Social behavior and genital swelling in pregnant chimpanzees (Pan troglodytes)

Previous reports of increased sexual activity associated with genital swelling during chimpanzee pregnancy have suggested that the tumescent pregnant female may be regarded as a regularly cycling estrous female. Analysis of additional data from seven pregnant and eight cycling females, however, indicated that tumescent pregnant females differed from cycling females in some social interactions with males. As in earlier reports, pregnant females were more receptive to male-initiated copulatory bouts. In addition, pregnant females initiated grooming bouts with males much less frequently than did cycling females, and males spent less time grooming and less frequently inspected the genitals of pregnant than cycling cagemates. This tendency to decrease social interaction during pregnancy is consistent with field reports that pregnant chimpanzees prefer the company of their offspring and other noncycling females. In addition, the genital swelling patterns of 40 chimpanzee pregnancies were analyzed. Length of gestation averaged 231.5 days. Although a high degree of individual variation existed, females were swollen an average of 41% of the days observed, yet reached maximum swelling only 8.7% of the time. Genital swelling was less frequent during the third trimester, averaging 50.5, 47.9, and 22.6% tumescent days in each trimester. Young pregnant females showed maximum swelling more often than did older females. This finding is discussed with respect to field reports of intercommunity transfer of young, tumescent female chimpanzees.     

Anogenital swelling in pregnant chimpanzees of Gombe National Park

Previous studies show that anogenital swelling patterns of pregnant chimpanzees in captivity are correlated with the female's age during pregnancy and are associated with changes in social behavior. The purpose of the present study was to investigate whether the anogenital swelling patterns of free-ranging pregnant chimpanzees resemble those reported for captive chimpanzees. We reviewed the records of 47 pregnancies experienced by 20 female chimpanzees living in Gombe National Park from 1975 to 1992. Percentage scores for swelling during each trimester and for the overall occurrence of various levels of swelling were obtained by dividing the frequency by the total number of days the subject was observed (i.e., 28 days of swelling/165 days observation = 17.2% anogenital swelling for that pregnancy). Subjects exhibited anogenital swelling an average of 25.1% of the days observed during pregnancy, yet reached maximum only 9.4% of the time. Anogenital swelling occurred less frequently as pregnancy progressed, averaging 40.8, 27.3, and 2.7% in the first, second, and third trimesters, respectively. As seen in captive chimpanzees, young pregnant females at Gombe showed anogenital swelling more often than did older ones (r = ?.65, P < .0001). These findings are consistent with reports of behavior associated with anogenital swelling (e.g., intercommunity transfer of young females exhibiting anogenital swelling). © 1993 Wiley-Liss, Inc.     

Male chimpanzee behavior in relation to female ano-genital swelling

This study examined the relationships between male agonistic, affiliative, and sexual behaviors and female estrus condition in captive adolescent and young-adult chimpanzees (Pan troglodytes). Data on agonistic, affiliative, and sexual behaviors of 11 males living in three social groups were collected during daily 45 minute observations over a 5 month period. Female estrus condition was assessed daily using the relative size of the female's ano-genital swelling. It was hypothesized that the presence of maximally tumescent females would generate conflicts between males, so an increase in inter-male agonism was predicted. Males exhibited higher rates of agonism toward other males when at least one female in the group was maximally tumescent. Male affiliative behavior directed toward other males and social play with males were affected by the presence and number of maximally tumescent females. Male sexual behavior increased when maximally tumescent females were present.     

Diurnal Variation in Nutrients and Chimpanzee Foraging Behavior

Primate feeding behavior varies over long (e.g., weekly, seasonally, yearly) and short (e.g., hourly) scales of time due to changes in resource availability and the nutritional composition of foods. While the factors that affect long‐term changes in feeding behavior have received considerable attention, few data exist regarding what drives variability in feeding behavior over the course of a single day. To address this problem, we investigated diurnal variation in chimpanzee feeding on the leaves of two species of saplings, Pterygota mildbraedii and Celtis africana, at Ngogo, Kibale National Park, Uganda. Specifically, we related short‐term changes in chimpanzee feeding behavior on these leaves to diurnal variation in their nutritional composition. Results showed that chimpanzees fed on the leaves of both saplings more in the evening than they did in the morning. The nutritional quality of leaves also improved over the course of the day. Concentrations of cellulose and lignin were lower and total nonstructural carbohydrates (including sugars and starch) were higher in the evening for P. mildbraedii, and sugars were higher in the evening for C. africana. These data suggest that chimpanzees consume these resources when their quality is highest, and consequently, may track the nutrient composition of their foods over very short periods that span only a few hours. In the future, foods collected for analyses must control for time of sampling to ensure biologically meaningful assays of nutrient composition. Am. J. Primatol. 75:342‐349, 2013. © 2013 Wiley Periodicals, Inc.     

