Stability and Change of Social Relationship Quality in Captive Chimpanzees (Pan troglodytes)

In social animals an individual’s fitness depends partly on the quality of relationships with others. Qualitative variation in relationships has been conceptualized according to a three-dimensional structure, consisting of relationship value, compatibility, and security. However, the determinants of the components and their temporal stability are not well understood. We studied relationship quality in a newly formed group of 20 captive chimpanzees made up of several previously existing social groups. We assessed dyadic relationship quality 2?yr and again 7?yr after grouping. We confirmed the existence and stability of three relationship components and labeled them value, compatibility, and approach symmetry. Previously familiar dyads had a higher value than unfamiliar dyads, especially when they were maternally or paternally related. Compatibility was higher in dyads with only females than in dyads containing a male, but familiarity did not influence compatibility. Approach symmetry was initially higher, but later lower, in familiar than unfamiliar dyads, indicating that approach symmetry of familiar dyads decreased over time. Dyadic value and compatibility were highly stable over time, which is similar to the long relationship duration found in wild chimpanzees. In sum, relationships formed earlier in life became more valuable than those formed in later adulthood, whereas nonaggressive, compatible relationships could be formed throughout life. This suggests that for immigrating individuals, high-value relationships may be relatively difficult to establish, partly explaining why wild female chimpanzees have relatively few high-quality relationships with other females. Our study supports the multicomponent structure and durability of relationships in social species.     

Problem Solving in the Presence of Others: How Rank and Relationship Quality Impact Resource Acquisition in Chimpanzees (Pan troglodytes)

In the wild, chimpanzees (Pan troglodytes) are often faced with clumped food resources that they may know how to access but abstain from doing so due to social pressures. To better understand how social settings influence resource acquisition, we tested fifteen semi-wild chimpanzees from two social groups alone and in the presence of others. We investigated how resource acquisition was affected by relative social dominance, whether collaborative problem solving or (active or passive) sharing occurred amongst any of the dyads, and whether these outcomes were related to relationship quality as determined from six months of observational data. Results indicated that chimpanzees obtained fewer rewards when tested in the presence of others compared to when they were tested alone, and this loss tended to be greater when paired with a higher ranked individual. Individuals demonstrated behavioral inhibition; chimpanzees who showed proficient skill when alone often abstained from solving the task when in the presence of others. Finally, individuals with close social relationships spent more time together in the problem solving space, but collaboration and sharing were infrequent and sessions in which collaboration or sharing did occur contained more instances of aggression. Group living provides benefits and imposes costs, and these findings highlight that one cost of group living may be diminishing productive individual behaviors.     

The Context and Quality of Social Relationships Affect Vigilance Behaviour in Wild Chimpanzees

Previous studies of vigilance behaviour have focused mainly on the influence of predation threat, whereas the influences of conspecific factors, such as within‐group threats, are relatively unstudied. To elucidate the influences of conspecific factors, this study examined vigilance behaviour in wild chimpanzees (Pan troglodytes schweinfurthii) in Mahale Mountains National Park, Tanzania. Vigilance level was lower during foraging than during resting, which indicated a conflict between vigilance and foraging activity. In addition, vigilance level was higher when chimpanzees were on the ground where an encounter with leopards (Panthera pardus) is likely than when the chimpanzees were in trees. Males, but not females, increased their level of vigilance as the number of individuals within 3 m increased. In both males and females, daily party size – an index of group cohesion – did not affect the vigilance level. The level of maternal vigilance was higher when a dependent infant was separated from its mother than when the offspring was in contact with its mother. Both males and females increased their vigilance when a less‐associated group member was nearby, when compared with when there was no less‐associated group member nearby. This finding suggests that variation in relationship quality influences the vigilance level and that individuals need to increase their level of vigilance when the level of within‐group threats is high. This study indicated that variation in vigilance cannot be understood unless conspecific factors, such as variation in the relationship quality with associates, are considered.     

Chimpanzees Today

Chimpanzees Today. 2001. 36 minutes, color. video by Anne Zeller. For more information, contact Insight Media, 2162 Broadway, New York, NY 10024-0621.     

Experimental Toxoplasmosis in Chimpanzees

Two chimpanzees were given by mouth large numbers of viable oocysts of Toxoplasma gondii obtained from the faeces of experimentally infected cats. Before the experiment the first chimpanzee had a positive dye test reaction (1:250), an indication that it had undergone an earlier infection of toxoplasmosis; the serum antibody titres remained unchanged, no evidence of illness was found, and oocysts did not appear in its faeces during the subsequent six weeks. The second chimpanzee showed a negative dye test reaction before infection, and this converted to positive on the 7th day, rose to a peak on the 35th day, and remained high for six months. This animal appeared unwell during the first week, and on the 7th day its blood proved infective to mice; on the 40th day the lymph nodes became enlarged and biopsy specimens of a node and muscle in the 11th week were also infective to mice. No oocysts were passed in the faeces. The presumed cycle in the chimpanzee and in man and the relationships between Toxoplasma and Isospora are discussed.     

