Abstract: Chimpanzees, Pan troglodytes, are the most sophisticated tool-users among all nonhuman primates. From an evolutionary perspective, it is therefore puzzling that the tool use behaviour of their closest living primate relative, the bonobo, Pan paniscus, has been described as particularly poor. However, only a small number of bonobo groups have been studied in the wild and only over comparably short periods. Here, we show that captive bonobos and chimpanzees are equally diverse tool-users in most contexts. Our observations illustrate that tool use in bonobos can be highly complex and no different from what has been described for chimpanzees. The only major difference in the chimpanzee and bonobo data was that bonobos of all age–sex classes used tools in a play context, a possible manifestation of their neotenous nature. We also found that female bonobos displayed a larger range of tool use behaviours than males, a pattern previously described for chimpanzees but not for other great apes. Our results are consistent with the hypothesis that the female-biased tool use evolved prior to the split between bonobos and chimpanzees.

Abstract: Studies of ape tool use have been conducted in captivity since the early 1900s and in the wild since the 1960s. Chimpanzees are the most prolific tool users among the apes, and are known to use more tools than any other nonhuman animal. In contrast, reports of gorilla tool use are rare both in wild and captive settings. Studies of the processes involved in tool use learning have been limited in the wild by the lack of ability to control several unpredictable variables, and in captivity by tool use opportunities that are often presented in non-naturalistic contexts. We attempted to address both of these limitations by providing naïve subjects with a naturalistic tool use device (built to simulate a termite mound) while housed in a more natural social setting to approximate how learning would occur in the wild. Both gorillas and chimpanzees participated in the experiment to allow comparative analyses of acquisition of tool behaviour and the factors that may affect acquisition. Both species showed low frequencies of interaction with the mound in the baseline condition, before baiting with a food reward. Once baited, chimpanzees both attempted and succeeded to extract the reward more quickly than did gorillas. The number of social group members at the mound was significantly higher for chimpanzees than for gorillas and may have affected skill acquisition. We advocate that comparative approaches to skill acquisition and learning are valuable, but that researchers need to be cognizant of species differences in social structure that may affect results.

Abstract: Despite numerous attempts to define animal tool use over the past four decades, the definition remains elusive and the behaviour classification somewhat subjective. Here, we provide a brief review of the definitions of animal tool use and show how those definitions have been modified over time. While some aspects have remained constant (i.e., the distinction between 'true' and 'borderline' tool use), others have been added (i.e., the distinction between 'dynamic' and 'static' behaviours). We present an updated, comprehensive catalog of documented animal tool use that indicates whether the behaviours observed included any 'true' tool use, whether the observations were limited to captive animals, whether tool manufacture has been observed, and whether the observed tool use was limited to only one individual and, thus, 'anecdotal' (i.e., N = 1). Such a catalog has not been attempted since Beck (1980). In addition to being a useful reference for behaviourists, this catalog demonstrates broad tool use and manufacture trends that may be of interest to phylogenists, evolutionary ecologists, and cognitive evolutionists. Tool use and tool manufacture are shown to be widespread across three phyla and seven classes of the animal kingdom. Moreover, there is complete overlap between the Aves and Mammalia orders in terms of the tool use categories (e.g., food extraction, food capture, agonism) arguing against any special abilities of mammals. The majority of tool users, almost 85% of the entries, use tools in only one of the tool use categories. Only members of the Passeriformes and Primates orders have been observed to use tools in four or more of the ten categories. Thus, observed tool use by some members of these two orders (e.g., Corvus, Papio) is qualitatively different from that of all other animal taxa. Finally, although there are similarities between Aves and Mammalia, and Primates and Passeriformes, primate tool use is qualitatively different. Approximately 35% of the entries for this order demonstrate a breadth of tool use (i.e., three or more categories by any one species) compared to other mammals (0%), Aves (2.4%), and the Passeriformes (3.1%). This greater breadth in tool use by some organisms may involve phylogenetic or cognitive differences � or may simply reflect differences in length and intensity of observations. The impact that tool usage may have had on groups' respective ecological niches and, through niche-construction, on their respective evolutionary trajectories remains a subject for future study.

Abstract: Summary Are complex, species-specific behaviors in animals reinforced by material reward alone or do they also induce positive emotions? Many adaptive human behaviors are intrinsically motivated: they not only improve our material outcomes, but improve our affect as well [1, 2, 3, 4, 5, 6, 7, 8]. Work to date on animal optimism, as an indicator of positive affect, has generally focused on how animals react to change in their circumstances, such as when their environment is enriched [9, 10, 11, 12, 13, 14] or they are manipulated by humans [15, 16, 17, 18, 19, 20, 21, 22, 23], rather than whether complex actions improve emotional state. Here, we show that wild New Caledonian crows are optimistic after tool use, a complex, species-specific behavior. We further demonstrate that this finding cannot be explained by the crows needing to put more effort into gaining food. Our findings therefore raise the possibility that intrinsic motivation (enjoyment) may be a fundamental proximate cause in the evolution of tool use and other complex behaviors. Video Abstract

Abstract: To determine whether tool use varied in relation to food availability in bearded capuchin monkeys, we recorded anvil and stone hammer use in two sympatric wild groups, one of which was provisioned daily, and assessed climatic variables and availability of fruits, invertebrates and palm nuts. Capuchins used tools to crack open encased fruits, mostly palm nuts, throughout the year. Significant differences between wet and dry seasons were found in rainfall, abundance of invertebrates and palm nuts, but not in fruit abundance. Catulè nuts were more abundant in the dry season. We tested the predictions of the necessity hypothesis (according to which tool use is maintained by sustenance needs during resource scarcity) and of the opportunity hypothesis (according to which tool use is maintained by repeated exposure to appropriate ecological conditions, such as preferred food resources necessitating the use of tools). Our findings support only the opportunity hypothesis. The rate of tool use was not affected by provisioning, and the monthly rate of tool use was not correlated with the availability of fruits and invertebrates. Conversely, all capuchins cracked food items other than palm nuts (e.g. cashew nuts) when available, and adult males cracked nuts more in the dry season when catulè nuts (the most common and exploited nut) are especially abundant. Hence, in our field site capuchins use tools opportunistically.