Interspecific interactions between wild pygmy chimpanzees (Pan paniscus) and red colobus (Colobus badius)

Interspecific interactions accompanied by physical contacts between wild pygmy chimpanzees (Pan paniscus) and red colobus (Colobus badius) were observed on three occasions at Wamba, Republic of Zaire. In all cases, the red colobus initiated the interactions by approaching the pygmy chimpanzees. Most of the pygmy chimpanzees, which were within 5 m of the red colobus, were juveniles or infants but the adult male pygmy chimpanzees never showed any interest in the red colobus. The red colobus groomed the chimpanzees in two cases, but the latter never groomed the former. No true aggressive interactions were observed between the two species. The lack of any evidence of hunting of red colobus through longitudinal studies of the pygmy chimpanzees of Wamba, together with the present observations, suggests that red colobus are probably not targets of hunting by the pygmy chimpanzees.     

The behavior of a group of captive pygmy chimpanzees (Pan paniscus)

A group of captive pygmy chimpanzees (Pan paniscus) was studied in the San Diego Zoological Gardens. The behavior patterns that these animals exhibit are described. Each of these behavior patterns is compared to those described for wild and captive common chimpanzees (P. troglodytes). Differences in behavior between these two species are attributed to specialization of the pygmy chimpanzee to a rain forest habitat and to a monogamous social system.     

Seed dispersal by pygmy chimpanzees (Pan paniscus): A preliminary report

The role in seed dispersal played by the pygmy chimpanzees (Pan paniscus) inhabiting Wamba, Republic of Zaïre, was studied. Germination was tested for seeds of 17 plant species recovered from the feces of pygmy chimpanzees at Wamba. The fecal seeds of 13 species germinated, and in six of the species the germination rate for the fecal seeds was higher than that of control seeds. Although five other species showed a higher germination rate in the control seeds than in the fecal seeds, the remaining two species revealed no difference in germination rate between the fecal and control seeds. There was no great difference in germination velocity between the fecal and control seeds of the same species. For comparison, seeds of four plant species collected from the feces of common chimpanzees (Pan troglodytes) and gibbons (Hylobates lar) in captivity in Okinawa were tested for their germinability. In this test, although the seeds had passed through the digestive tract, their germinability demonstrated little change. Based on the behavioral characteristics of the pygmy chimpanzee at Wamba and observations of the captive primates on Okinawa, it seems that pygmy chimpanzees may play an important role in the seed dispersal of fruit plant species at Wamba.     

Social behavior of wild pygmy chimpanzees (Pan paniscus) of Wamba: A preliminary report

The author studied wild pygmy chimpanzees (Pan paniscus) for about eight months on two separate expeditions between 1975 and 1977, in the study area located at Wamba, Zone de Djolu, in the Republic of Zaire. Most of this paper is based on data obtained through observing the pygmy chimpanzees in their natural surroundings. However, some of them were provisioned and frequently visited the feeding area. Some portion of the data on groupings and food sharing were gathered there. Pygmy chimpanzees groupings, although similar to the common chimpanzee groupings, have larger temporary associations that are almost exclusively bisexual. At least in some greeting contexts, they exhibit patterns of behavior distinctly different from the common chimpanzee. A preliminary investigation of their social interactions revealed strong bonds between males and females, and high female sociability, in contrast to the strong male bonds and female unsociability seen among common chimpanzees.     

Epidermal patterns of the hands and feet of the pygmy chimpanzee (Pan paniscus)

The characteristics of the epidermal ridge system were studied in a series of eighteen lesser or pygmy chimpanzees (Pan paniscus). The general ridge alignments are very similar to those of the chimpanzee (Pan troglodytes); Biegert ('61). On the average the pattern intensity (P.I.) of the palm configurations is considerably higher in the pygmy chimpanzee than in the chimpanzee, thus representing the highest total palm pattern intensity of all species of the Hominoidea. The sole configurations show parallel main results to those of the palm; however, the decreased sole pattern frequency of the pygmy chimpanzee is of a smaller predominance only as compared to the values of the other species of this superfamily. The preliminary data on the finger tip patterns, translated into P.I. values, are much higher than in chimpanzees and within the range of the mean values of gorillas (Brehme, '73), while those of the toes of pygmy chimpanzees seem to possess the lowest P.I. values of the African apes.     

Reactions of a group of pygmy chimpanzees (Pan paniscus) to their mirror-images: Evidence of self-recognition

A group of seven pygmy chimpanzees (Pan paniscus) was tested for their mirror-image reactions during a ten-day experiment. The time spent viewing the mirror waned quickly. Little social responses directed towards the mirror were observed. Self-directed behaviors were shown from testday one on. It was concluded that four out of seven animals could correctly identify their mirror-image, one infant was not (yet) able to do so, and for two other individuals the results were inconclusive.     

