Female genital swellings have no effect on chimpanzee agonism in well-established groups

Reducing the incidence of aggression and wounding is a major concern for those managing socially housed chimpanzees (Pan troglodytes). Competition among adult males for access to sexually receptive females is believed to be one factor contributing to undesirably high levels of agonism. An observational analysis of two chimpanzee groups (n = 9, n = 8) was initiated to study this phenomenon. All-events sampling was used to record agonistic and sexual behavior on days when one female in the group showed a maximal genital swelling and on other days when no female in the group had a tumescent swelling. The result of the multivariate analysis of variance indicated that the presence of a female with a tumescent swelling had no effect on bluff displays or aggressive, submissive, or reconciliatory behavior but that sexual behavior was significantly increased. These results may be explained by the fact that long-term stability in group membership may reduce male–male competition for sexual access to females and the sometimes associated aggression and wounding. Colony managers dealing with well-established groups of chimpanzees should invest more of their effort in controlling aggression elicited by factors other than the cycling of a single female group member.     

Injury risks among chimpanzees in three housing conditions

Meeting the psychological needs of chimpanzees (Pan troglodytes) can be a challenge given their aggressiveness on the one hand and the complexity of their social lives on the other. It is unclear how to balance the need to provide opportunities for species-appropriate behavior against potential risks of injury chimpanzees may inflict on each other. This study evaluates the suggestion that simpler social environments protect chimpanzees from wounding. Over a two-year period all visible injuries to 46 adult males, 64 adult females, and 25 immature chimpanzees were recorded at the Yerkes Regional Primate Research Center. Approximately half of the subjects were mother-reared, and the rest were nursery-reared. Housing included compounds containing about 20 chimpanzees, interconnected indoor-outdoor runs for groups of up to 12 individuals, and smaller indoor-outdoor runs for pairs and trios. Annual wounding rates were calculated for serious wounds (extensive injuries and all those requiring veterinary intervention) as well as for minor wounds. Compound-housed chimpanzees incurred the highest level of minor wounding, but serious wounding levels were not affected by housing condition. Even with a period of dominance instability and elevated levels of wounding in one compound, compound chimpanzees were not injured more than those in smaller social groups over the long term. Nursery-reared females in moderate-sized groups were wounded more than mother-reared females. Also, nursery-reared males and females were wounded less often when paired with mother-reared companions. Overall, this study indicates that maintaining chimpanzees in pairs and trios would not be an effective means for reducing injuries. The management of wounding in chimpanzee colonies is influenced more by the sex and rearing composition of a colony.     

Factors affecting wounding aggression in a colony of captive chimpanzees (Pan troglodytes)

Previous research has suggested that chimpanzees (Pan troglodytes) display higher levels of aggression in captivity than in the wild. One of the challenges of captive management, therefore, is to balance the chimpanzees' need for social interaction with managements' desire to minimize wounding and aggression. Various captive studies have examined the effects of individual and social variables on the frequency of wounding aggression, but none have examined these variables simultaneously. We collected retrospective wounding data for severe wounds from 83 captive chimpanzees (36 males, 47 females) from January 1993 to December 2003. The context of the wounding event, including individual age and sex, group age and sex composition, group duration, and portion of the week (weekday vs. weekend) were collected. Logistic regression analysis was performed to determine which variables had a significant effect on the probability of a severe wounding event. The sex and age composition of the group, group duration, and portion of the week had a statistically significant association with wounding. All‐male groups (Odds Ratio (OR)=6.738) had the highest risk of wounding aggression, with uni‐male groups (OR=3.311) having the next largest. Compared to individuals in all sub‐adult groups, individuals in either all‐adult (OR=4.516) or mixed‐age (OR=3.587) groups had a higher risk of wounding. There was an inverse association between group duration and wounding (OR=0.821). Finally, there was an increased risk of wounding during the work week (OR=1.653). These results suggest that captive management should pay close attention to group composition, as well as levels of human activity, when devising strategies to reduce captive chimpanzee aggression. Zoo Biol 29:351–364, 2010. © 2009 Wiley‐Liss, Inc.     

