Abstract: Innovative problem solving enables individuals to deal with novel social and ecological challenges. However, our understanding of the importance of innovation for animals in their natural habitat is limited because experimental investigations of innovation have historically focused on captive animals. To determine how captivity affects innovation, and whether captive studies of animal innovation suffer from low external validity, we need experimental investigations of innovation in both wild and captive populations of the same species in diverse taxa. Here we inquired whether wild and captive spotted hyaenas differ in their ability to solve the same novel technical problem, and in the diversity of exploratory behaviours they exhibit when first interacting with the problem. Our results suggest that wild and captive populations show important differences in their innovative problem-solving abilities. Captive hyaenas were significantly more successful at solving the novel problem, and significantly more diverse in their initial exploratory behaviour, than were wild hyaenas. We were able to rule out hypotheses suggesting that these differences result from excess energy or time available to captive animals. We conclude that captive hyaenas were more successful because captive individuals were less neophobic and more exploratory than their wild counterparts. These results have important implications for our interpretation of studies on innovative problem solving in captive animals and aid our attempts to gain a broader understanding of the importance of innovation for animals in their natural habitat.

Abstract: Under certain models of animal competition, individuals are expected to gather information about opponent quality in order to determine whether they should fight or withdraw. However, the ability to process complex information differs between individuals and across brain hemispheres: a feature of vertebrate cognition known as lateralization that is not anticipated by contest models. I investigated the relationship between aggressive behaviour and mating success during the fallow deer, Dama dama, rut and a measure of lateralization derived from eye preference during parallel walking. Results show that there was no relationship between the tendency to escalate to fighting or predictability in the tendency to engage in fighting and lateralization. Conversely, there was a quadratic relationship between third-party intervention behaviour and lateralization: the greater the tendency to intervene in ongoing fights the lower the degree of lateralization. However, individuals that showed lateralization for right-eye use were least likely to be targeted by the intervening male; thus lateralization is beneficial in this context because targeted males are highly likely to lose this subsequent encounter. The relationship between lateralization and mating success was also nonlinear: males that showed little evidence for an eye bias during lateral displays had the greatest mating success. Taken together, individuals that showed lateralization benefited from avoiding being targeted after third-party intervention; conversely, individuals that showed little evidence for lateralization actively intervened during ongoing fights and had higher mating success. These results suggest that, although lateralization does appear to confer a fitness advantage on individuals, this is not as extensive as anticipated.

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.

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: Aggression can generate anxiety, create uncertainty about its aftermath and jeopardise social relationships. Postconflict interactions serve as conflict management strategies to mitigate these consequences. Whereas postconflict interactions are well characterized in many animals, their functions are still insufficiently investigated. Four functional hypotheses have been proposed: stress reduction, relationship repair, self-protection and benign intent. We aimed to test these hypotheses in females of a tolerant macaque species, the crested macaque, Macaca nigra, under natural conditions, for three postconflict interactions: reconciliation, affiliation and aggression with third parties. Our results provide meaningful contrasts compared with findings in other species. We found no evidence that aggression had consequences for individuals' behavioural indicators of anxiety, although it increased the likelihood of secondary aggression with third parties. There was little evidence for the stress reduction hypothesis as the occurrence of any of the three postconflict interactions investigated had little effect on the measured behavioural indicators of anxiety. Conflict and dyad characteristics also had limited influence on anxiety. The relationship repair function was only partly validated: dyads with stronger bonds or that exchanged more support did not reconcile more often, but dyads with attributes related to the symmetry, stability and predictability (i.e. security) within relationships did. Patterns of initiation and directionality of postconflict interactions in this study population suggest that reconciliation may constitute the signalling of appeasement and benign intent. Furthermore, we found that aggression towards third parties may serve as a source of self-protection and reassertion of the females' social status. The distinctive pattern of postconflict management strategies revealed in wild female crested macaques appears to be related to their typically tolerant social style. These results demonstrate the usefulness of concomitantly studying aggression, postconflict interactions and their functions, to understand conflict management strategies comprehensively, while taking into account the level of social tolerance characterizing the studied society.