Naïve, adult, captive chimpanzees do not socially learn how to make and use sharp stone tools.

Although once regarded as a unique human feature, tool-use is widespread in the animal kingdom. Some of the most proficient tool-users are our closest living relatives, chimpanzees. These repertoires however consist primarily of tool use, rather than tool manufacture (for later use). Furthermore, most populations of chimpanzees use organic materials, such as sticks and leaves, rather than stones as tools. This distinction may be partly ecological, but it is also important as chimpanzees are often used as models for the evolution of human material culture, the oldest traces of which consist of manufactured sharp stone tools (so-called "flakes"). Thus, examining the conditions (if any) under which chimpanzees may develop flake manufacture and use can provide insight into the drivers of these behaviours in our own lineage. Previous studies on non-human apes' ability to make and use flakes focused on enculturated apes, giving them full demonstrations of the behaviour immediately, without providing social information on the task in a stepwise manner. Here we tested naïve, captive chimpanzees (N = 4; three potentially enculturated and one unenculturated subject) in a social learning experimental paradigm to investigate whether enculturated and/or unenculturated chimpanzees would develop flake making and use after social information of various degrees (including a human demonstration) was provided in a scaffolded manner. Even though social learning opportunities were provided, neither the unenculturated subject nor any of the potentially enculturated subjects made or used flakes, in stark contrast to previous studies with enculturated apes. These data suggest that flake manufacture and use is outside of our tested group of captive chimpanzees' individual and social learning repertoires. It also suggests that high levels of enculturation alongside human demonstrations (and/or training) may be required before captive chimpanzees can develop this behaviour.

Evolved Open-Endedness in Cultural Evolution: A New Dimension in Open-Ended Evolution Research.

The goal of Artificial Life research, as articulated by Chris Langton, is "to contribute to theoretical biology by locating life-as-we-know-it within the larger picture of life-as-it-could-be." The study and pursuit of open-ended evolution in artificial evolutionary systems exemplify this goal. However, open-ended evolution research is hampered by two fundamental issues: the struggle to replicate open-endedness in an artificial evolutionary system and our assumption that we only have one system (genetic evolution) from which to draw inspiration. We argue not only that cultural evolution should be seen as another real-world example of an open-ended evolutionary system but that the unique qualities seen in cultural evolution provide us with a new perspective from which we can assess the fundamental properties of, and ask new questions about, open-ended evolutionary systems, especially with regard to evolved open-endedness and transitions from bounded to unbounded evolution. Here we provide an overview of culture as an evolutionary system, highlight the interesting case of human cultural evolution as an open-ended evolutionary system, and contextualize cultural evolution by developing a new framework of (evolved) open-ended evolution. We go on to provide a set of new questions that can be asked once we consider cultural evolution within the framework of open-ended evolution and introduce new insights that we may be able to gain about evolved open-endedness as a result of asking these questions.

Bone-related behaviours of captive chimpanzees (Pan troglodytes) during two excavating experiments.

After stone tools, bone tools are the most abundant artefact type in the Early Pleistocene archaeological record. That said, they are still relatively scarce, which limits our understanding of the behaviours that led to their production and use. Observations of extant primates constitute a unique source of behavioural data with which to construct hypotheses about the technological forms and repertoires exhibited by our hominin ancestors. We conducted two different experiments to investigate the behavioural responses of two groups of captive chimpanzees (Pan troglodytes; n = 33 and n = 9) to disarticulated, defleshed, ungulate bones while participating in a foraging task aimed at eliciting excavating behaviour. Each chimpanzee group was provided with bone specimens with different characteristics, and the two groups differed in their respective experience levels with excavating plant tools. We found that several individuals from the inexperienced group used the provided bones as tools during the task. In contrast, none of the individuals from the experienced group used bones as excavating tools, but instead continued using plant tools. These chimpanzees also performed non-excavating bone behaviours such as percussion and tool-assisted extraction of organic material from the medullary cavity. Our findings serve as a proof-of-concept that chimpanzees can be used to investigate spontaneous bone tool behaviours such as bone-assisted excavation. Furthermore, our results raise interesting questions regarding the role that bone characteristics, as well as previous tool-assisted excavating experience with other raw materials, might have in the expression of bone tool-assisted excavation.

The method of exclusion (still) cannot identify specific mechanisms of cultural inheritance.