Chimpanzee (Pan troglodytes) birth patterns and human presence in zoological settings

In response to work demonstrating a negative correlation between human staff activity and parturition in laboratory-managed primates, this study examined the distribution of 231 captive chimpanzee (Pan troglodytes) births that occurred in accredited American zoological institutions by the day of the week on which the birth was discovered. We hypothesized that if chimpanzee parturition patterns displayed sensitivity to human presence, then fewer births per day would be reported during the weekend period, when the visitor density was high, as compared with the lower density working week. Analyses indicated that chimpanzee births were randomly distributed throughout the week. In the context of the questionable sensitivity of primate parturition to external influence, results suggest that variations in human presence do not affect the fine-level timing of birth in chimpanzees managed in a zoological setting.     

The quality of chimpanzee T-cell activation and simian immunodeficiency virus/human immunodeficiency virus susceptibility achieved via antibody-mediated T-cell receptor/CD3 stimulation is a function of the anti-CD3 antibody isotype

While human immunodeficiency virus type 1 (HIV-1) infection is associated with hyperimmune activation and systemic depletion of CD4⁺ T cells, simian immunodeficiency virus (SIV) infection in sooty mangabeys or chimpanzees does not exhibit these hallmarks. Control of immune activation is thought to be one of the major components that govern species-dependent differences in the disease pathogenesis. A previous study introduced the idea that the resistance of chimpanzees to SIVcpz infection-induced hyperimmune activation could be the result of the expression of select sialic acid-recognizing immunoglobulin (Ig)-like lectin (Siglec) superfamily members by chimpanzee T cells. Siglecs, which are absent on human T cells, were thought to control levels of T-cell activation in chimpanzees and were thus suggested as a cause for the pathogenic differences in the course of SIVcpz or HIV-1 infection. As in human models of T-cell activation, stimulation had been attempted using an anti-CD3 monoclonal antibody (MAb) (UCHT1; isotype IgG1), but despite efficient binding, UCHT1 failed to activate chimpanzee T cells, an activation block that could be partially overcome by MAb-induced Siglec-5 internalization. We herein demonstrate that anti-CD3 MAb-mediated chimpanzee T-cell activation is a function of the anti-CD3 MAb isotype and is not governed by Siglec expression. While IgG1 anti-CD3 MAbs fail to stimulate chimpanzee T cells, IgG2a anti-CD3 MAbs activate chimpanzee T cells in the absence of Siglec manipulations. Our results thus imply that prior to studying possible differences between human and chimpanzee T-cell activation, a relevant model of chimpanzee T cell activation needs to be established.     

Evaluation of introduction procedures in captive chimpanzees

Chimpanzee introductions can be stressful and even dangerous for the individuals involved, and different facilities use different approaches to group formation. However, little empirical evidence has been available to evaluate the different factors related to successful or unsuccessful chimpanzee introductions. This study evaluated a variety of factors, such as age, sex, previous experience, and the behavior displayed during the initial introduction period, to determine any influence on the outcome of the introduction in 165 dyadic introduction events. The direct introduction of one chimpanzee to a group or to another individual resulted in an overall success rate of 86.7%. The first few social interactions were characterized by more aggressive and submissive behavior than afffiliative behavior, but this trend was quickly reversed. Unsuccessful outcomes were more common if wounding occurred, if either individual was male, and if the newcomer had high levels of submissive behavior. Among male chimpanzees, the behaviors during the first 10 min of the introductions were characterized by more submissive behavior of the newcomer and more aggressive behavior of the introduction partner. The frequency of aggression by the introduction partner was higher when the newcomer was introduced to one chimpanzee at a time rather than to the group as a whole. This information may have important consequences for how we view chimpanzee group formation, and factors such as previous experience and aggression may be less important than the method of introduction and the level of submissive behavior. Zoo Biol 16:335–342, 1997. © 1997 Wiley-Liss, Inc.     

The Long‐term Impact of Timber Harvesting on the Resource Base of Chimpanzees in Kibale National Park,Uganda

Commercial timber harvesting results in the loss of critical habitat for tropical forest fauna, and large‐bodied frugivores (including chimpanzees and most other apes) may experience particularly detrimental effects. Few quantitative data, however, are available to evaluate the long‐term impact of harvesting on chimpanzees and other apes. In particular, few data are available to compare population demographics and/or forest composition before and after timber harvesting at the same site. Utilizing detailed forestry department records of logging operations conducted in the late 1960s, present‐day botanical surveys, and long‐term data on the feeding ecology of chimpanzees in Kibale National Park (KNP), Uganda, I examined the impact that logging has had on KNP chimpanzee communities of known size and demography. Although some important chimpanzee food resources were harvested in high abundance during commercial logging operations, the overall impact on the most predominant dietary items (those making up roughly 75% of the chimpanzees' diet) and on presumably critical subsistence resources was limited. Furthermore, the low density of chimpanzees inhabiting the logged region of KNP is apparently not attributable to the impact of logging at the site: comparisons of resource densities at this ‘low‐chimpanzee‐density’ site with that of an unlogged and ‘high‐chimpanzee‐density’ KNP site did not differ when logging concessions at the low‐chimpanzee‐density site were excluded from the analysis. This study suggests that low‐intensity logging can be compatible with the conservation of large‐bodied frugivores, provided that dietary data are taken into account in forest management planning.     