Variations in the serum esterases of chimpanzees

Seventy chimpanzees, representing Pan paniscus (the pygmy chimp) and all four races of the common chimpanzee (Pan troglodytes), were examined for electrophoretic variation in their serum esterases. Only three variant electromorphs were observed at these six loci. One of these was at the albumin-associated esterase locus, and the other two were at esterase 4. This is the first report of any variation in an esterase or albumin in chimpanzees.     

Ontogeny and the evolution of adult body size dimorphism in apes

This analysis investigates the ontogeny of body size dimorphism in apes. The processes that lead to adult body size dimorphism are illustrated and described. Potential covariation between ontogenetic processes and socioecological variables is evaluated. Mixed-longitudinal growth data from 395 captive individuals (representing Hylobates lar [gibbon], Hylobates syndactylus [siamang], Pongo pygmaeus [orangutan], Gorilla gorilla [gorilla], Pan paniscus [pygmy chimpanzee], and Pan troglodytes [“common” chimpanzee]) form the basis of this study. Results illustrate heterogeneity in the growth processes that produce ape dimorphism. Hylobatids show no sexual differentiation in body weight growth. Adult body size dimorphism in Pongo can be largely attributed to indeterminate male growth. Dimorphism in African apes is produced by two different ontogenetic processes. Both pygmy chimpanzees (Pan paniscus) and gorillas (Gorilla gorilla) become dimorphic primarily through bimaturism (sex differences in duration of growth). In contrast, sex differences in rate of growth account for the majority of dimorphism in common chimpanzees (Pan troglodytes). Diversity in the ontogenetic pathways that produce adult body size dimorphism may be related to multiple evolutionary causes of dimorphism. The lack of sex differences in hylobatid growth is consistent with a monogamous social organization. Adult dimorphism in Pongo can be attributed to sexual selection for indeterminate male growth. Interpretation of dimorphism in African apes is complicated because factors that influence female ontogeny have a substantial effect on the resultant adult dimorphism. Sexual selection for prolonged male growth in gorillas may also increase bimaturism relative to common chimpanzees. Variation in female growth is hypothesized to covary with foraging adaptations and with differences in female competition that result from these foraging adaptations. Variation in male growth probably corresponds to variation in level of sexual selection. © 1995 Wiley-Liss, Inc.     

Comparing maternal styles in bonobos (Pan paniscus) and chimpanzees (Pan troglodytes)

Studies on Cercopithecine primate maternal styles, using factor analysis on a set of maternal behaviors, commonly render two factors that describe separate dimensions of maternal behavior: protectiveness and rejection. The aims of this study were to 1) investigate whether this method for determining maternal styles in Cercopithecine species can be applied to bonobos (Pan paniscus) and chimpanzees (Pan troglodytes), 2) determine whether they follow the same pattern, and 3) assess whether species differences in maternal style are apparent. We performed a factor analysis on nine maternal behaviors using data on eight mother-infant pairs of each species. This resulted in three factors: protectiveness, distance, and refusal. Protectiveness is positively correlated with time spent in ventral contact, making contact, approaching, and restraining. Distance is positively related with breaking contact and leaving. Refusal is positively correlated with rejecting and nipple-rejecting. The pattern of protectiveness corresponds with the pattern found in Cercopithecine species, suggesting a high consistency of this dimension across species and higher taxa. The retention of the other two factors indicates that in the Pan species, breaking contact and leaving represent another dimension, apart from rejecting and nipple-rejecting, which usually fall under one dimension in Cercopithecine species. An interspecific comparison of the factor scores for each dimension of maternal behavior reveals that, on average, bonobos and chimpanzees score equally on protectiveness. Scores on distance increase positively with infant age in chimpanzees, and negatively in bonobos, and on average bonobos have higher scores on refusal. These interspecies differences in maternal style are discussed in the light of interspecies differences in infant development, infant vulnerability to aggression, interbirth intervals, and female sociality.     

