Food calling by captive bonobos (Pan paniscus): An experiment

We examined (i) whether bonobos display a specific food-calling behavior when discovering a hidden food resource, (ii) whether the presence of competitors affects this behavior, and (iii) whether food quantity or gender influences its appearance. We carried out experiments (n = 108) within a captive group of eight bonobos at the Animal Park Planckendael (Mechelen,Belgium). We hid highly preferred food items (n = 7 or 25) in their enclosure and recorded vocal behavior and interactions between discoverer and group members. As a control, we gave the same number of items to the individuals when isolated from the group, a situation without potential food competition (n = 38). The only vocalization frequently uttered by the discoverer was the food peep. They uttered food peeps significantly more often when no food competition was possible. The amount of food had no significant influence on whether food peeps were uttered. The same applies to the individuals’ identity or gender. Although the costs of food calling behavior seemed much higher for males, both sexes uttered food calls to the same extent. We hypothesize thai males signal food presence in order to attract potential mates and are willing to give up the discovered food resource in return for sex: sex for food exchange. In contrast, females may vocalize to attract coalition partners. Through these coalitions, they can monopolize food resources vis-à-vis males. It is also possible that females have less reason to suppress food calk, since they are dominant to males. This study suggests that bonobos are able to give shaded signals about their environment and have the potential to communicate this information in order to promote their sexual strategy.     

A review of terminology on aggregation patterns in bonobos (Pan paniscus)

We review terms that describe the levels of group structure in Pan paniscus.We discuss points of confusion that result from unclear definition of terms, which could lead to inaccurate intra- and interspecific comparisons. We recommend terms that are applicable to captive populations.     

Structure and use of distance calls in wild bonobos (Pan paniscus)

We report the physical structure and use of a distance call (high-hoot) by wild bonobos (Pan paniscus).Although spectrographic analyses reveal high structural variability, the total sample can be subdivided according to the composition of units—the presence or absence of an initial segment—and the range of the lowest harmonic. Analyses of samples from male—female pairs,vocalizing simultaneously and in close proximity, reveal that both animals utter calls in more or less precise temporal alternation but with different spectral ranges. Whether these differences are gender-specific or related to other factors, such as age or the social relations between particular individuals, is not clear. We suggest that (a) individuals of the same party may coordinate their vocal activity on both the temporal and the spectral level and (b) high hootings stimulate emission of equal vocalizations by members of other parties and may increase cohesion among community members. Comparison of a restricted number of spectrograms from known individuals indicates that bonobos may be able to adjust spectral parameters of one type of distance calls (high- hoot) according to corresponding calls of conspecifics.     

Comparison of behavioral sequence of copulation between chimpanzees and bonobos

We compared sex differences in behaviors leading to copulation of chimpanzees (Pan troglodytes) in the Kalinzu Forest, Uganda with those of bonobos (Pan paniscus) at Wamba, D.R. Congo, using the same definition. Female chimpanzees were more likely to initiate copulation than female bonobos. While most of copulations (96%) were initiated by males in bonobos, among chimpanzees only 63% of copulations were initiated by males. Female bonobos initiated an interaction leading to copulation when males approached them within a short distance. On the other hand, both male and female chimpanzees initiated behavior at a longer distance. Higher proceptivity and a higher copulation rate during the maximal swelling period of female chimpanzees might suggest that they gain greater benefits from a high frequency of copulations than do female bonobos.     

More reliable estimates of divergence times in Pan using complete mtDNA sequences and accounting for population structure

Here, we report the sequencing and analysis of eight complete mitochondrial genomes of chimpanzees (Pan troglodytes) from each of the three established subspecies (P. t. troglodytes, P. t. schweinfurthii and P. t. verus) and the proposed fourth subspecies (P. t. ellioti). Our population genetic analyses are consistent with neutral patterns of evolution that have been shaped by demography. The high levels of mtDNA diversity in western chimpanzees are unlike those seen at nuclear loci, which may reflect a demographic history of greater female to male effective population sizes possibly owing to the characteristics of the founding population. By using relaxed-clock methods, we have inferred a timetree of chimpanzee species and subspecies. The absolute divergence times vary based on the methods and calibration used, but relative divergence times show extensive uniformity. Overall, mtDNA produces consistently older times than those known from nuclear markers, a discrepancy that is reduced significantly by explicitly accounting for chimpanzee population structures in time estimation. Assuming the human–chimpanzee split to be between 7 and 5 Ma, chimpanzee time estimates are 2.1–1.5, 1.1–0.76 and 0.25–0.18 Ma for the chimpanzee/bonobo, western/(eastern + central) and eastern/central chimpanzee divergences, respectively.     

