Jorg J. M. Massen

A behavioral view on chimpanzee personality: Exploration tendency, persistence, boldness, and tool‐orientation measured with group experiments

Human and nonhuman animals show personality: temporal and contextual consistency in behavior patterns that vary among individuals. In contrast to most other species, personality of chimpanzees, Pan troglodytes, has mainly been studied with non‐behavioral methods. We examined boldness, exploration tendency, persistence and tool‐orientation in 29 captive chimpanzees using repeated experiments conducted in an ecologically valid social setting. High temporal repeatability and contextual consistency in all these traits indicated they reflected personality. In addition, Principal Component Analysis revealed two independent syndromes, labeled exploration‐persistence and boldness. We found no sex or rank differences in the trait scores, but the scores declined with age. Nonetheless, there was considerable inter‐individual variation within age‐classes, suggesting that behavior was not merely determined by age but also by dispositional effects. In conclusion, our study complements earlier rating studies and adds new traits to the chimpanzee personality, thereby supporting the existence of multiple personality traits among chimpanzees. We stress the importance of ecologically valid behavioral research to assess multiple personality traits and their association, as it allows inclusion of ape studies in the comparison of personality structures across species studied behaviorally, and furthers our attempts to unravel the causes and consequences of animal personality. Am. J. Primatol. 75:947–958, 2013.

Ravens notice dominance reversals among conspecifics within and outside their social group

A core feature of social intelligence is the understanding of third-party relations, which has been experimentally demonstrated in primates. Whether other social animals also have this capacity, and whether they can use this capacity flexibly to, for example, also assess the relations of neighbouring conspecifics, remains unknown. Here we show that ravens react differently to playbacks of dominance interactions that either confirm or violate the current rank hierarchy of members in their own social group and of ravens in a neighbouring group. Therefore, ravens understand third-party relations and may deduce those not only via physical interactions but also by observation.

Male Yawning Is More Contagious than Female Yawning among Chimpanzees (Pan troglodytes)

Yawn contagion is not restricted to humans and has also been reported for several non-human animal species, including chimpanzees (Pan troglodytes). Contagious yawning may lead to synchronisation of behaviour. However, the function of contagious yawning is relatively understudied. In this study, we investigated the function of contagious yawning by focusing on two types of signal providers: close social associates and leaders. We provided a captive chimpanzee colony with videos of all individuals of their own group that were either yawning, or at rest. Consistent with other studies, we demonstrated that yawning is contagious for chimpanzees, yet we did not find any effect of relationship quality on yawn contagion. However, we show that yawn contagion is significantly higher when the video model is a yawning male than when the video model was a yawning female, and that this effect is most apparent among males. As males are dominant in chimpanzee societies, male signals may be more relevant to the rest of the group than female signals. Moreover, since chimpanzees form male-bonded societies, male signals are especially relevant for other males. Therefore, we suggest that the sex-differences of yawning contagion among chimpanzees reflect the function of yawning in the synchronisation of behaviour.

Tolerance and reward equity predict cooperation in ravens (Corvus corax).

Cooperative decision rules have so far been shown experimentally mainly in mammal species that have variable and complex social networks. However, these traits should not necessarily be restricted to mammals. Therefore, we tested cooperative problem solving in ravens. We showed that, without training, nine ravens spontaneously cooperated in a loose-string task. Corroborating findings in several species, ravens’ cooperative success increased with increasing inter-individual tolerance levels. Importantly, we found this in both a forced dyadic setting, and in a group setting where individuals had an open choice to cooperate with whomever. The ravens, moreover, also paid attention to the resulting reward distribution and ceased cooperation when being cheated upon. Nevertheless, the ravens did not seem to pay attention to the behavior of their partners while cooperating, and future research should reveal whether this is task specific or a general pattern. Given their natural propensity to cooperate and the results we present here, we consider ravens as an interesting model species to study the evolution of, and the mechanisms underlying cooperation.

Ravens Intervene in Others’ Bonding Attempts

Summary The competition for power in a complex social world is hypothesized to be a driving force in the evolution of intelligence [1]. More specifically, power may be obtained not only by brute force but also by social strategies resembling human politics [2]. Most empirical evidence comes from primate studies that report unprovoked aggression by dominants to maintain power by spreading fear [3] and third-party interventions in conflicts [4–6]. Coalitionary support has also been described in other animals [7, 8] and is often linked to social bonding [9, 10]. As coalitions can lead to a gain in power [5, 11] and fitness benefits [12], individuals may try to prevent coalitionary support or indirectly prevent others from forming social bonds that might lead to coalitions. Although there is some empirical evidence that coalitionary support can be manipulated [13], little is known about the indirect strategy. We show here that wild ravens (Corvus corax) regularly intervene in affiliative interactions of others even though such interventions are potentially risky and without immediate benefits. Moreover, the identities of both interveners and intervened pairs are not randomly distributed. Ravens with existing ties initiate most interventions, and ravens that are creating new ties are most likely to be the targets of interventions. These patterns are consistent with the idea that interventions function to prevent others from forming alliances and consequently becoming future competitors. We thus show previously undescribed social maneuvers in the struggle for power. These maneuvers are likely to be of importance in other social species as well.

