Bernard Thierry

In this paper, I chronicle the Strasbourg population of Tonkean macaques (Macaca tonkeana) over a period of half a century. In 1972, Tonkean macaques were imported from Sulawesi, Indonesia, to eastern France, leading to the establishment of two social groups in the Strasbourg region several years later. Our research team studied the social behavior and cognitive abilities of these Tonkean macaques for four decades. The species is characterized by a high degree of social tolerance. This has proven to be very informative in comparative studies of macaque social behavior, opening a new perspective on the evolution of primate societies. Over the years, the population has grown, and more social groups have been formed. However, the fact that some of the Tonkean macaques were healthy carriers of the herpes B virus led to disagreements over their management and eventually to the elimination of the positive individuals. Many individuals from the Strasbourg population are now kept in sanctuaries, and the number of captive breeding groups is limited. We still have much to learn about Tonkean macaques and there is a need for studies carried out in their native habitat in Sulawesi.

While tool use has been widely reported in non-human animals for food acquisition, the use of tools for drinking has been largely overlooked, with primates being a notable exception. We documented tool-assisted water scooping and drinking in several Balinese long-tailed macaques (Macaca fascicularis), living in Ubud, Indonesia, over a period of four years. We observed repeated tool-assisted water scooping using leaves, nuts, pits, and stones. Our results indicate that this behaviour is associated with manual drinking and can be performed in a playful context. This population habitually engages in a cultural form of stone-assisted object play known as stone handling, and it has an overall propensity to manipulate objects in water. We discuss the relationship between instrumental and non-instrumental object-assisted actions, as well as the possibility for this behaviour to be a tradition in this population. This report offers new insights into the limited literature on tool-assisted drinking in monkeys.

Recruitment is a process by which animals can initiate collective movements: the action of an individual prompts conspecifics to follow. Although it has been hypothesized that animals may be able to intentionally recruit others, there is no experimental evidence of this to date. We tested this hypothesis in two pairs of Tonkean macaques in a situation requiring the subjects to find a food site in a 2800 m² area, and approach the site together to release rewards. Each subject was informed of the location of either highly or little-valued rewards. We recorded attention-action sequences in which an individual checked that his partner was attending to him before moving, and also simple departures (i.e., not preceded by eye contact). Analyses showed that sequences were more often followed by recruitment and leading the partner to a baited site than simple departures were. Moreover, subjects used attention-action sequences more frequently when informed of the location of the highly valued reward. This may be explained by the fact that the more motivated they were by the expected rewards, the more likely they were to actively recruit their partner. No such effect was found when subjects performed simple departures. We conclude that Tonkean macaques are capable of intentional recruitment because the subjects voluntarily behaved with the goal of influencing their partner's movement: they checked that the partner was paying attention to them and prompted him to follow by moving. Such performances can be accounted for either by associative learning or by intentional communication.

Comparative study of the social systems of macaques has revealed correlated variations between species in multiple social traits such as the asymmetry of dominance relationships, preferential treatment of kin, patterns of aggression and reconciliation, modes of socialization, and access to food resources. Macaques can be classified on a scale of four categories of social styles, ranging from the least to the most tolerant species. This led to the development of the covariation framework, which addresses the constraints responsible for the linkages between social traits, and their consequences on the evolution of primate social systems. Decades of research have provided a wealth of information that supports, complements, expands, or challenges the covariation framework. In this article, I review this body of knowledge and consider covariation in its two aspects, that is, as a pattern and as a hypothesis. I first consider the extent to which social styles can be invariant, the strength of correlations between traits, and the possible extension of the framework to nonhuman primates other than macaques. I then discuss how to formulate hypotheses, identify sources of linkage between traits, make predictions about the effects of social constraints, assess the tolerance dimension of social styles, and consider the breaking of linkages between traits. Whereas socioecological studies aim to understand how adaptation to the ecological environment determines the shape of social systems, the covariation framework is a complementary research program that seeks to unravel the internal processes that restrict or channel change in social behavior.

Large-scale studies of individual differences in innovative behavior among nonhuman animals are rare because of logistical difficulties associated with obtaining observational data on a large number of innovative individuals across multiple locations. Here, we take a different approach, using observer ratings to study individual differences in innovative behavior in 127 brown capuchin monkeys (Sapajus [Cebus] sp.) from 15 social groups and 7 facilities. Capuchins were reliably rated by 1 to 7 raters (mean 3.2 ± 1.6 raters/monkey) on a 7-point Likert scale for levels of innovative behavior, task motivation, sociality, and dominance. In a subsample, we demonstrate these ratings are valid: Rated innovation predicted performance on a learning task, rated motivation predicted participation in the task, rated dominance predicted social rank based on win/loss aggressive outcomes, and rated sociality predicted the time that monkeys spent in proximity to others. Across all 127 capuchins, individuals that were rated as being more innovative were significantly younger, more social, and more motivated to engage in tasks. Age, sociality, and task motivation all had independent effects on innovativeness, whereas sex, dominance, and group size were nonsignificant. Our findings are consistent with long-term behavioral observations of innovation in wild white-faced capuchins. Observer ratings may, therefore, be a valid tool for studies of animal innovation. (PsycInfo Database Record (c) 2021 APA, all rights reserved).

It was long assumed that only humans can distinguish the living from the dead. Renewed interest in this question over the last decade has led several authors to assert that non-human primates are also aware of death. We investigate this issue by comparing the behaviours of monkeys and great apes toward helpless conspecifics, basing our analysis on published reports. We first examine the behaviours of mothers towards the body of their dead offspring. They may carry the corpse for days or more before abandoning it. They groom, inspect and protect it, sometimes allowing group members to explore it, and rare cases of cannibalism have been reported. No significant difference is observed in the way that monkeys and great apes treat the bodies of infants. We then examine responses to collapsed (still able to move and react) and inanimate (unresponsive or dead) conspecifics. Monkeys and great apes guard, care for and inspect their helpless partners, and also manipulate and mobilise them. Through these actions, individuals may inform themselves about the state of their partners, test their responsiveness and/or attempt to rouse them. It appears that only chimpanzees and gorillas show violent action such as display behaviours and the rough treatment of bodies. They can also make distress calls, and periods of “stunned silence” sometimes occur in chimpanzees, indicating that they are experiencing intense emotion. Finally, we argue that while both monkeys and great apes detect body dysfunction through the victims’ inability to wake up and move, only great apes can understand that something serious has happened. The signs of emotional disturbance reported in them indicate that they may believe that inanimate conspecifics have entered a state of “dormancy”, meaning that they are unlikely to regain wakefulness. However, there is no evidence that any non-human primates are aware of mortality.