Ontogeny of body size variation in African apes

Size variation in African apes (Gorilla gorilla [gorilla], Pan paniscus [pygmy chimpanzee], and Pan troglodytes [“common” chimpanzee]) is substantial, both within and between species. We investigate the possible evolutionary significance of this variation through an analysis of the ontogeny of size variation in this group. In addition, we highlight possible areas of future endocrinological research, and evaluate recently proposed alternative models that attempt to account for ontogenetic variation in apes. The present study shows that intergeneric variation in size is largely a consequence of differences among species in the rate of body weight growth. Interspecific size variation in Pan is a product of both rate and duration differences in growth. The ontogenetic bases of sexual dimorphism vary in this group. Dimorphism is largely a result of sex differences in the duration of body weight growth in gorillas and pygmy chimpanzees, but results from differences in the rate of growth in common chimpanzees. Ontogenetic analyses largely confirm earlier interpretations, but with better data and methods. The great degree of ontogenetic variation within and among these species, especially in the timing and magnitude of “pubertal” growth spurts, implies that studies of endocrine growth control in African apes could be a productive line of future research. We also suggest that ontogenetic variation can be understood with respect to ecological risks. Growth rates seem to be negatively correlated with ecological risk in African apes, suggesting links between ontogenetic patterns and social and ecological variables. High growth rates in gorillas compared to Pan are most consistent with this model. Variation between chimpanzees and pygmy chimpanzees (especially females) also seem to fit predictions of this model. © 1996 Wiley-Liss, Inc.     

Grouping patterns of wild pygmy chimpanzees (Pan paniscus) observed at a marsh grassland amidst the tropical rain forest of Yalosidi,Republic of Zaïre

Fixed point observation to ascertain the grouping patterns of wild pygmy chimpanzees was carried out at a marsh grassland known locally as Iyoko amidst the tropical rain forest of Yalosidi, Republic of Zaïre. The chimpanzees were seen alone or in parties consisting of up to 32 individuals. The mean size of parties which arrived at Iyoko was 7.9 (N=67), although the modal party size was 2–5. The majority (76%) of all observed parties including those that reformed after joining/parting while staying at Iyoko (N=96) was of the mixed type, i.e., consisting of adult male(s), adult female(s), and dependent individual(s). There was a tendency for parties to be composed of approximately equal numbers of adult males and females. All “social” activities such as sexual behavior and branch-dragging displays were recorded only in mixed parties consisting of more than ten individuals. The joining of parties of pygmy chimpanzees after arriving separately at Iyoko was seen 13 times and the parting of those before departing from Iyoko occurred seven times in total. In contrast, antagonistic encounters between two parties were recorded twice. These observations suggest that the joining/parting between parties is an intra-unit-group phenomenon while antagonistic encounters between parties are inter-unit-group interactions. It was assumed that at least two such unit-groups of pygmy chimpanzees consisting of 80–90 individuals in total utilized Iyoko without intermingling with each other. A comparison on grouping patterns among three pygmy chimpanzee populations (Yalosidi, Wamba, and Lomako) indicates that in terms of basic social organization they show many similarities except for the mean unit-group size, the mean party size, and the modal party size. Perhaps differences in the unit-group size were simply reflected as a whole in the differences of the mean party size as well as the modal party size observed across the three populations.     

The social group of pygmy chimpanzees (Pan paniscus) of Wamba

This is a sociological study of the pygmy chimpanzees of Wamba which have been provisioned sufficiently since 1977. The society of the pygmy chimpanzee is basically similar to that of the common chimpanzee in the flexibility in forming parties and in the existence of a definite social unit, i.e.,unit group. It also resembles in that young females play an important role in exchanging members among unit groups, thus, as transmitters of genes from group to group, while males do in integrating groups, delimiting the social boundaries of unit groups through their inter-group antagonism. Pygmy chimpanzees differ from common chimpanzees in fission and fusion pattern of parties. The former generally forms parties larger in size and more homogeneous in composition than the latter. Any party of the former has both reproductive and nursing functions, while the parties of the latter are categorized into various types according to functions and compositions. This uniformity in pygmy chimpanzee parties seems to be well maintained by the females' high sexual ability to manifest semi-continual estrus.     

Hand abnormalities in pygmy chimpanzees (Pan paniscus)

Hand deformities in 2 wild-born pygmy chimpanzees were investigated through dissection, linear measurements, bone mineral analysis and X-rays. Weight of tissue (skin, muscle, bone) was determined for each body segment (hand, forearm, upper arm), permitting comparison between the abnormal and normal upper limbs. In one individual, the right and left forearms and hands differed in tissue weights and bone mineral content. In the other, the hands differed in tissue composition, but not the forearms.     

Social relationships between immigrant and resident bonobo (Pan paniscus) females at Wamba.