Procedures for improving maternal behavior in captive chimpanzees

Captive female chimpanzees who have had no opportunity to observe mothers with infants or to interact with infants often show inappropriate maternal behavior, particularly with their first-born infant, and this usually results in the removal of the infant to be human-reared. The present study used two techniques to encourage appropriate maternal behavior in ten pregnant female chimpanzees. These females were housed together with unrelated infant chimpanzees to adopt, or with lactating female chimpanzees and infants to observe. In five cases both techniques were used, in two cases only the first technique was used, and in three cases only the second technique was used. All ten female chimpanzees showed appropriate maternal behavior when their infants were born, in contrast to a group of eight female chimpanzees who had no such experience whose infants had to be removed for human-rearing. It is suggested that these techniques, or adaptations of them, could be applied to many other captive female chimpanzees with similar results.     

Sources of variation in the nesting behavior of chimpanzees (Pan troglodytes schweinfurthii) in the Budongo forest, Uganda

Data on nesting behavior and nests of the Sonso community of chimpanzees (Pan troglodytes schweinfurthii) living in the Budongo Forest, Uganda, were collected between October 1995 and December 1996. Analysis of the selection of nests shows a preference for particular tree species, especially Cynometra alexandri. Male chimpanzees nested lower than females. Day nests were found to be structurally simpler than night nests and were built in tree species used for feeding, at the same height as feeding activity. Day and night nests serve different functions, and should be clearly distinguished in studies of nesting behavior.     

Chimpanzee identification and social network construction through an online citizen science platform

Citizen science has grown rapidly in popularity in recent years due to its potential to educate and engage the public while providing a means to address a myriad of scientific questions. However, the rise in popularity of citizen science has also been accompanied by concerns about the quality of data emerging from citizen science research projects. We assessed data quality in the online citizen scientist platform Chimp&See, which hosts camera trap videos of chimpanzees (Pan troglodytes) and other species across Equatorial Africa. In particular, we compared detection and identification of individual chimpanzees by citizen scientists with that of experts with years of experience studying those chimpanzees. We found that citizen scientists typically detected the same number of individual chimpanzees as experts, but assigned far fewer identifications (IDs) to those individuals. Those IDs assigned, however, were nearly always in agreement with the IDs provided by experts. We applied the data sets of citizen scientists and experts by constructing social networks from each. We found that both social networks were relatively robust and shared a similar structure, as well as having positively correlated individual network positions. Our findings demonstrate that, although citizen scientists produced a smaller data set based on fewer confirmed IDs, the data strongly reflect expert classifications and can be used for meaningful assessments of group structure and dynamics. This approach expands opportunities for social research and conservation monitoring in great apes and many other individually identifiable species.     

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.     

Leaf-pile pulling: an unusual play pattern in wild chimpanzees

Play in nonhuman animals has generally been viewed as being uniform among study sites. No studies have examined whether there are local variations in play. In this work we report an apparently locality-specific form of play that is basically solo locomotor play, but also has aspects of object play and social play. We describe this unusual "leaf-pile pulling" (LPL) pattern based on video footage of the chimpanzees of Mahale, Tanzania. Typically, when a party of chimpanzees moves in a procession down a slope in the dry season, a youngster will turn around and walk backward while raking many dry leaves with both hands. This activity accumulates many dry leaves while producing a lot of sound. After the player walks 1-15 m, he/she either turns around and walks forward or moves in a somersaulting fashion. The performer usually faces an individual that is immediately following him/her in the procession. The age of the performers ranges from 2 to 22 years, but 3-10 years are most typical. Compared to younger (< 8 years) individuals, older (> or = 8 years) individuals tend to cover longer distances during play, and to be more likely to play only on sloped surfaces and during travel. One of the authors (W.W.) has only seen the behavior exhibited by five different individuals on 10-15 occasions at Gombe. The behavior is exhibited less often at Gombe than at Mahale. To date, LPL has not been reported elsewhere. The absence of LPL at other sites may or may not be explained by environmental differences, such as differences in the availability of dead leaves and sloped terrain. Although the possibility that LPL is acquired by individual learning cannot be completely ruled out, the hypothesis that it is a tradition of the Mahale study group is more likely. This suggests that play may warrant more careful consideration in studies of nonhuman primate culture.     