Vocal Production by a Language-Competent Pan paniscus

Human spoken language and nonhuman primate vocalization systems have traditionally been regarded as qualitatively different from one another with respect to their semanticity and the way in which individuals acquire and utilize these signals. However, recent studies of the vocal behaviors of both captive and free-ranging monkeys and apes suggest that this dichotomy may not be unequivocal. We examined the vocalizations produced by a linguistically-competent adult male bonobo (Pan paniscus) named Kanzi. We analyzed his vocalizations during communicative interactions with humans in order to determine whether they vary systematically according to the semantic context in which they are produced. We determined semantic contexts based upon a vocalization's co-occurrence with predefined behavioral correlates. Spectrographic and statistical analyses revealed that acoustic structure is similar among the vocalizations that occurred within a specific semantic context and structural differences are evident between the vocalizations produced in different contexts. The results provide evidence that, during communicative interactions with humans, Kanzi modulates his vocal output on both the temporal and spectral levels.     

Sex differences in copulation attempts in wild bonobos at Wamba

We examined sex differences in copulation attempts in a group of wild bonobos at Wamba, Congo, by analyzing the behavioral sequence. Most copulation attempts were initiated by approach or courtship behaviors by males. Males showed these behaviors when they were more than 5 m from females, whereas females did so only when males solicited them from within 5 m. Most copulations involved females showing perineal swelling, because males solicited those females more frequently and those females accepted copulation more frequently than did females in the non-swelling phase. Nevertheless, males solicited females in the non-swelling phase in one-third of copulation attempts, and those females accepted copulation in half of those attempts. This is markedly different from chimpanzees, in which sexual behaviors almost exclusively involve females in the swelling phase. The perineum of female bonobos during the non-swelling phase is soft and wrinkled but fairly large, which may attract males to some extent. The low, but existing, attractiveness and receptivity of female bonobos during the non-swelling phase might have evolved to control sexual competition among males and provide higher social status for females.     

Sexual competition in a group of captive bonobos (Pan paniscus)

We describe the occurrence of sexual competition, expressed as harassment of sexual interactions in a captive group of bonobos. We monitored all aggressive and pestering interventions during sexual interactions of three captive adult females, one adolescent, and three adult males. The study period covered two complete menstrual cycles for each female, with continuous daily observations. There was relatively little overt sexual competition by the males, in analogy with other studies. Most male interventions occurred towards interactions with the alfa female. The alfa female performed the most intense and the highest number of interventions towards the sexual interactions of the other females. The data provide evidence for female intra-sexual competition in this female dominant species.     

Function and distribution of coalitions in captive bonobos (Pan paniscus)

We examined the distribution of support behaviour within a captive group of bonobos. Most support was evoked by inter-sexual conflicts with the two highest ranking females. Within a dyad, the usual winner was most often supported. Individuals that challenged the rank order by aggressions and pestering were aggressed more often by their targets in the company of an ally. The two lowest ranking males served as scapegoats, receiving 80% of the contra-support. In coalitions, inviduals did not aggress victims they would not dare to attack without supporters. However, the victims of coalitions reacted more strongly with fear and rarely counteraggressed than when being attacked alone, indicative of the high impact of aggression in support. The alpha female showed some control behaviour when intervening in conflicts. The data fitted with several functional hypotheses: coalitions functioned to maintain existing ranks, to acquire ranks, to reduce tension, and to test or strengthen the bond. We suggest that support behaviour fulfilled a crucial role in the maintenance of the power of the two highest ranking females over the males. Among the females themselves the dominance relationships were not based on coalitions, but on individual attributes.     

Vocal coordination of troop movement among white-faced capuchin monkeys,Cebus capucinus

Coordinated travel by social groups is well documented, often with evidence that cognitive spatial maps are employed. Yet the mechanisms by which movement decisions are made and implemented within social groups are poorly known. In a field study of white-faced capuchin monkeys in Costa Rica it was demonstrated that a specific call, the “trill,” is used by adults in the initiation and directing of troop movement. The trills of subadults were restricted to vocal exchanges with other subadults. Continuous vocal recordings were collected of the vocalizations of the 14 members of the study troop. A cumulative 33.7 h of continuous samples and 1,892 sonagrams were analyzed. In addition to vocalizations clearly associated with alarm, distress, or agonistic contexts, two distinct call types were identified, trills and huhs. Age-sex classes differed in the rate at which both types of calls were produced in different spatial positions within the troop. Adult females and males produced higher rates of trills when in the leading edge compared to all other spatial positions in a traveling troop. Trills at the edge of a stationary troop represented 36 “successful” and 3 “unsuccessful” start attempts; the troop usually moved in the trajectory predicted by a trilling adult's location on the troop periphery within 10 min of the initiation of trilling. Adults also altered the trajectory of traveling troops by trilling at the side and back of the troop (10 “successful” and 4 “unsuccessful” attempts). Huh vocalizations were most predictably produced when a capuchin is in a dense fruit patch. These results emphasize the role vocalizations serve in the coordination and trajectory of group movement in nonhuman primates, especially those populations that are arboreal or in which visual contact is otherwise impeded. © 1993 Wiley-Liss, Inc.