Chimpanzees and retirement

The subject of sanctuaries for chimpanzees has lately become the topic of a great deal of discussion (Brent, Butler, & Haberstroh, 1997; Committee on Long-Term Care of Chimpanzees, 1997; Dyke, Williams-Blangero, Mamelka, & Goodwin, 1995; Peterson & Goodall, 1993). In the United States, laboratories that use chimpanzees in research are facing a housing crisis. An increase in captive births caused by the initiation of the National Chimpanzee Breeding and Research Program in 1986 (Hobson, Graham, & Rowell, 1991), coupled with the diminished use of chimpanzees as experimental subjects, have led to a large population of chimpanzees considered to be surplus to demand (Blood, Wolfle, & Whitney, 1992). These chimpanzees, as well as an unknown number from the private sector, are candidates for what is currently being called retirement.     

Ontogeny of Ranging in Wild Chimpanzees

We examined the relationship between juvenile age and distance traveled per day, or day range, in Kanyawara chimpanzees. Because the energy cost of locomotion is greater for small-bodied animals, we predict that day range is constrained by body size, i.e., younger individuals tend to have shorter day ranges. To test this hypothesis, we measured day range for 200 day-ranges of groups in which we recorded the age of the youngest juvenile present. As predicted, day range correlated positively with age for juveniles. Comparisons of day range vs. estimated stature support the hypothesis that the increase in day range with age was a consequence of body size. To assess other sources of variation in day range, we also measured the effects of group size and the presence of a carried infant. While day range correlated significantly with group size, the presence of a carried infant had no effect on adult female day range. Our results suggest the size of a juvenile may constrain ranging for mothers and their offspring.     

The Relationship between Event-Based Prospective Memory and Ongoing Task Performance in Chimpanzees (Pan troglodytes)

Prospective memory is remembering to do something at a future time. A growing body of research supports that prospective memory may exist in nonhuman animals, but the methods used to test nonhuman prospective memory differ from those used with humans. The current work tests prospective memory in chimpanzees using a method that closely approximates a typical human paradigm. In these experiments, the prospective memory cue was embedded within an ongoing task. Tokens representing food items could be used in one of two ways: in a matching task with pictures of items (the ongoing task) or to request a food item hidden in a different location at the beginning of the trial. Chimpanzees had to disengage from the ongoing task in order to use the appropriate token to obtain a higher preference food item. In Experiment 1, chimpanzees effectively matched tokens to pictures, when appropriate, and disengaged from the ongoing task when the token matched the hidden item. In Experiment 2, performance did not differ when the target item was either hidden or visible. This suggested no effect of cognitive load on either the prospective memory task or the ongoing task, but performance was near ceiling, which may have contributed to this outcome. In Experiment 3, we created a more challenging version of the task. More errors on the matching task occurred before the prospective memory had been carried out, and this difference seemed to be limited to the hidden condition. This finding parallels results from human studies and suggests that working memory load and prospective memory may have a similar relationship in nonhuman primates.     

Chimpanzees coordinate in a negotiation game

A crucially important aspect of human cooperation is the ability to negotiate to cooperative outcomes when interests over resources conflict. Although chimpanzees and other social species may negotiate conflicting interests regarding travel direction or activity timing, very little is known about their ability to negotiate conflicting preferences over food. In the current study, we presented pairs of chimpanzees with a choice between two cooperative tasks—one with equal payoffs (e.g., 5-5) and one with unequal payoffs (higher and lower than in the equal option, e.g., 10-1). This created a conflict of interests between partners with failure to work together on the same cooperative task resulting in no payoff for either partner. The chimpanzee pairs cooperated successfully in as many as 78–94% of the trials across experiments. Even though dominant chimpanzees preferred the unequal option (as they would obtain the largest payoff), subordinate chimpanzees were able to get their way (the equal option) in 22–56% of trials across conditions. Various analyses showed that subjects were both strategic and also cognizant of the strategies used by their partners. These results demonstrate that one of our two closest primate relatives, the chimpanzee, can settle conflicts of interest over resources in mutually satisfying ways—even without the social norms of equity, planned strategies of reciprocity, and the complex communication characteristic of human negotiation.     

Subsistence Technology of Nigerian Chimpanzees

A trademark of Homo sapiens is the enormous variation in behavioral patterns across populations. Insight into the development of human cultures can be aided by studies on communities of Pan across Africa that display unique combinations of social behavior and elementary technology. Only cross-population comparisons can reveal whether the diversity reflects differential genetics, environmental constraints, or is a cultural variant. However, the recently recognized and most endangered subspecies, Pan troglodytes vellerosus, remains completely unstudied in this respect. We report first evidence from a new long-term study of Nigerian chimpanzees at Gashaka. Their dietary composition is highly varied and they have to cope with high concentrations of antifeedant defenses of plants against consumption. Gashaka chimpanzees use a varied tool kit for extractive foraging. For example, they harvest insects throughout the year, via digging sticks and probes, to obtain honey from stingless-bee and honeybee nests, dipping wands to prey on army ants, and fishing rods to eat arboreal ants. Tools appeared to be custom-made with a considerable degree of standardization in length, diameter, and preferential use of distal ends. Moreover, compared to the rainy season, tools were longer during the dry season when insects retreat further into their nests. Many of the expressions of subsistence technology seem to be environmentally constrained. Most notably, the absence of termite-eating could reflect a low abundance of mounds. Other traits may represent cultural variation. For example, the chimpanzees did not hammer open 2 types of hard-shelled nuts with tools, unlike what occurs elsewhere in West Africa. The prevalence of elementary technology may indicate that the material culture of Gashaka chimpanzees is most related to core cultural tendencies of Central African populations.     

