Matthew W. Campbell

Ingroup-Outgroup Bias in Contagious Yawning by Chimpanzees Supports Link to Empathy

Humans favor others seen as similar to themselves (ingroup) over people seen as different (outgroup), even without explicitly stated bias. Ingroup-outgroup bias extends to involuntary responses, such as empathy for pain. However, empathy biases have not been tested in our close primate relatives. Contagious yawning has been theoretically and empirically linked to empathy. If empathy underlies contagious yawning, we predict that subjects should show an ingroup-outgroup bias by yawning more in response to watching ingroup members yawn than outgroup. Twenty-three chimpanzees (Pan troglodytes) from two separate groups watched videos of familiar and unfamiliar individuals yawning or at rest (control). The chimpanzees yawned more when watching the familiar yawns than the familiar control or the unfamiliar yawns, demonstrating an ingroup-outgroup bias in contagious yawning. These results provide further empirical support that contagious yawning is a measure of empathy, which may be useful for evolutionary biology and mental health.

Computer animations stimulate contagious yawning in chimpanzees

People empathize with fictional displays of behaviour, including those of cartoons and computer animations, even though the stimuli are obviously artificial. However, the extent to which other animals also may respond empathetically to animations has yet to be determined. Animations provide a potentially useful tool for exploring non-human behaviour, cognition and empathy because computer-generated stimuli offer complete control over variables and the ability to program stimuli that could not be captured on video. Establishing computer animations as a viable tool requires that non-human subjects identify with and respond to animations in a way similar to the way they do to images of actual conspecifics. Contagious yawning has been linked to empathy and poses a good test of involuntary identification and motor mimicry. We presented 24 chimpanzees with three-dimensional computer-animated chimpanzees yawning or displaying control mouth movements. The apes yawned significantly more in response to the yawn animations than to the controls, implying identification with the animations. These results support the phenomenon of contagious yawning in chimpanzees and suggest an empathic response to animations. Understanding how chimpanzees connect with animations, to both empathize and imitate, may help us to understand how humans do the same.

Ape duos and trios: spontaneous cooperation with free partner choice in chimpanzees

The purpose of the present study was to push the boundaries of cooperation among captive chimpanzees (Pan troglodytes). There has been doubt about the level of cooperation that chimpanzees are able to spontaneously achieve or understand. Would they, without any pre-training or restrictions in partner choice, be able to develop successful joint action? And would they be able to extend cooperation to more than two partners, as they do in nature? Chimpanzees were given a chance to cooperate with multiple partners of their own choosing. All members of the group (N = 11) had simultaneous access to an apparatus that required two (dyadic condition) or three (triadic condition) individuals to pull in a tray baited with food. Without any training, the chimpanzees spontaneously solved the task a total of 3,565 times in both dyadic and triadic combinations. Their success rate and efficiency increased over time, whereas the amount of pulling in the absence of a partner decreased, demonstrating that they had learned the task contingencies. They preferentially approached the apparatus when kin or nonkin of similar rank were present, showing a preference for socially tolerant partners. The forced partner combinations typical of cooperation experiments cannot reveal these abilities, which demonstrate that in the midst of a complex social environment, chimpanzees spontaneously initiate and maintain a high level of cooperative behavior.

Chimpanzees empathize with group mates and humans, but not with baboons or unfamiliar chimpanzees

Human empathy can extend to strangers and even other species, but it is unknown whether non-humans are similarly broad in their empathic responses. We explored the breadth and flexibility of empathy in chimpanzees, a close relative of humans. We used contagious yawning to measure involuntary empathy and showed chimpanzees videos of familiar humans, unfamiliar humans and gelada baboons (an unfamiliar species). We tested whether each class of stimuli elicited contagion by comparing the effect of yawn and control videos. After including previous data on the response to ingroup and outgroup chimpanzees, we found that familiar and unfamiliar humans elicited contagion equal to that of ingroup chimpanzees. Gelada baboons did not elicit contagion, and the response to them was equal to that of outgroup chimpanzees. However, the chimpanzees watched the outgroup chimpanzee videos more than any other. The combination of high interest and low contagion may stem from hostility towards unfamiliar chimpanzees, which may interfere with an empathic response. Overall, chimpanzees showed flexibility in that they formed an empathic connection with a different species, including unknown members of that species. These results imply that human empathic flexibility is shared with related species.