1 This article is a US Government work and, as such, is in the public domain in the United States of America.

     

Peering in mature, captive bonobos (Pan paniscus)

“Peering”—close-proximity staring at the mouth of another—was observed in ten (three males and seven females) mature (at least 7 years old) bonobos (Pan paniscus) living in three social groups at the San Diego Zoo and Wild Animal Park. Instantaneous scan samples, taken at 2-min intervals, over a three-and-a-half year period, yielded 617 observations of peering (1.4 per observation hour). Food was exchanged in only 15 of these scans. Peering was most often performed by younger animals and was primarily directed toward older females (“matrons”). In a given dyad, the animal more likely to peer at the other was also more like to both peer and be peered at if they frequently groomed and infrequently displayed aggression at a given female. An adolescent male showed the highest frequency of peering when living with two older females, but dropped to adult male levels when later housed with two younger (albeit mature) females. A reversal in which animal was more likely to peer, follow, and groom occurred in one female dyad, after the birth of the younger animal's first infant. After a similar birth in the other group, no such changes were observed. We discuss how these and related findings, in conjunction with what is known of the social structure of this species, suggest that one possible function of peering in bonobos may be as a signal acknowledging female status.     

Association and social interactions between strangers and residents in bonobos (Pan paniscus)

This study reports on close spatial association and repeated behavioural interactions between two strange adult male bonobos with residents of another community. Over a period of 12 months one of the two males developed friendly social relations to some of the females and other residents, which were indistinguishable from those existing between co-residents. Aggression by resident males against the strangers decreased but the former remained intolerant. The strange males appeared at a time when the number of adult resident males was lower as in the years before and when the adult sex ratio (number of adult females per male) was higher as in the years before. Using definitions from studies on dispersal patterns of male gorillas (Harcourt, 1978) and female bonobos (Furuichi, 1989) the spatial association between the two strange males and residents could be described as male transfer.     

Population Dynamics of Wild Bonobos (Pan paniscus) at Wamba

We analyzed population dynamics and birth seasonality of wild bonobos at Wamba, Democratic Republic of the Congo, based on 20 years of observations (1976–1996). Wamba Bonobo infant mortality is much lower than that reported for chimpanzees. This seemes to be related to several socioecological characteristics of bonobos: the use of abundant fruit and herbaceous foods, larger food patch size, female feeding priority, and the absence of infanticide. The mean interval between live births of 4.8 years is shorter than those reported for chimpanzees, and some females simultaneously carried and nursed two successive offspring. Mother–offspring conflicts, such as refusal of suckling attempts and interference with mothers' copulation, which are common in chimpanzees, are rare in Wamba bonobos. A birth peak seems to occur during the light rainy season from March to May, just after the season with the least rainfall. This timing of births is similar to those reported for chimpanzee populations, and might benefit both mother and offspring by maximizing the amount of time before the next dry season.     

Seasonality and Socioecology: The Importance of Variation in Fruit Abundance to Bonobo Sociality

The assumption that nonseasonal, evergreen, rain forests contain more continuously available food resources than seasonal rain forests is fundamental to comparisons made between the socioecology of the male-bonded Pan troglodytes and the female-based social system of the Pan paniscus. Chimpanzee females may be less social due to the high costs of feeding competition, whereas in the more food-rich central African rain forests such as the Lomako forest, female bonobos can associate and socially bond. The Lomako Forest experiences two wet and two dry seasons a year. Data on fruit abundance and sociality show that despite monthly variation in fruit availability, there was no consistent seasonal variation in fruit abundance or dietary breadth. Bonobo use of nonfig fruits, figs, THV, and leaves did not follow seasonal patterns. Leaves and THV may act as complementary sources of plant protein and their use was inversely correlated. Monthly variation in fruit abundance was associated with a significant decrease in the number of males in a party but not in the number of females. Focal males were frequently solitary during 1 of the 3 months with the smallest party sizes. In contrast, females remained social with each other throughout the year. Therefore, seaonality at Lomako appeared to be less marked than at comparable chimpanzee sites, such that the variation in fruit abundance did not fall below a level that prohibits female sociality.     

The responses of bonobos (Pan paniscus) to their mirror images: Evidence of selfrecognition

This research examined the responses of bonobos (Pan paniscus) to their mirror images. Nine bonobos were presented alternately with the reflective and non-reflective sides of a mirror. The apes exhibited considerable interest in the mirror, and immature animals exhibited higher frequencies of contingent action and inactive looking than did adults. four animals used the mirror to inspect parts of their bodies that were otherwise not visible to them, indicating that bonobos are capable of self-recognition.