The method of exclusion identifies patterns of distributions of behaviours and/or artefact forms among different groups, where these patterns are deemed unlikely to arise from purely genetic and/or ecological factors. The presence of such patterns is often used to establish whether a species is cultural or not-i.e. whether a species uses social learning or not. Researchers using or describing this method have often pointed out that the method cannot pinpoint which specific type(s) of social learning resulted in the observed patterns. However, the literature continues to contain such inferences. In a new attempt to warn against these logically unwarranted conclusions, we illustrate this error using a novel approach. We use an individual-based model, focused on wild ape cultural patterns-as these patterns are the best-known cases of animal culture and as they also contain the most frequent usage of the unwarranted inference for specific social learning mechanisms. We built a model that contained agents unable to copy specifics of behavioural or artefact forms beyond their individual reach (which we define as "copying"). We did so, as some of the previous inference claims related to social learning mechanisms revolve around copying defined in this way. The results of our model however show that non-copying social learning can already reproduce the defining-even iconic-features of observed ape cultural patterns detected by the method of exclusion. This shows, using a novel model approach, that copying processes are not necessary to produce the cultural patterns that are sometimes still used in an attempt to identify copying processes. Additionally, our model could fully control for both environmental and genetic factors (impossible in real life) and thus offers a new validity check for the method of exclusion as related to general cultural claims-a check that the method passed. Our model also led to new and additional findings, which we likewise discuss.

Free hand hitting of stone-like objects in wild gorillas.

The earliest stone tool types, sharp flakes knapped from stone cores, are assumed to have played a crucial role in human cognitive evolution. Flaked stone tools have been observed to be accidentally produced when wild monkeys use handheld stones as tools. Holding a stone core in hand and hitting it with another in the absence of flaking, free hand hitting, has been considered a requirement for producing sharp stone flakes by hitting stone on stone, free hand percussion. We report on five observations of free hand hitting behavior in two wild western gorillas, using stone-like objects (pieces of termite mound). Gorillas are therefore the second non-human lineage primate showing free-hand hitting behavior in the wild, and ours is the first report for free hand hitting behavior in wild apes. This study helps to shed light on the morphofunctional and cognitive requirements for the emergence of stone tool production as it shows that a prerequisite for free hand percussion (namely, free hand hitting) is part of the spontaneous behavioral repertoire of one of humans' closest relatives (gorillas). However, the ability to combine free hand hitting with the force, precision, and accuracy needed to facilitate conchoidal fracture in free hand percussion may still have been a critical watershed for hominin evolution.

Early knapping techniques do not necessitate cultural transmission.

Early stone tool production, or knapping, techniques are claimed to be the earliest evidence for cultural transmission in the human lineage. Previous experimental studies have trained human participants to knap in conditions involving opportunities for cultural transmission. Subsequent knapping was then interpreted as evidence for a necessity of the provided cultural transmission opportunities for these techniques. However, a valid necessity claim requires showing that individual learning alone cannot lead to early knapping techniques. Here, we tested human participants (N = 28) in cultural isolation for the individual learning of early knapping techniques by providing them with relevant raw materials and a puzzle task as motivation. Twenty-five participants were technique naïve according to posttest questionnaires, yet they individually learned early knapping techniques, therewith producing and using core and flake tools. Early knapping techniques thus do not necessitate cultural transmission of know-how and could likewise have been individually derived among premodern hominins.

Experimental investigation of orangutans' lithic percussive and sharp stone tool behaviours.

Early stone tools, and in particular sharp stone tools, arguably represent one of the most important technological milestones in human evolution. The production and use of sharp stone tools significantly widened the ecological niche of our ancestors, allowing them to exploit novel food resources. However, despite their importance, it is still unclear how these early lithic technologies emerged and which behaviours served as stepping-stones for the development of systematic lithic production in our lineage. One approach to answer this question is to collect comparative data on the stone tool making and using abilities of our closest living relatives, the great apes, to reconstruct the potential stone-related behaviours of early hominins. To this end, we tested both the individual and the social learning abilities of five orangutans to make and use stone tools. Although the orangutans did not make sharp stone tools initially, three individuals spontaneously engaged in lithic percussion, and sharp stone pieces were produced under later experimental conditions. Furthermore, when provided with a human-made sharp stone, one orangutan spontaneously used it as a cutting tool. Contrary to previous experiments, social demonstrations did not considerably improve the stone tool making and using abilities of orangutans. Our study is the first to systematically investigate the stone tool making and using abilities of untrained, unenculturated orangutans showing that two proposed pre-requisites for the emergence of early lithic technologies-lithic percussion and the recognition of sharp-edged stones as cutting tools-are present in this species. We discuss the implications that ours and previous great ape stone tool experiments have for understanding the initial stages of lithic technologies in our lineage.