Chimpanzees share forbidden fruit

The sharing of wild plant foods is infrequent in chimpanzees, but in chimpanzee communities that engage in hunting, meat is frequently used as a 'social tool' for nurturing alliances and social bonds. Here we report the only recorded example of regular sharing of plant foods by unrelated, non-provisioned wild chimpanzees, and the contexts in which these sharing behaviours occur. From direct observations, adult chimpanzees at Bossou (Republic of Guinea, West Africa) very rarely transferred wild plant foods. In contrast, they shared cultivated plant foods much more frequently (58 out of 59 food sharing events). Sharing primarily consists of adult males allowing reproductively cycling females to take food that they possess. We propose that hypotheses focussing on 'food-for-sex and -grooming' and 'showing-off' strategies plausibly account for observed sharing behaviours. A changing human-dominated landscape presents chimpanzees with fresh challenges, and our observations suggest that crop-raiding provides adult male chimpanzees at Bossou with highly desirable food commodities that may be traded for other currencies.     

Chimpanzees and the mathematics of battle

Recent experiments have demonstrated the importance of numerical assessment in animal contests. Nevertheless, few attempts have been made to model explicitly the relationship between the relative number of combatants on each side and the costs and benefits of entering a contest. One framework that may be especially suitable for making such explicit predictions is Lanchester's theory of combat, which has proved useful for understanding combat strategies in humans and several species of ants. We show, with data from a recent series of playback experiments, that a model derived from Lanchester's 'square law' predicts willingness to enter intergroup contests in wild chimpanzees (Pan troglodytes). Furthermore, the model predicts that, in contests with multiple individuals on each side, chimpanzees in this population should be willing to enter a contest only if they outnumber the opposing side by a factor of 1.5. We evaluate these results for intergroup encounters in chimpanzees and also discuss potential applications of Lanchester's square and linear laws for understanding combat strategies in other species.     

Development of the Laryngeal Air Sac in Chimpanzees

Though many nonhuman primates possess a laryngeal sac, the great apes are unique in their great size. Though an enlarged sac probably arose in their common ancestor, its functional adaptations remain a matter of debate. Its development in extant great apes is likely to provide valuable information to clarify the issue. We used magnetic resonance imaging to examine the development of the laryngeal sac in 3 living chimpanzees, age 4 mo–5 yr, and identified 2 distinct growth phases of the sac. A gradual growth of the sac in early infancy results in a configuration so that it occupies the ventral region of the neck; many adult nonhominoid primates having a sac show the configuration. The subsequent rapid expansion of the sac in late infancy causes the final configuration in chimpanzees, wherein the sac expands into the pectoral, clavicular, and axillary regions. The latter phase possibly arose at latest in the last common ancestor of extant great apes and contributed to the evolution of the enlarged sac, despite the later evolutionary diversification in adult sac anatomy and growth. As many studies have advocated, the enlarged sac probably plays a role in vocalization in adults. However, physiological modifications in the laryngeal region during infancy are likely to provide valuable information to evaluate the functional adaptations of the enlarged sac in the great apes.     

Noninvasive Monitoring of Respiratory Viruses in Wild Chimpanzees

To diagnose respiratory disease among wild great apes, there is a need for noninvasive diagnostic methods. Therefore, we analyzed fecal samples from habituated chimpanzees from Taï National Park, Côte d’Ivoire. Samples had been collected during four distinct outbreaks: two with known aetiology (March 2004 and February 2006) and two with unknown aetiology (October 2004 and August 2005). Fecal samples were screened by polymerase chain reaction (PCR) for the presence of human metapneumovirus (HMPV) and human respiratory syncytial virus (HRSV), two paramyxoviruses previously found in lung tissue of chimpanzees that died due to respiratory disease. In the March 2004 outbreak, 72% of the tested individuals were positive for HMPV, and during the 2006 epidemic, 25% tested HRSV-positive. In the outbreaks where no causative pathogen was previously known, fecal samples tested positive for either HRSV or HMPV, showing that reinfection occurred. Virus sequences were generated and compared with sequences previously found in tissue; nearly identical virus sequences in both tissue and fecal samples were found. These results demonstrate that fecal samples collected during outbreak times can be used for the diagnostic and phylogenetic analysis of HMPV and HRSV. Using such diagnostic tools, systematic noninvasive disease investigation of respiratory outbreaks in wild great apes becomes possible. The methods presented here may also be applied for the investigation of further acute diseases in great apes and other species.