Vocal Response of Captive-reared Saguinus oedipus During Mobbing

Mobbing is an important component of antipredator behavior for animals from many taxa. Callitrichids are small-bodied primates that mob multiple types of predators. Though there have been several observations of callitrichids mobbing predators in the wild, their anecdotal nature provides only rough descriptions of behavior and vocalizations. Researchers could neither spectrographically identify nor quantify vocalizations owing to the limitations of observing predation in the field. We examined in detail the mobbing response of 1 callitrichid species, the cotton-top tamarin, in captivity. We recorded vocalizations for quantitative analysis and observed behavior qualitatively. We report 3 new vocalizations that had not been described in the original repertoire for the cotton-top tamarin. Analysis of the time course of a mobbing session yielded a pattern in which the highest intensity mobbing vocalizations decreased over the session even though lower intensity vocalizing continued, which may reflect a shifting strategy from mobbing to vigilance. The rate of calling during mobbing sessions differed from the rate of calling during control sessions. We discuss the vocalizations in relation to 2 hypotheses of form and function of antipredator calls. The newly described mobbing vocalizations may have an important impact on the study of mobbing because they represent a class of vocalizations that researchers have largely ignored in studies of callitrichids, thus raising new issues concerning past and future research on antipredator behavior in the family.

How chimpanzees cooperate in a competitive world

Significance Competitive tendencies may make it hard for members of a group to cooperate with each other. Humans use many different “enforcement” strategies to keep competition in check and favor cooperation. To test whether one of our closest relatives uses similar strategies, we gave a group of chimpanzees a cooperative problem that required joint action by two or three individuals. The open-group set-up allowed the chimpanzees a choice between cooperation and competitive behavior like freeloading. The chimpanzees used a combination of partner choice and punishment of competitive individuals to reduce competition. In the end, cooperation won. Our results suggest that the roots of human cooperation are shared with other primates. Our species is routinely depicted as unique in its ability to achieve cooperation, whereas our closest relative, the chimpanzee (Pan troglodytes), is often characterized as overly competitive. Human cooperation is assisted by the cost attached to competitive tendencies through enforcement mechanisms, such as punishment and partner choice. To examine if chimpanzees possess the same ability to mitigate competition, we set up a cooperative task in the presence of the entire group of 11 adults, which required two or three individuals to pull jointly to receive rewards. This open-group set-up provided ample opportunity for competition (e.g., freeloading, displacements) and aggression. Despite this unique set-up and initial competitiveness, cooperation prevailed in the end, being at least five times as common as competition. The chimpanzees performed 3,565 cooperative acts while using a variety of enforcement mechanisms to overcome competition and freeloading, as measured by (attempted) thefts of rewards. These mechanisms included direct protest by the target, third-party punishment in which dominant individuals intervened against freeloaders, and partner choice. There was a marked difference between freeloading and displacement; freeloading tended to elicit withdrawal and third-party interventions, whereas displacements were met with a higher rate of direct retaliation. Humans have shown similar responses in controlled experiments, suggesting shared mechanisms across the primates to mitigate competition for the sake of cooperation.

Methodological Problems in the Study of Contagious Yawning

The recent interest in contagious yawning has raised several challenges as the varied methods of testing have left some unresolved issues. We do not know how differences in key variables affect the observed rates of yawning, and we highlight these as being in need of direct testing. Different researchers analyze their results differently, and we make some recommendations for more rigorous, thorough and informative analyses. Ultimately, problems arise when authors compare studies that used different methods and different analyses without acknowledging how these differences may have affected the results. In these cases, authors make inappropriate comparisons, which lead to conclusions that add confusion to the literature. Our goal in raising awareness of these issues is to generate new experiments and improve the discussion of existing research. With its link to empathy, a more standardized study of contagious yawning may be a useful tool for a variety of disciplines.