Captive great apes tend to innovate simple tool behaviors quickly.

Recent studies have highlighted the important role that individual learning mechanisms and different forms of enhancenment play in the acquisition of novel behaviors by naïve individuals. A considerable subset of these studies has focused on tool innovation by our closest living relatives, the great apes, to better undestand the evolution of technology in our own lineage. To be able to isolate the role that individual learning plays in great ape tool innovation, researchers usually employ what are known as baseline tests. Although these baselines are commonly used in behavioral studies in captivity, the length of these tests in terms of number of trials and duration remains unstandarized across studies. To address this methodological issue, we conducted a literature review of great ape tool innovation studies conducted in zoological institutions and compiled various methodological data including the timing of innovation. Our literature review revealed an early innovation tendency in great apes, which was particularly pronounced when simple forms of tool use were investigated. In the majority of experiments where tool innovation took place, this occurred within the first trial and/or the first hour of testing. We discuss different possible sources of variation in the latency to innovate such as testing setup, species and task. We hope that our literature review helps researchers design more data-informed, resource-efficient experiments on tool innovation in our closest living relatives.

Clarifying Misconceptions of the Zone of Latent Solutions Hypothesis: A Response to Haidle and Schlaudt: Miriam Noël Haidle and Oliver Schlaudt: Where Does Cumulative Culture Begin? A Plea for a Sociologically Informed Perspective (Biological Theory 15: 161-174, 2020).

The critical examination of current hypotheses is one of the key ways in which scientific fields develop and grow. Therefore, any critique, including Haidle and Schlaudt's article, "Where Does Cumulative Culture Begin? A Plea for a Sociologically Informed Perspective," represents a welcome addition to the literature. However, critiques must also be evaluated. In their article, Haidle and Schlaudt (Biol Theory 15:161-174, 2020. 10.1007/s13752-020-00351-w; henceforth H&S) review some approaches to culture and cumulative culture in both human and nonhuman primates. H&S discuss the "zone of latent solutions" (ZLS) hypothesis as applied to nonhuman primates and stone-toolmaking premodern hominins. Here, we will evaluate whether H&S's critique addresses its target.

A proof of concept for machine learning-based virtual knapping using neural networks.

Prehistoric stone tools are an important source of evidence for the study of human behavioural and cognitive evolution. Archaeologists use insights from the experimental replication of lithics to understand phenomena such as the behaviours and cognitive capacities required to manufacture them. However, such experiments can require large amounts of time and raw materials, and achieving sufficient control of key variables can be difficult. A computer program able to accurately simulate stone tool production would make lithic experimentation faster, more accessible, reproducible, less biased, and may lead to reliable insights into the factors that structure the archaeological record. We present here a proof of concept for a machine learning-based virtual knapping framework capable of quickly and accurately predicting flake removals from 3D cores using a conditional adversarial neural network (CGAN). We programmatically generated a testing dataset of standardised 3D cores with flakes knapped from them. After training, the CGAN accurately predicted the length, volume, width, and shape of these flake removals using the intact core surface information alone. This demonstrates the feasibility of machine learning for investigating lithic production virtually. With a larger training sample and validation against archaeological data, virtual knapping could enable fast, cheap, and highly-reproducible virtual lithic experimentation.

Naïve orangutans (Pongo abelii and Pongo pygmaeus) individually acquire nut-cracking using hammer tools.