Positional behavior of Pan troglodytes in the Mahale Mountains and Gombe Stream National Parks, Tanzania.

The positional behavior of habituated adult chimpanzees and baboons was observed for 784 hr in a year-long study. Comparisons between species were made to establish the distinctiveness of chimpanzee positional behavior and habitat use. Brachiation (sensu stricto, i.e., hand-over-hand suspensory locomotion) was observed in low frequencies among chimpanzees, and its significance for chimpanzee anatomy is judged slight. Although no significant differences were found between sympatric baboons and chimpanzees in the proportion of time spent in the terminal branches, or in the mean diameter of weight-bearing strata, chimpanzees exhibited evidence of a terminal branch adaptation in that they, unlike baboons, used postures among smaller supporting strata different from those used among larger supports. Among chimpanzees, unimanual arm-hanging was most common among the smallest strata and was associated with smaller mean and median support diameter than other postures. Unimanual arm-hanging was the only common behavior among chimpanzees that usually involved complete abduction of the humerus. A number of behaviors often subsumed under the label "quadrumanous climbing" were distinguished in this study. Compared to baboons and other cercopithecoids, chimpanzees did not show increased frequencies of large-stratum vertical climbing, and their vertical climbing did not involve significant humeral abduction. Arm-hanging (i.e., unimanual suspension) and vertical climbing distinguish chimpanzee positional behavior from that of monkeys.     

Wounding aggression during the formation and maintenance of captive,multimale chimpanzee groups

Although the multimale community is the natural social organization of chimpanzees, both wild and captive adult males have killed other adult males and infants in intercommunity conflicts and intragroup aggression. Despite the potential for serious aggression, the formation of captive, multimale social groups is desirable for the efficient, long-term, humane housing of chimpanzees in socially and physically enriched environments and for the education of zoo visitors. The University of Texas Science Park (UTSP) has maintained multimale groups of chimpanzees for 14 years. In the UTSP outdoor corral housing, multimale/multifemale social groups of unrelated adult and adolescent chimpanzees (42 F, 46 M) were formed by a series of 397 individual introductions. Wounding aggression was minimal during introductions of females to males or other females and upon male-male introductions of formerly single-caged adolescent and young adult males having had long-term prior visual familiarity. Serious wounding occurred during male-male introductions when there were major discrepancies in the age and social experience of the subjects or when adult, socially experienced males were reintroduced to former group mates following lengthy separations. Male wounding in the eight established long-term groups of 5–11 adults (2–7 males) averaged 1.4 episodes per male-year of residence; 14% of male wounding episodes required surgical therapy. Adult wounding was significantly associated with the presence of one or more group females with maximally tumescent genital swellings. No male-perpetrated infanticides occurred. Not all multimale groupings are successful, but the majority of formerly laboratory-housed chimpanzees may live and reproduce safely in multimale groups. Experience with all-male groups at UTSP suggests that bachelor groups are also practical for long-term housing. © 1995 Wiley-Liss, Inc.     

Attempting to reduce regurgitation and reingestion in a captive chimpanzee through increased feeding opportunities: a case study

Persistent regurgitation and reingestion is an abnormal behavior seen in many captive NHPs, particularly gorillas and chimpanzees. Successful reduction of regurgitation in captive gorillas has been achieved by providing browse, such as branches and leaves, and feeding continuously throughout the day. However, this modality has not been tested in chimpanzees. The authors tested these methods in a 15-year-old male chimpanzee (Pan troglodytes) with a seven-year history of regurgitation and reingestion by alternately providing additional opportunities to forage or giving daily browse. The results show that providing browse on a daily basis models the feeding behavior of wild chimpanzees and is a successful way to reduce regurgitation and reingestion in a chimpanzee unresponsive to previous dietary changes.     