Social relations and behaviours of adolescent female pygmy chimpanzees (Pan paniscus) that migrated between unit groups were studied at Wamba, Za?re. Each immigrant female selected one particular resident female, a 'specific senior female' (SSF) and frequently approached and followed her. Affinitive behaviours (e.g. grooming, genitogenital rubbing) were frequently performed between the immigrant and her SSF. Most affinitive behaviours were more likely to be initiated by the immigrant than by her SSF. The immigrant's relationship with her SSF seemed to facilitate the immigrant's integration into the new unit group. Resident males actively approached and mated with the immigrant females. Copulation also appeared to further the immigrant's stable relationship with the resident males.     

An ecological study of the pygmy chimpanzees (Pan paniscus) of Yalosidi,Republic of Zaire

The pygmy chimpanzees of Yalosidi, near the southeastern limit of the range of this species, differ in group size, food habit, nesting behavior, response to humans, etc., from those of other localities (Wamba, Lomako, and Lake Tumba). It is inferred that the pygmy chimpanzee has a wide range of ecological variation. A comprehensive use of the habitat was characteristic of the Yalosidi pygmy chimpanzees. Every stratum of every vegetation type in their habitat was used by them for feeding, resting, or sleeping. The wide ecological niche of pygmy chimpanzees probably permits their survival in competition with other nonhuman primates of Yalosidi, which are more agile but confined to more limited niches.     

Development of guidelines for assessing obesity in captive chimpanzees (Pan troglodytes)

Many captive chimpanzees (Pan troglodytes) are subjectively considered to be overweight or obese. However, discussions of obesity in chimpanzees are rare in the literature, despite the acknowledged problem. No study to date has systematically examined obesity in captive chimpanzees. This project develops guidelines for defining obesity in captive chimpanzees through the examination of morphometric and physiologic characteristics in 37 adult female and 22 adult male chimpanzees. During each animal's biannual physical exam, morphometric data was collected including seven skinfolds (mm), body mass index (BMI), waist‐to‐hip ratio (WHR), and total body weight (kg). The morphometric characteristics were correlated with triglycerides and serum glucose concentration, to test the utility of morphometrics in predicting relative obesity in captive chimpanzees. Abdominal skinfold (triglyceride: F=3.83, P=0.05; glucose: F=3.83, P=0.05) and BMI (triglyceride: F=10.42, p=0.003; glucose: F=6.20, P=0.02) were predictive of increased triglycerides and serum glucose in females; however no morphometric characteristics were predictive of relative obesity in males. Results suggest that no males in this population are overweight or obese. For females, there were additional significant differences in morphometric (skinfolds, BMI, WHR, total body weight) and physiologic measurements (systolic and diastolic blood pressure, red blood cells) between individuals classified overweight and those classified non‐overweight. Skinfold measurements, particularly abdominal, seem to be an accurate measure of obesity and thus potential cardiovascular risk in female chimpanzees, but not males. By establishing a baseline for estimated body fat composition in female captive chimpanzees, institutions can track individuals empirically determined to be obese, as well as obesity‐related health problems. Zoo Biol 0:1–12, 2007. © 2007 Wiley‐Liss, Inc.     

Predation on mammals by the chimpanzee (Pan troglodytes)

With respect to prey selectivity and predation frequency, chimpanzees (Pan troglodytes) show local differences as well as diachronic variability within the same population. When data on predation from three long-term studies at Mahale, Gombe, and Tai are compared, some differences and similarities emerge; Mahale is more like Gombe than Tai in regard of prey selection but features of hunting at Tai with respect to predation frequency are not conspicuous. The most responsible factor for diversity in prey selectivity is a distinct “prey image” maintained by chimpanzees of different populations, although it is necessary to clarify in future studies why and how such tradition develops. Relative body size of chimpanzees to prey species and/or the degree of cooperation among members of a hunting party may explain the variability in prey size selected at each site, the latter influencing the frequency of successful hunts at the same time. Although various degrees of habituation and different sampling methods including artificial feeding might have obscured the real differences, recent data from the three populations do not seem to be biased greatly by such factors. Nevertheless, it is still difficult to make strict comparisons due to the lack of sufficient standardized data across the three populations on the frequency of hunting and predation. It is suggested that the size or demographic trend of a chimpanzee unit-group, especially the number of adult males included, necessarily influences its hunting frequency as well as its prey profile. It is also suggested that factors which bring these males together into a party (e.g. fruit abundance, swollen females, conflict between unit-groups etc.) strongly affect theactual hunting and kill rates. Other possible factors responsible for the local differences are forest structure (e.g. tree height), skilful “hero” chimpanzees, and competition with sympatric carnivorous animals. A total of at least 32 species have been recorded as prey mammals of chimpanzees from 12 study sites and the most common prey mammals are primates (18 species), of which 13 species are forest monkeys. Forest monkeys, colobine species in particular, are often the most common victims of the predation by chimpanzees at each site. We may point out a tendency toward selective hunting for the forest monkeys in terms of the selectivity of prey fauna among all three subspecies of chimpanzees, including populations living in drier environment. The mode of chimpanzee hunting seems to correspond to the highest available biomass of gregarious, arboreal monkeys in the forest, colobine species in particular. In contrast, bonobos (P. paniscus) are less carnivorous than chimpanzees, only rarely preying on a few species of small mammals. The sharp contrast of the two allied species in their predatory tendencies appears to have something to do with the differences in the structure of primary production between their habitats.     