Nut-cracking with hammer tools (henceforth: nut-cracking) has been argued to be one of the most complex tool-use behaviors observed in nonhuman animals. So far, only chimpanzees, capuchins, and macaques have been observed using tools to crack nuts in the wild (Boesch and Boesch, 1990; Gumert et al., 2009; Mannu and Ottoni, 2009). However, the learning mechanisms behind this behavior, and the extent of nut-cracking in other primate species are still unknown. The aim of this study was two-fold. First, we investigated whether another great ape species would develop nut-cracking when provided with all the tools and appropriate conditions to do so. Second, we examined the mechanisms behind the emergence of nut-cracking by testing a naïve sample. Orangutans (Pongo abelii and Pongo pygmaeus) have the second most extensive tool-use repertoire among the great apes (after chimpanzees) and show flexible problem-solving capacities. Orangutans have not been observed cracking nuts in the wild, however, perhaps because their arboreal habits provide limited opportunities for nut-cracking. Therefore, orangutans are a valid candidate species for the investigation of the development of this behavior. Four nut-cracking-naïve orangutans at Leipzig zoo (P. abelii; Mage = 16; age range = 10-19; 4F; at the time of testing) were provided with nuts and hammers but were not demonstrated the nut-cracking behavioral form. Additionally, we report data from a previously unpublished study by one of the authors (Martina Funk) with eight orangutans housed at Zürich zoo (six P. abelii and two P. pygmaeus; Mage = 14; age range = 2-30; 5F; at the time of testing) that followed a similar testing paradigm. Out of the twelve orangutans tested, at least four individuals, one from Leipzig (P. abelii) and three from Zürich (P. abelii and P. pygmaeus), spontaneously expressed nut-cracking using wooden hammers. These results demonstrate that nut-cracking can emerge in orangutans through individual learning and certain types of non-copying social learning.

Evaluating the influence of action- and subject-specific factors on chimpanzee action copying.

The ability to imitate has been deemed crucial for the emergence of human culture. Although non-human animals also possess culture, the acquisition mechanisms underlying behavioural variation between populations in other species is still under debate. It is especially controversial whether great apes can spontaneously imitate. Action- and subject-specific factors have been suggested to influence the likelihood of an action to be imitated. However, few studies have jointly tested these hypotheses. Just one study to date has reported spontaneous imitation in chimpanzees (Persson et al. 2017 Primates 59, 19-29), although important methodological limitations were not accounted for. Here, we present a study in which we (i) replicate the above-mentioned study addressing their limitations in an observational study of human-chimpanzee imitation; and (ii) aim to test the influence of action- and subject-specific factors on action copying in chimpanzees by providing human demonstrations of multiple actions to chimpanzees of varying rearing background. To properly address our second aim, we conducted a preparatory power analysis using simulated data. Contrary to Persson et al.'s study, we found extremely low rates of spontaneous chimpanzee imitation and we did not find enough cases of action matching to be able to apply our planned model with sufficient statistical power. We discuss possible factors explaining the lack of observed action matching in our experiments compared with previous studies.

The Method of Local Restriction: in search of potential great ape culture-dependent forms.

Humans possess a perhaps unique type of culture among primates called cumulative culture. In this type of culture, behavioural forms cumulate changes over time, which increases their complexity and/or efficiency, eventually making these forms culture-dependent. As changes cumulate, culture-dependent forms become causally opaque, preventing the overall behavioural form from being acquired by individuals on their own; in other words, culture-dependent forms must be copied between individuals and across generations. Despite the importance of cumulative culture for understanding the evolutionary history of our species, how and when cumulative culture evolved is still debated. One of the challenges faced when addressing these questions is how to identify culture-dependent forms that result from cumulative cultural evolution. Here we propose a novel method to identify the most likely cases of culture-dependent forms. The 'Method of Local Restriction' is based on the premise that as culture-dependent forms are repeatedly transmitted via copying, these forms will unavoidably cumulate population-specific changes (due to copying error) and therefore must be expected to become locally restricted over time. When we applied this method to our closest living relatives, the great apes, we found that most known ape behavioural forms are not locally restricted (across domains and species) and thus are unlikely to be acquired via copying. Nevertheless, we found 25 locally restricted forms across species and domains, three of which appear to be locally unique (having been observed in a single population of a single species). Locally unique forms represent the best current candidates for culture-dependent forms in non-human great apes. Besides these rare exceptions, our results show that overall, ape cultures do not rely heavily on copying, as most ape behaviours appear across sites and/or species, rendering them unlikely to be culture-dependent forms resulting from cumulative cultural evolution. Yet, the locally restricted forms (and especially the three locally unique forms) identified by our method should be tested further for their potential reliance on copying social learning mechanisms (and in turn, for their potential culture-dependence). Future studies could use the Method of Local Restriction to investigate the existence of culture-dependent forms in other animal species and in the hominin archaeological record to estimate how widespread copying is in the animal kingdom and to postulate a timeline for the emergence of copying in our lineage.

Naïve, unenculturated chimpanzees fail to make and use flaked stone tools.