Captive‐born cotton‐top tamarins (Saguinus oedipus) respond similarly to vocalizations of predators and sympatric nonpredators

What types of cues do callitrichid primates use to detect and respond to predators? Do they respond to predator‐specific cues or to more general cues? The evidence for these questions remains conflicting. We presented captive‐born and reared cotton‐top tamarins with no previous exposure to predators (or predator cues) with vocalizations from three potential predators of cotton‐top tamarin in the wild (white hawk, jaguar, and tayra) and with vocalizations from sympatric nonpredators (black‐faced antthrush and red howler monkey). Vocalizations from predators and from nonpredator mammals elicited equivalent arousal, fear, and vocal responses. Howler monkey roars produced the strongest responses. The results suggest that predator‐naïve cotton‐top tamarins do not recognize specific predator vocalizations, but may respond to vocal qualities (low‐frequency, noisy sounds) that indicate large body size, threat, or aggression. On the other hand, tamarins responded much more strongly to the higher frequency calls from the hawk than the antthrush, suggesting another mechanism must also be involved. The failure of captive‐reared tamarins to distinguish between vocalizations of predators and nonpredator mammals has important implications for reintroduction studies. Am. J. Primatol. 70:707–710, 2008.

The roles of food quality and sex in chimpanzee sharing behavior (Pan troglodytes)

Both wild and captive chimpanzees (Pan troglodytes) share food with non-relatives. Researchers have proposed several hypotheses to explain this behavior, including ‘food for sex’, ‘food for grooming or agonistic support’, and ‘sharing under pressure’. We examined food sharing in two captive groups of socially-housed chimpanzees. In contrast to previous captive studies, which only examined transfers of low-quality foods, we conducted seven trials with high-quality food and seven with low-quality food for each group to directly compare transfers of different food qualities. We recorded how male chimpanzees shared food, including active transfers, passive transfers, and co-feeding. We also noted all instances of copulations, female estrous states, benign attempts to access food (termed ‘perseverance’), and aggressive attempts (termed ‘harassment’) to examine whether any of these factors influenced food sharing. Male food possessors shared at the same rate in both food quality conditions, but seemingly for different reasons, indicating that food quality may affect the exchange of social benefits in chimpanzees. In the low-quality condition, there was an interaction with rank and perseverance: while low- and middle-ranking females received more food the more they persevered, high-ranking females received more food without perseverance and gained relatively little benefit from persevering. In the high-quality condition, there was an interaction between copulations and perseverance: females who copulated with the male food possessor received more food during that trial with less perseverance. Non-copulating females received more transfers the more they persevered. This result was only observed in the shortterm — copulations over the previous year were not correlated with food transfers. Further, the copulations observed here were unusual for these chimpanzees in that they were not confined to peak fertility, suggesting a non-conceptive function for copulations in chimpanzees. Copulations in this study may have functioned to reduce tension and increase short-term tolerance, allowing females better access to food.

Chimpanzees Prefer African and Indian Music Over Silence

[Correction Notice: An Erratum for this article was reported in Vol 40(4) of Journal of Experimental Psychology: Animal Learning and Cognition (see record 2014-35305-001). For the article, the below files were used to create the audio used in this study. The original West African akan and North Indian raga pieces were used in their entirety and the Japanese taiko piece was used from the 0:19 second mark through the end. The tempo of each piece was adjusted so that they maintained an identical base tempo of 90 beats per minute, then looped to create 40 minutes of continuous music. Additionally, the volume of the music was standardized at 50 dB so that the all music maintained the same average amplitude. All audio manipulations were completed using GarageBand