Use of "entertainment" chimpanzees in commercials distorts public perception regarding their conservation status

Chimpanzees (Pan troglodytes) are often used in movies, commercials and print advertisements with the intention of eliciting a humorous response from audiences. The portrayal of chimpanzees in unnatural, human-like situations may have a negative effect on the public's understanding of their endangered status in the wild while making them appear as suitable pets. Alternatively, media content that elicits a positive emotional response toward chimpanzees may increase the public's commitment to chimpanzee conservation. To test these competing hypotheses, participants (n?=?165) watched a series of commercials in an experiment framed as a marketing study. Imbedded within the same series of commercials was one of three chimpanzee videos. Participants either watched 1) a chimpanzee conservation commercial, 2) commercials containing "entertainment" chimpanzees or 3) control footage of the natural behavior of wild chimpanzees. Results from a post-viewing questionnaire reveal that participants who watched the conservation message understood that chimpanzees were endangered and unsuitable as pets at higher levels than those viewing the control footage. Meanwhile participants watching commercials with entertainment chimpanzees showed a decrease in understanding relative to those watching the control footage. In addition, when participants were given the opportunity to donate part of their earnings from the experiment to a conservation charity, donations were least frequent in the group watching commercials with entertainment chimpanzees. Control questions show that participants did not detect the purpose of the study. These results firmly support the hypothesis that use of entertainment chimpanzees in the popular media negatively distorts the public's perception and hinders chimpanzee conservation efforts.     

The Nuts and Bolts of Captive Chimpanzee Diets and Food as Enrichment: A Survey

Beyond planning for good nutrition, one of the Cornerstones of a successful captive-chimpanzee wellness program is the opportunity for the nonhuman primates to express normal feeding and foraging behaviors. This study presents the results of a survey of 11 institutions—8 zoological and 3 research facilities—given to ascertain how daily diets and feeding enrichments were accomplished. Information represents 806 captive chimpanzees, and entails a composite of captive chimpanzee daily diets, forage and browse materials, periodic treats, medicating and supplemental foods, alternative feeding techniques, and food devices. Results suggest facilities are moving away from timed feeding schedules, which tend to increase prefeeding levels of agonism, to feeding a wide variety of foods throughout the day and using a wide variety of feeding techniques.     

Chimpanzee vocal signaling points to a multimodal origin of human language

The evolutionary origin of human language and its neurobiological foundations has long been the object of intense scientific debate. Although a number of theories have been proposed, one particularly contentious model suggests that human language evolved from a manual gestural communication system in a common ape-human ancestor. Consistent with a gestural origins theory are data indicating that chimpanzees intentionally and referentially communicate via manual gestures, and the production of manual gestures, in conjunction with vocalizations, activates the chimpanzee Broca's area homologue--a region in the human brain that is critical for the planning and execution of language. However, it is not known if this activity observed in the chimpanzee Broca's area is the result of the chimpanzees producing manual communicative gestures, communicative sounds, or both. This information is critical for evaluating the theory that human language evolved from a strictly manual gestural system. To this end, we used positron emission tomography (PET) to examine the neural metabolic activity in the chimpanzee brain. We collected PET data in 4 subjects, all of whom produced manual communicative gestures. However, 2 of these subjects also produced so-called attention-getting vocalizations directed towards a human experimenter. Interestingly, only the two subjects that produced these attention-getting sounds showed greater mean metabolic activity in the Broca's area homologue as compared to a baseline scan. The two subjects that did not produce attention-getting sounds did not. These data contradict an exclusive "gestural origins" theory for they suggest that it is vocal signaling that selectively activates the Broca's area homologue in chimpanzees. In other words, the activity observed in the Broca's area homologue reflects the production of vocal signals by the chimpanzees, suggesting that this critical human language region was involved in vocal signaling in the common ancestor of both modern humans and chimpanzees.     

The nuts and bolts of captive chimpanzee diets and food as enrichment: a survey

Beyond planning for good nutrition, one of the Cornerstones of a successful captive-chimpanzee wellness program is the opportunity for the nonhuman primates to express normal feeding and foraging behaviors. This study presents the results of a survey of 11 institutions—8 zoological and 3 research facilities—given to ascertain how daily diets and feeding enrichments were accomplished. Information represents 806 captive chimpanzees, and entails a composite of captive chimpanzee daily diets, forage and browse materials, periodic treats, medicating and supplemental foods, alternative feeding techniques, and food devices. Results suggest facilities are moving away from timed feeding schedules, which tend to increase prefeeding levels of agonism, to feeding a wide variety of foods throughout the day and using a wide variety of feeding techniques.     