A thermal window for yawning in humans: Yawning as a brain cooling mechanism

The thermoregulatory theory of yawning posits that yawns function to cool the brain in part due to counter-current heat exchange with the deep inhalation of ambient air. Consequently, yawning should be constrained to an optimal thermal zone or range of temperature, i.e., a thermal window, in which we should expect a lower frequency at extreme temperatures. Previous research shows that yawn frequency diminishes as ambient temperatures rise and approach body temperature, but a lower bound to the thermal window has not been demonstrated. To test this, a total of 120 pedestrians were sampled for susceptibly to self-reported yawn contagion during distinct temperature ranges and seasons (winter: 1.4°C, n=60; summer: 19.4°C, n=60). As predicted, the proportion of pedestrians reporting yawning was significantly lower during winter than in summer (18.3% vs. 41.7%), with temperature being the only significant predictor of these differences across seasons. The underlying mechanism for yawning in humans, both spontaneous and contagious, appears to be involved in brain thermoregulation.

Memory, transmission and persistence of alternative foraging techniques in wild common marmosets.

Experimental studies on traditions in animals have focused almost entirely on the initial transmission phase in captive populations. We conducted an open diffusion field experiment with 13 groups of wild common marmosets, Callithrix jacchus. Seven groups contained individuals that were already familiar with the task (‘push or pull’ box) and thus served as potential models for naïve individuals. Additionally, in four groups one individual was trained for one of the two possible techniques and in two control groups no skilled individuals were present. First, we investigated whether experienced individuals would remember how to solve the task even after 2 years without exposure and whether they would still prefer their learned technique. Second, we tested whether naïve individuals would learn socially from their skilled family members and, more importantly, whether they would use the same technique. Third, we conducted several test blocks to see whether the individual and/or group behaviour would persist over time. Our results show that wild common marmosets were able to memorize, learn socially and maintain preferences of foraging techniques. This field experiment thus reveals a promising approach to studying social learning in the wild and provides the basis for long-term studies on tradition formation.

Stability and Durability of Intra- and Intersex Social Bonds of Captive Rhesus Macaques ( Macaca mulatta )

Many social species maintain differentiated relationships with their group members. These social bonds may reflect short-term interactions satisfying immediate needs. Alternatively, individuals of some species may entertain stable and valuable bonds that last much longer than opportunistic interactions would predict. This claim is supported by recent research that revealed clear fitness benefits for long-lasting social bonds. Here, we investigated the stability and equitability of social relationships in captive rhesus macaques, measured using their proximity and grooming patterns over a 3-yr period. We show that, in general, proximity patterns of the whole group were stable over 3 yr. More specifically, each individual’s top two nonkin within and between sex relationships based on proximity were relatively stable and some of these relationships lasted the full 3 yr. Female grooming relationships within and between sex were also stable over 3 yr. Moreover, strong social bonds in females were not based on age similarity and were not sex specific, suggesting that a general underlying mechanism of social bonding in female rhesus macaques. In contrast, patterns of male grooming of other males and of females were relatively unstable and male social bonds were less equitable than those of females. The differences in stability of the relationships between the sexes may reflect the dispersal pattern of rhesus macaques, although we have limited data on non-natal males in our study. With these data we add to a growing body of literature describing the value and durability of relationships among primates and other species.

Auditory Contagious Yawning in Humans: An Investigation into Affiliation and Status Effects.

While comparative research on contagious yawning has grown substantially in the past few years, both the interpersonal factors influencing this response and the sensory modalities involved in its activation in humans remain relatively unknown. Extending upon previous studies showing various in-group and status effects in non-human great apes, we performed an initial study to investigate how the political affiliation (Democrat vs. Republican) and status (high vs. low) of target stimuli influences auditory contagious yawning, as well as the urge to yawn, in humans. Self-report responses and a subset of video recordings were analyzed from 118 undergraduate students in the US following exposure to either breathing (control) or yawning (experimental) vocalizations paired with images of former US Presidents (high status) and their respective Cabinet Secretaries of Commerce (low status). The overall results validate the use of auditory stimuli to prompt yawn contagion, with greater response in the experimental than the control condition. There was also a negative effect of political status on self-reported yawning and the self-reported urge to yawn irrespective of the condition. In contrast, we found no evidence for a political affiliation bias in this response. These preliminary findings are discussed in terms of the existing comparative evidence, though we highlight limitations in the current investigation and we provide suggestions for future research in this area.

No costly prosociality among related long-tailed macaques (Macaca fascicularis).

Altruism, benefiting another at a cost to the donor, may be achieved through prosocial behavior. Studies of nonhuman animals typically investigate prosocial behavior with paradigms in which the donor can choose to give a recipient a food item, and the choice does not affect the donors reward (which is either present or absent). In such tasks, long-tailed macaques (Macaca fascicularis) show prosocial behavior, especially toward kin. Here, we tested captive long-tailed macaques with related recipients in an alternative task, in which the donor had to give up a preferred reward to benefit the recipient; that is, they had to choose a lower valued reward for themselves to provide food to their kin. Overall, the macaques did not provide their kin with food. The task forced the donor to balance its prosocial behavior with its selfish choice for a higher value reward, a balance that turned out to favor selfish motives. Consequently, our study shows that a prosocial tendency is not sufficient to elicit costly prosocial behavior in long-tailed macaques. Subsequently, we feel that tasks in which the donor must choose a lower value reward to benefit another individual may allow the titration of the strength of prosocial behavior, and thus provides interesting possibilities for future comparative studies.