Comparing infant and juvenile behavior in bonobos (Pan paniscus) and chimpanzees (Pan troglodytes): a preliminary study

The dichotomy between the two Pan species, the bonobo (Pan paniscus) and chimpanzee (Pan troglodytes) has been strongly emphasized until very recently. Given that most studies were primarily based on adult individuals, we shifted the “continuity versus discontinuity” discussion to the infant and juvenile stage. Our aim was to test quantitatively, some conflicting statements made in literature considering species differences between immature bonobos and chimpanzees. On one hand it is suggested that infant bonobos show retardation in motor and social development when compared with chimpanzees. Additionally it is expected that the weaning process is more traumatic to chimpanzee than bonobo infants. But on the other hand the development of behaviors is expected to be very similar in both species. We observed eight mother–infant pairs of each species in several European zoos. Our preliminary research partially confirms that immature chimpanzees seem spatially more independent, spending more time at a larger distance from their mother than immature bonobos. However, the other data do not seem to support the hypothesis that bonobo infants show retardation of motor or social development. The development of solitary play, environmental exploration, social play, non-copulatory mounts and aggressive interactions do not differ between the species. Bonobo infants in general even groom other group members more than chimpanzee infants. We also found that older bonobo infants have more nipple contact than same aged chimpanzees and that the weaning process seems to end later for bonobos than for immature chimpanzee. Additionally, although immature bonobos show in general more signs of distress, our data suggest that the weaning period itself is more traumatic for chimpanzees.     

Natural pathology of the captive chimpanzee (Pan troglodytes): A 35‐year review

We present the spontaneous pathological lesions identified as a result of necropsy or biopsy for 245 chimpanzees (Pan troglodytes) over a 35‐year period. A review of the pathology database was performed for all diagnoses on chimpanzees from 1980 to 2014. All morphologic diagnoses, associated system, organ, etiology, and demographic information were reviewed and analyzed. Cardiomyopathy was the most frequent lesion observed followed by hemosiderosis, hyperplasia, nematodiasis, edema, and hemorrhage. The most frequently affected systems were the gastrointestinal, cardiovascular, urogenital, respiratory, and lymphatic/hematopoietic systems. The most common etiology was undetermined, followed by degenerative, physiologic, neoplastic, parasitic, and bacterial. Perinatal and infant animals were mostly affected by physiologic etiologies and chimpanzee‐induced trauma. Bacterial and physiologic etiologies were more common in juvenile animals. Degenerative and physiologic (and neoplastic in geriatric animals) etiologies predominated in adult, middle aged, and geriatric chimpanzees.     

Dimensions and moment arms of the hind- and forelimb muscles of common chimpanzees (Pan troglodytes).

This paper supplies quantitative data on the hind- and forelimb musculature of common chimpanzees (Pan troglodytes) and calculates maximum joint moments of force as a contribution to a better understanding of the differences between chimpanzee and human locomotion. We dissected three chimpanzees, and recorded muscle mass, fascicle length, and physiological cross-sectional area (PCSA). We also obtained flexion/extension moment arms of the major muscles about the limb joints. We find that in the hindlimb, chimpanzees possess longer fascicles in most muscles but smaller PCSAs than are predicted for humans of equal body mass, suggesting that the adaptive emphasis in chimpanzees is on joint mobility at the expense of tension production. In common chimpanzee bipedalism, both hips and knees are significantly more flexed than in humans, necessitating muscles capable of exerting larger moments at the joints for the same ground force. However, we find that when subject to the same stresses, chimpanzee hindlimb muscles provide far smaller moments at the joints than humans, particularly the quadriceps and plantar flexors. In contrast, all forelimb muscle masses, fascicle lengths, and PCSAs are smaller in humans than in chimpanzees, reflecting the use of the forelimbs in chimpanzee, but not human, locomotion. When subject to the same stresses, chimpanzee forelimb muscles provide larger moments at the joints than humans, presumably because of the demands on the forelimbs during locomotion. These differences in muscle architecture and function help to explain why chimpanzees are restricted in their ability to walk, and particularly to run bipedally.