No evidence of short-term exchange of meat for sex among chimpanzees

The meat-for-sex hypothesis posits that male chimpanzees (Pan troglodytes) trade meat with estrous females in exchange for short-term mating access. This notion is widely cited in the anthropological literature and has been used to construct scenarios about human evolution. Here we review the theoretical and empirical basis for the meat-for-sex hypothesis. We argue that chimpanzee behavioral ecology does not favor the evolution of such exchanges because 1) female chimpanzees show low mate selectivity and require little or no material incentive to mate, violating existing models of commodity exchange; and 2) meat-for-sex exchanges are unlikely to provide reproductive benefits to either partner. We also present new analyses of 28 years of data from two East African chimpanzee study sites (Gombe National Park, Tanzania; Kanyawara, Kibale National Park, Uganda) and discuss the results of previously published studies. In at least three chimpanzee communities, 1) the presence of sexually receptive females did not increase hunting probability, 2) males did not share preferentially with sexually receptive females, and 3) sharing with females did not increase a male's short-term mating success. We acknowledge that systematic meat sharing by male chimpanzees in expectation of, or in return for, immediate copulations might be discovered in future studies. However, current data indicate that such exchanges are so rare, and so different in nature from exchanges among humans, that with respect to chimpanzees, sexual bartering in humans should be regarded as a derived trait with no known antecedents in the behavior of wild chimpanzees.     

High frequency of leaf swallowing and its relationship to intestinal parasite expulsion in "village" chimpanzees at Bulindi, Uganda

Self-medication by great apes to control intestinal parasite infections has been documented at sites across Africa. Chimpanzees (Pan troglodytes) swallow the leaves of certain plant species whole, without chewing. Previous studies demonstrated a relationship between chimpanzee leaf swallowing and expulsion of nematode worms (Oesophagostomum sp.) and tapeworms (Bertiella sp.) in dung. We investigated the relationship between leaf swallowing and parasite expulsion in chimpanzees inhabiting a fragmented forest-farm mosaic at Bulindi, Uganda. During 13 months whole undigested leaves occurred in chimpanzee dung at a considerably higher frequency (10.4% of dungs) than at other sites (0.4-4.0%). Leaf swallowing occurred year-round and showed no pronounced seasonality. Chimpanzees egested adults of multiple species of Oesophagostomum (including O. stephanostomum) and proglottids of two tapeworms-Bertiella sp. and probably Raillietina sp. The latter may not be a true infection, but the byproduct of predation on domestic fowl. Compared to previous studies, the co-occurrence of whole leaves and parasites in chimpanzee dung was low. Whereas the presence of leaves in dung increased the probability of adult nematode expulsion, no association between leaf swallowing and the shedding of tapeworm proglottids was apparent. Anthropogenic habitat changes have been linked to alterations in host-parasite interactions. At Bulindi, deforestation for agriculture has increased contact between apes and people. Elevated levels of leaf swallowing could indicate these chimpanzees are especially vulnerable to parasite infections, possibly due to environmental changes and/or increased stress levels arising from a high frequency of contact with humans. Frequent self-medication by chimpanzees in a high-risk environment could be a generalized adaptation to multiple parasite infections that respond differently to the behavior. Future parasitological surveys of apes and humans at Bulindi are needed for chimpanzee health monitoring and management, and to investigate the potential for disease transmission among apes, people, and domestic animals.     

Gene flow in wild chimpanzee populations: what genetic data tell us about chimpanzee movement over space and time