The development of affiliative and coercive reproductive tactics in male chimpanzees

Like many animals, adult male chimpanzees often compete for a limited number of mates. They fight other males as they strive for status that confers reproductive benefits and use aggression to coerce females to mate with them. Nevertheless, small-bodied, socially immature adolescent male chimpanzees, who cannot compete with older males for status nor intimidate females, father offspring. We investigated how they do so through a study of adolescent and young adult males at Ngogo in Kibale National Park, Uganda. Adolescent males mated with nulliparous females and reproduced primarily with these first-time mothers, who are not preferred as mating partners by older males. Two other factors, affiliation and aggression, also influenced mating success. Specifically, the strength of affiliative bonds that males formed with females and the amount of aggression males directed toward females predicted male mating success. The effect of male aggression toward females on mating success increased as males aged, especially when they directed it toward females with whom they shared affiliative bonds. These results mirror sexual coercion in humans, which occurs most often between males and females involved in close, affiliative relationships.     

Comparison of instantaneous and locomotor bout sampling methods: a case study of adult male chimpanzee locomotor behavior and substrate use.

Currently two methods, instantaneous and locomotor bout sampling, are used most commonly in studies of locomotor behavior. To date, no study has addressed how comparable the results of the two methods are. This paper considers whether different sampling methods of locomotor behavior produce different results. Continuous locomotor bout and instantaneous sampling were carried out simultaneously on each focal animal during a seven month study of chimpanzee positional behavior in the Tai Forest of the Ivory Coast. Results provide two independent sets of data which describe the same events. Results indicate that as locomotor bouts are frequently presented (the percentage of bouts spent in an activity), they overrepresent the frequencies of activities that occur relatively often, but for short distances, and underrepresent activities that have a relatively large mean distance per bout. However, when bouts are weighted with distance, as originally defined by Fleagle (1976b), there are no significant differences in the results obtained by the two methods. Both provide similar results for the frequencies of locomotor activities, frequency of substrate use, and the relationship between substrate and locomotor activity. The advantage of instantaneous sampling is that because it is done at regular intervals, it can easily be carried out in conjunction with sampling other behavioral and ecological data. The advantages of locomotor bout sampling are that it permits the sampling of rare or brief locomotor events and allows for an analysis of sequences of locomotor activities. This paper demonstrates that the two methods can be conducted simultaneously and thus provide the richest return from which the effect of environment and morphology on locomotion can be addressed.     

Mountain gorilla life histories,reproductive competition,and sociosexual behavior and some implications for captive husbandry

New data on modal patterns of, and variability in, mountain gorilla life history tactics and sociosexual behavior, collected during long-term fieldwork at the Karisoke Research Centre, are presented here. These data show that immature males and females develop sociosexual relationships with both peers and adults, and that these relationships—which contribute to more complex social relationships—often lead to mating between natal females and males who are “familiar” partners. They also show that within-group reproductive competition between males can reach considerable levels, but that immature males are sometimes able to copulate with both nulliparous and parous females; that unrelated silverbacked males can reside in the same all-male group but become intolerant of each other if females join the group; and that, whereas females sometimes compete with each other for opportunities of copulate, their probability of conceiving is probably not decreased by any inability to control the mating situation. This information may contribute to continued improvement in the husbandry of captive gorillas.     

Comparative locomotor behavior of chimpanzees and bonobos: The influence of morphology on locomotion

Results from a 10 month study of adult male and female bonobos (Pan paniscus) in the Lomako Forest, Zaire, and those from a 7 month study of adult male and female chimpanzees in the Tai Forest, Ivory Coast (Pan troglodytes verus), were compared in order to determine whether there are species differences in locomotor behavior and substrate use and, if so, whether these differences support predictions made on the basis of interspecific morphological differences. Results indicate that bonobos are more arboreal than chimpanzees and that male bonobos are more suspensory than their chimpanzee counterpart. This would be predicted on the basis of male bonobo's longer and more narrow scapula. This particular finding is contrary to the prediction that the bonobo is a “scaled reduced version of a chimpanzee” with little or no positional behavior difference as had been suggested. This study provides the behavioral data necessary to untangle contradictory interpretations of the morphological differences between chimpanzees and bonobos, and raises a previously discussed (Fleagle: Size and Scaling in Primate Biology, pp. 1–19, 1985) but frequently overlooked point–that isometry in allometric studies does not necessarily equate with behavioral equivalence. Several researchers have demonstrated that bonobos and chimpanzees follow the same scaling trends for many features, and are in some sense functionally equivalent, since they manage to feed and reproduce. However, as reflected in their morphologies, they do so through different types and frequencies of locomotor behaviors. © 1993 Wiley-Liss, Inc.     

