that young orang-utans showed interest in nest building from 6 months of age and begin to construct day nests at around 1 year of age, whereas night nests were not practised until close to the third year of life. Nest-building practice peaked around age 3e4 years and then steadily decreased as immatures approached the age of nutritional independence, around age 8 years. By then, immature orang-utans were competent nest-builders, but their nests differed from adult nests in several aspects, such as fewer multitree nests and additional comfort elements, which seemed to be mastered later in development. All age classes demonstrated stronger selectivity towards tree species used for night nests and immatures eventually had similar preferences to mothers. We conclude that the ontogeny of nest-building behaviour and the selection of nest tree species in Sumatran orang-utans is a multiyear learning process that requires intense practice.
... However, while the method of exclusion allows cata loging and identifying cultural traits, ge ne tic and ecological factors are often difficult to entirely exclude (Laland and Hoppitt 2003;Schuppli and van Schaik 2019). For example, subsequent studies on ant-dipping techniques in chimpanzees revealed that the performed variants are influenced by the nature of the ants' be hav ior (Humle and Matsuzawa 2002). ...
Non-human animal (henceforth animal) culture has been demonstrated across a variety of species and, for some, includes technical traditions. Like human culture, animal culture must strike a balance between flexible and rigid behaviour. On one hand, culture must allow for the creation, acquisition, or application of cultural behaviours. On the other hand, culture must ensure a high degree of similarity between individual performances, leading to stable traditions that persist across generations despite disruptive influences. The dominant argument for understanding the rigidity of cultural transmission within animals has focused on social learning, with mechanisms such as imitation leading to the highest fidelity of transmission. Thereby animal culture has often been limited to examples explained by social learning alone and has historically excluded behaviours that could be explained by genetic or ecological factors. In this chapter we argue that these factors, rather than be excluded, may assist social learning in explaining cultural differences. To explore these concepts, we will examine and define flexible and rigid behaviour at the individual level, then expand to the group and population level. We then describe the current perspectives from social learning theory as well as cultural attraction theory in understanding animal cultural behaviour, highlighting the objectives and limitations of each. Further, we propose a framework that combines social learning and cultural attraction theory to understand the origin and stability of chimpanzee cultural techniques. We argue that cultural traits are the product of three classes of factors of attraction: individual predispositions, social influences, and ecological contexts. This new framework can contribute to better understanding the origins, persistence, and eventual demise of cultural techniques.
... However, while the method of exclusion allows cata loging and identifying cultural traits, ge ne tic and ecological factors are often difficult to entirely exclude (Laland and Hoppitt 2003;Schuppli and van Schaik 2019). For example, subsequent studies on ant-dipping techniques in chimpanzees revealed that the performed variants are influenced by the nature of the ants' be hav ior (Humle and Matsuzawa 2002). ...
A novel, interdisciplinary exploration of the relative contributions of rigidity and flexibility in the adoption, maintenance, and evolution of technical traditions.
Techniques can either be used in rigid, stereotypical ways or in flexibly adaptive ways, or in some combination of the two. The Evolution of Techniques, edited by Mathieu Charbonneau, addresses the impacts of both flexibility and rigidity on how techniques are used, transformed, and reconstructed, at varying social and temporal scales. The multidisciplinary contributors demonstrate the important role of the varied learning contexts and social configurations involved in the transmission, use, and evolution of techniques. They explore the diversity of cognitive, behavioral, sociocultural, and ecological mechanisms that promote and constrain technical flexibility and rigidity, proposing a deeper picture of the enablers of, and obstacles to, technical transmission and change.
In line with the extended evolutionary synthesis, the book proposes a more inclusive and materially grounded conception of technical evolution in terms of promiscuous, dynamic, and multidirectional causal processes. Offering new evidence and novel theoretical perspectives, the contributors deploy a diversity of methods, including ethnographies, field and laboratory experiments, cladistics and phylogenetic tree building, historiography, and philosophical analysis. Examples of the wide range of topics covered include field experiments with potters from five cultures, stability and change in Paleolithic toolmaking, why children lack flexibility when making tools, and cultural techniques in nonhuman animals.