Background: Despite substantial research on early hominin lithic technologies, the learning mechanisms underlying flake manufacture and use are contested. To draw phylogenetic inferences on the potential cognitive processes underlying the acquisition of both of these abilities in early hominins, we investigated if and how one of our closest living relatives, chimpanzees ( Pan troglodytes), could learn to make and use flakes. Methods: Across several experimental conditions, we tested eleven task-naïve chimpanzees (unenculturated n=8, unknown status n=3) from two independent populations for their abilities to spontaneously make and subsequently use flakes as well as to use flakes made by a human experimenter. Results: Despite the fact that the chimpanzees seemed to understand the requirements of the task, were sufficiently motivated and had ample opportunities to develop the target behaviours, none of the chimpanzees tested made or used flakes in any of the experimental conditions. Conclusions: These results differ from all previous ape flaking experiments, which found flake manufacture and use in bonobos and one orangutan. However, these earlier studies tested human-enculturated apes and provided test subjects with flake making and using demonstrations. The contrast between these earlier positive findings and our negative findings (despite using a much larger sample size) suggests that enculturation and/or demonstrations may be necessary for chimpanzees to acquire these abilities. The data obtained in this study are consistent with the hypothesis that flake manufacture and use might have evolved in the hominin lineage after the split between Homo and Pan 7 million years ago, a scenario further supported by the initial lack of flaked stone tools in the archaeological record after this split. We discuss possible evolutionary scenarios for flake manufacture and use in both non-hominin and hominin lineages.

Chimpanzees' (Pan troglodytes) problem-solving skills are influenced by housing facility and captive care duration.

Although a large body of primate cognition research is done in captive institutions, little is known about how much individuals from different facilities vary in their experiences and cognitive skills. Here we present the results of an experimental study investigating how physical cognitive skills vary between chimpanzees in relation to captive settings and their time in captivity. We tested 59 chimpanzees housed at two different captive facilities (a rehabilitation center (sanctuary) and a zoo) in three problem-solving tasks. Our results showed that chimpanzees at the two housing facilities significantly differed in overall task performance. On average, the sanctuary chimpanzees outperformed the chimpanzees housed at the zoo in the detour reaching task and the honey trap task. However, the zoo chimpanzees performed slightly better on average in the learning task. We propose that, for this particular sample, the documented differences result from a combination of factors, such as prior experience with cognitive testing, motivation levels and varying degrees of human exposure. Within the sanctuary sample, we found that chimpanzees who arrived at an earlier age at the sanctuary and had therefore spent a larger percentage of their lives in a captive environment, were better problem-solvers than those that arrived at a later age to the sanctuary. Thus, rehabilitation and time in captivity contributed to improved physical cognitive skills in sanctuary chimpanzees. Our results highlight the importance of studying intraspecific variation and the effect that previous experience and living conditions might have on physical cognitive skills in non-human apes. Accordingly, we should be cautious when extrapolating findings of cognitive studies from one population to the species as a whole.

The zone of latent solutions and its relevance to understanding ape cultures.

The zone of latent solutions (ZLS) hypothesis provides an alternative approach to explaining cultural patterns in primates and many other animals. According to the ZLS hypothesis, non-human great ape (henceforth: ape) cultures consist largely or solely of latent solutions. The current competing (and predominant) hypothesis for ape culture argues instead that at least some of their behavioural or artefact forms are copied through specific social learning mechanisms ("copying social learning hypothesis") and that their forms may depend on copying (copying-dependent forms). In contrast, the ape ZLS hypothesis does not require these forms to be copied. Instead, it suggests that several (non-form-copying) social learning mechanisms help determine the frequency (but typically not the form) of these behaviours and artefacts within connected individuals. The ZLS hypothesis thus suggests that increases and stabilisations of a particular behaviour's or artefact's frequency can derive from socially-mediated (cued) form reinnovations. Therefore, and while genes and ecology play important roles as well, according to the ape ZLS hypothesis, apes typically acquire the forms of their behaviours and artefacts individually, but are usually socially induced to do so (provided sufficient opportunity, necessity, motivation and timing). The ZLS approach is often criticized-perhaps also because it challenges the current null hypothesis, which instead assumes a requirement of form-copying social learning mechanisms to explain many ape behavioural (and/or artefact) forms. However, as the ZLS hypothesis is a new approach, with less accumulated literature compared to the current null hypothesis, some confusion is to be expected. Here, we clarify the ZLS approach-also in relation to other competing hypotheses-and address misconceptions and objections. We believe that these clarifications will provide researchers with a coherent theoretical approach and an experimental methodology to examine the necessity of form-copying variants of social learning in apes, humans and other species.