The isolation of phylogenetically distinct primate immunodeficiency viruses from at least seven wild-born, captive chimpanzees indicates that viruses closely related to HIV-1 may be endemic in some wild chimpanzee populations. The search for the chimpanzee population or populations harbouring these viruses is therefore on. This paper attempts to answer the question of whether or not such populations of chimpanzees are likely to exist at all, and, if so, where they are likely to be found. We summarize what is known about gene flow in wild populations of chimpanzees, both between major phylogeographical subdivisions of the species, and within these subdivisions. Our analysis indicates that hitherto undocumented reproductively isolated chimpanzee populations may in fact exist. This conclusion is based on the observation that, despite limited geographical sampling and limited numbers of genetic loci, conventional notions of the nature and extent of chimpanzee gene flow have recently been substantially revised. Molecular genetic studies using mitochondrial DNA sequences and hypervariable nuclear microsatellite markers have indicated the existence of heretofore undocumented barriers to chimpanzee gene flow. These studies have identified at least one population of chimpanzees genetically distinct enough to be classified into a new subspecies (Pan troglodytes vellerosus). At the same time, they have called into question the long-accepted genetic distinction between eastern chimpanzees (Pan troglodytes schweinfurthii) and western equatorial chimpanzees (Pan troglodytes troglodytes). The same studies have further indicated that gene flow between local populations is more extensive than was previously thought, and follows patterns sometimes inconsistent with those documented through direct behavioural observation. Given the apparently incomplete nature of the current understanding of chimpanzee gene flow in equatorial Africa, it seems reasonable to speculate that a chimpanzee population or populations may exist which both harbour the putative HIV-1 ancestor, and which have remained reproductively isolated from other chimpanzee populations over the time-scale relevant to the evolution of the SIVcpz-HIV-1 complex of viruses. Continued extensive sampling of wild chimpanzee populations, both for their genes and their viruses, should be performed quickly considering the high probability of extinction that many wild chimpanzee populations face today. The history of human-chimpanzee contacts is discussed.     

The chimpanzee cytochrome P450 3A subfamily: Is our closest related species really that similar?

With the release of the chimpanzee genomic database, much work has been accomplished to understand more fully the closest related species to humans. This study investigates the cytochrome P450 3A (CYP3A) subfamily and examines differences which may be expected between chimpanzees and humans in regards to CYP3A metabolism. A previous publication had reported the presence of five putative chimpanzee CYP3A isoforms, as compared to the four in humans (Williams ET et al., Mol Phylogenet Evol 33, 300–8). Based on the previous report, the chimpanzee CYP3A5 should have had a different C-terminus than its human counterpart; therefore, CYP3A5 and CYP3A67 were cloned. The CYP3A5 clone obtained disputes the previous prediction and confirms that the nucleotide similarity between the two species is 99.7%. While CYP3A67 is most closely related to CYP3A7, with significant differences in the amino acid sequences. Also, the mRNA expression of CYP3A67 can rival the expression of CYP3A4 in the tissues analyzed. CYP3A7 was not found to be expressed in any chimpanzee tissue examined. Total CYP3A protein expression was not significantly different between chimpanzees and humans. Metabolism assays using benzphetamine and erythromycin with chimpanzee liver microsomes did not reveal major differences between chimpanzees and humans. In conclusion, adult CYP3A metabolism may not be significantly different between chimpanzees and humans.     

The perception of self-agency in chimpanzees (Pan troglodytes)

The ability to distinguish actions and effects caused by oneself from events occurring in the external environment is a fundamental aspect of human cognition. Underlying such distinctions, self-monitoring processes are often assumed, in which predicted events accompanied by one's own volitional action are compared with actual events observed in the external environment. Although many studies have examined the absence or presence of a certain type of self-recognition (i.e. mirror self-recognition) in non-human animals, the underlying cognitive mechanisms remain unclear. Here, we provide, to our knowledge, the first behavioural evidence that chimpanzees can perform self/other distinction for external events on the basis of self-monitoring processes. Three chimpanzees were presented with two cursors on a computer display. One cursor was manipulated by a chimpanzee using a trackball, while the other displayed motion that had been produced previously by the same chimpanzee. Chimpanzees successfully identified which cursor they were able to control. A follow-up experiment revealed that their performance could not be explained by simple associative responses. A further experiment with one chimpanzee showed that the monitoring process occurred in both temporal and spatial dimensions. These findings indicate that chimpanzees and humans share the fundamental cognitive processes underlying the sense of being an independent agent.     

Pan paniscus 1973 to 1996: Twenty-three years of field research

Although the pygmy chimpanzee is not smaller than a chimpanzee, it has long intrigued anthropologists with its more gracile build and greater arboreality. The social organization of the pygmy chimpanzee is unique among primates in showing female-bonding among nonrelatives, little male bonding despite male residence, and relative inability of males to outrank females. The crucial ecological basis for this difference from the male-bonded system of chimpanzees is that pygmy chimpanzees do not go through an extended season of low food availability during which party sizes fall to low levels.