The volume's three thematic sections are:
• Timescales of technical rigidity and flexibility
• Rigid copying to flexible reconstruction
• Exogenous factors of technical rigidity and flexibility
The volume closes with a discussion by philosopher Kim Sterelny.
Contributors
Rita Astuti, Adam Howell Boyette, Blandine Bril, Josep Call, Mathieu Charbonneau, Arianna Curioni, Nicola Cutting, Bert De Munck, György Gergely, Anne-Lise Goujon, Ildikó Király, Catherine Lara, Sébastien Manem, Luke McEllin, Helena Miton, Giulio Ongaro, Sarah Pope-Caldwell, Valentine Roux, Manon Schweinfurth, Dan Sperber, Kim Sterelny, Dietrich Stout, James W. A. Strachan, Sadie Tenpas
... Additionally, the careful study required to document culture with certainty may be unethical or impossible with endangered species (Brakes et al., 2021b), and restrictive or exclusionary de nitions are likely to result in large underestimates of the prevalence of cultural behaviours (Schuppli & van Schaik, 2019). Therefore I will refer to any socially learned, group-speci c behaviour as culture (Laland & Hoppitt, 2003;Laland & Janik, 2006;Whitehead et al., 2004;Whiten et al., 2017). ...
This handbook is currently in development, with individual articles publishing online in advance of print publication. At this time, we cannot add information about unpublished articles in this handbook, however the table of contents will continue to grow as additional articles pass through the review process and are added to the site. Please note that the online publication date for this handbook is the date that the first article in the title was published online. For more information, please read the site FAQs.
... From the early observation of one group of Japanese macaques on Koshima Island washing sweet potatoes in the water before eating them (Imanishi, 1957), scientists then came to identify culture in chimpanzees (Whiten et al., 1999), whales and dolphins (Whitehead & Rendell, 2014), and birds (Aplin, 2018). Today a number of scientists now endorse the idea that there is culture in a large number of animal species (Schuppli & van Schaik, 2019;Whiten, 2019Whiten, , 2021, including fish (Mathis et al., 1996), bumble bees (Alem et al., 2016), and fruit flies (Danchin et al., 2018). If the function of over-imitation is to learn the cultural affordances and social norms of one's cultural group, and many species have culturally distinct behaviors, then there is reason to ask whether some version of overimitation is found in these other species as well. ...
Human culture is seen as more cumulative, cooperative, and normative, in contrast to animal cultures. One hypothesis to explain these differences is the over-imitation hypothesis—that the differences between human culture and animal cultures can be traced to the human unique tendency to over-imitate. In this paper we analyze the current state of the literature on animal over-imitation and challenge the adequacy of the over-imitation hypothesis. To make this argument, we first argue that the function of human over-imitation is norm-learning. Then we review the empirical evidence against animal over-imitation and argue that these studies do not take into account relevant variables given the normative nature of over-imitation. We then analyze positive empirical evidence of over-imitation in great apes and canids and conclude they may have some capacity for over-imitation. In addition to the methodological suggestion for how to study animal over-imitation, a theoretical suggestion is that over-imitation might be much more widely found among species. The larger implication is that if we do find widespread evidence of over-imitation across species, many of the current theories of human uniqueness that focus on human hyper-cooperation or social norms may have only identified a difference of degree between humans and other animals.
... Ethological research has produced studies on the pace of animal innovations, as found in seasonal changes in songs of humpback whales (Schulze et al., 2022), on cultural transmission in the manufacture of tools, as in New Caledonian crows (Hunt & Gray, 2003), and on adoption of new behavioural elements from resident individuals when they must adapt to new habitats after dispersal, as in male orangutans (Mörchen et al., 2023). The current state of the field suggests there is a variety of basic cultural performances in a wide array of animal orders from invertebrates to birds and mammals (Whiten, 2021) and that the number of (unspecified) cases of 'culture' is underestimated (Schuppli & Van Schaik, 2019 In contrast to field studies on animal behaviour and ethnography, which include direct observations of performances, primary data from archaeological field studies provide only indirect insights into past human behaviour through trace material evidence. Performances, and thus also group-specific patterns and their development, must be inferred from finds via multiple analytical steps before interpretation (Perrault, 2019). ...