Sex differences in adult chimpanzee positional behavior: The influence of body size on locomotion and posture

Focal animal instantaneous sampling of adult male and female chimpanzee positional behavior was conducted during a 7-month study in the Tai Forest, Ivory Coast, in order to determine whether there are sex differences in the locomotion, posture, substrate use, and height preference of sexually dimorphic adult chimpanzees, and if so, whether these differences support predictions based on body size differences. Results indicate that as predicted, adult male and female chimpanzees differ in their arboreal locomotor behavior, with the larger males using less quadrupedalism and more climbing, scrambling, and aided bipedalism than females during feeding locomotion. There is a sex difference in height preference as well, with female chimpanzees consistently using more arboreal behavior than males, primarily during resting. Although it has been previously demonstrated that separate primate species of differing body size differ in locomotor and postural activities (Fleagle and Mittermeier, 1980; Crompton, 1984), this study clearly demonstrates that body size differences within a species can also be correlated with differences in locomotor behavior. These findings may influence how we interpret sex differences in body size of extinct species. © 1993 Wiley-Liss, Inc.     

The diet of chimpanzees in the Budongo Forest Reserve, Uganda

Chimpanzee diets are highly variable, but in all cases dominated by ripe fruit. Previous work has suggested that in the Budongo Forest Reserve, Uganda, ripe fruit remains available throughout the year, but dietary data for chimpanzees at this site remain sparse. This paper details the diet of a recently habituated community of Budongo chimpanzees living in the Sonso region. Chimpanzees of this community spent 64.5% of their feeding time eating fruit, and 19.7% eating arboreal leaves. Terrestrial herbaceous vegetation formed only a minor component of the diet. In these general dietary characteristics, Budongo chimpanzees show greater similarity to those of the Gombe National Park than they do to those in the geographically closer and floristically more similar Kibale Forest National Park. The diet was dominated by the fruit of four species and leaves of two species, although the composition of the diet varied from month to month, remaining diverse. Figs were consumed throughout much of the year, and in consequence, should perhaps be regarded as a staple, rather than fallback, food. A period of food scarcity was not apparent in this study and future comparative nutritional investigation will be required to determine whether these chimpanzees face a time of dietary hardship.     

Deciduous dentition eruption sequence of the laboratory-reared chimpanzee (Pan troglodytes)

Eruption sequence and age ranges for the maxillary and mandibular deciduous dentition were generated from serial dental study models from 28 laboratory-reared chimpanzees (Pan troglodytes). These data can serve as additional normative, baseline information for the evaluation of fetal insults on neonatal growth and development in the laboratory-reared chimpanzee.     

Some observations on the sociosexual behavior of a captive male Yangtze finless porpoise calf (Neophocaena phocaenoides asiaeorientalis)

Sociosexual behavior in a captive male Yangtze finless porpoise calf (Neophocaena phocaenoides asiaeorientalis) was studied during its first year postpartum. This behavior was first observed at 1 month postpartum. During the year, (1) each event of the sociosexual behavior typically lasted 6–9 s and the frequency was 2.8 occurrences h^?1; (2) ventral–ventral was the predominant sociosexual gesture; (3) prior to sociosexual interactions, the calf generally bonded with the receiver; (4) the mother was his preferred sociosexual partner but other adults of both sexes were also involved; (5) the frequency of sociosexual behavior decreased until the 9th month postpartum and increased thereafter.     

The effects of infant births on the sociosexual behavior and hormonal patterns of a cooperatively breeding primate (Cebuella pygmaea)

This study examined changes in the behavioral and hormonal patterns of cooperatively breeding pairs in a primate species with the passing of time and with specific reproductive events. We (1) compared patterns of sexual, agonistic, and affiliative behavior of newly paired pygmy marmosets with the same behavioral patterns immediately after the birth of their first set of infants; (2) determined if postpartum behavioral differences existed between pairs whose infants lived and those whose infants died; and (3) examined whether behavioral patterns changed over the course of ovarian cycles in parous pygmy marmosets as had been documented in nulliparous pairs. The behavior of pairs was recorded during daily half-hour focal samples for 60 days after pairing, and 30 or 60 days after the birth of infants for pairs whose litters died or lived, respectively. Daily urine samples from females during the study were analyzed for luteinizing hormone and pregnanediol glucuronide concentrations to determine dates of ovulation. The results indicated that males consistently altered their sexual behavior and olfactory monitoring of mates during periovulatory periods in the females' cycles both postpairing and postpartum, while similar rates of social and sexual behavior were maintained between the conditions. Sexual behavior occurred throughout the females' ovarian cycles. Peaks in sexual behavior during the periovulatory period in nulliparous pairs disappeared after the birth of infants. Pairs whose infants died showed higher rates of sexual behavior than pairs with surviving infants. Social and sexual behavior may function to maintain the relationship both during and outside of ovulation, especially with the loss of infants. © 1996 Wiley-Liss, Inc.