The rapid growth of cultural evolutionary science, its expansion into numerous fields, its use of diverse methods, and several conceptual problems have outpaced corollary developments in theory and philosophy of science. This has led to concern, exemplified in results from a recent survey conducted with members of the Cultural Evolution Society, that the field lacks ‘knowledge synthesis’, is poorly supported by ‘theory’, has an ambiguous relation to biological evolution, and uses key terms (e.g. ‘culture’, ‘social learning’, ‘cumulative culture’) in ways that hamper operationalization in models, experiments, and field studies. Though numerous review papers in the field represent and categorize its empirical findings, the field's theoretical challenges receive less critical attention even though challenges of a theoretical or conceptual nature underlie most of the problems identified by Cultural Evolution Society members. Guided by the heterogeneous ‘grand challenges’ emergent in this survey, this paper restates those challenges and adopts an organizational style requisite to discussion of them. The paper's goal is to contribute to increasing conceptual clarity and theoretical discernment around the most pressing challenges facing the field of cultural evolutionary science. It will be of most interest to cultural evolutionary scientists, theoreticians, philosophers of science, and interdisciplinary researchers.
... 61 In contrast to other studies, by investigating peering behavior as an indicator of social learning, we analyzed social learning by directly looking at the moment of potential knowledge transmission, rather than indirectly by studying its potential results. 73 This allowed us to narrow down the mechanism underlying social transmission. The setup of our study allowed us first to differentiate between the immediate proximate effects of food availability on the utilization of social learning opportunities and second to assess the effects of experienced food availability on the development of social learning propensities, even though differentiating between developmental and genetic effects was not possible. ...
Humans’ colonization of diverse habitats relied on our ancestors' abilities to innovate and share innovations with others. While ecological impacts on innovations are well studied, their effect on social learning remains poorly understood. We examined how food availability affects social learning in migrant orangutan unflanged males, who may learn from local orangutans through peering (i.e., observational social learning). We analyzed 1,384 dyadic associations, including 360 peering events, among 46 wild Sumatran orangutan and 25 Bornean orangutan males, collected over 18 years. Migrants’ peering rates significantly increased with higher food availability and time spent in proximity to others. Furthermore, migrants in the more sociable Sumatran population exhibited significantly higher peering rates compared to the Borneans, suggesting intrinsic and/or developmental effects of food availability on social learning. These findings emphasize the importance of investigating ecological effects on social learning on the immediate, developmental, and intrinsic levels for our understanding of cultural evolution.
... This is probably the most commonly used definition in the animal culture literature [e.g. van Schaik et al., 2003;Laland & Janik, 2006;Koops et al., 2015;; although see Schuppli & van Schaik (2019) and Hopper & Carter (2020) for important qualifications about excluding environmental differences]. ...
Social norms – rules governing which behaviours are deemed appropriate or inappropriate within a given community – are typically taken to be uniquely human. Recently, this position has been challenged by a number of philosophers, cognitive scientists, and ethologists, who have suggested that social norms may also be found in certain non-human animal communities. Such claims have elicited considerable scepticism from norm cognition researchers, who doubt that any non-human animals possess the psychological capacities necessary for normative cognition. However, there is little agreement among these researchers about what these psychological prerequisites are. This makes empirical study of animal social norms difficult, since it is not clear what we are looking for and thus what should count as behavioural evidence for the presence (or absence) of social norms in animals. To break this impasse, we offer an approach that moves beyond contested psychological criteria for social norms. This approach is inspired by the animal culture research program, which has made a similar shift away from heavily psychological definitions of ‘culture’ and to become organized around a cluster of more empirically tractable concepts of culture. Here, we propose an analogous set of constructs built around the core notion of a normative regularity, which we define as a socially maintained pattern of behavioural conformity within a community. We suggest methods for studying potential normative regularities in wild and captive primates. We also discuss the broader scientific and philosophical implications of this research program with respect to questions of human uniqueness, animal welfare and conservation.
