Innovation in wild chimpanzees (Pan troglodytes)

Innovations of behavior have major implications for the concept of culture in animals. Innovation has rarefy been documented in wild animal populations. The chimpanzees of the Ta? National Park, C?te d’Ivoire, spontaneously included new patterns of behavior in their repertoire during our study. Some innovations were incorporated into a minority of the group members’ repertoire, whereas others, such as building day nests in trees and on the ground more frequently, became general behaviors. Similarly, new contexts for use of leaf- clipping behavior and novel ways of feeding on some leaves appeared and rapidly became part of the behavior of most group members. The environmental parameters were too stable to explain these new forms of behavior, as most of them took place within 1 month during the same dry season. In a similar way, leaf- grooming acquired a new function in Gombe chimpanzees. A process similar to social conventions could explain the emergence of these new functions for an existing behavior in a way rather similar to human “fashion. ” Three major characteristics of human culture — the absence of individual variations in the performance of the behavior, the general use of the behavior by group members, and the ability to modify the function of a behavior — characterize leaf-clipping in Ta? chimpanzees and leaf- grooming in Gombe chimpanzees.     

Can the Patterns of Sexual Swelling Cycles in Female Taï Chimpanzees be Explained by the Cost-of-Sexual-Attraction Hypothesis?

Chimpanzee females of East and West African populations differ in the average number of cycles per interbirth interval. Whereas females in Gombe, Mahale, and Kibale (eastern chimpanzees, Pan troglodytes schweinfurthii) average <9 cycles before they conceive, females at Taï (western chimpanzees; Pan troglodytes verus) average 29 cycles. We examined data from 2 different study groups (North and South) at Taï. By showing that Taï females interrupt cyclic activity for, on average, 7.4 mo between the end of the postpartum amenorrhea period (PPA) and the subsequent conception, and by calculating the number of cycles to conception based on a probability distribution of cycles over the interbirth interval, we show that Taï females average 19.4 cycles (North Group) and 11.7 cycles (South Group) to conception; therefore, the earlier calculation of 29 cycles was an overestimate. Further, at Taï young parous females have a significantly shorter PPA than those of older females, but the number of cycles to conception does not differ significantly between primiparous and nulliparous females. Some of our results are therefore not in line with the predictions of the cost-of-sexual-attraction hypothesis, which proposes that the factor ultimately responsible for the intensity of female sexual attractiveness is female intragroup scramble competition. We discuss an alternative hypothesis—the social passport hypothesis—which is compatible with the results of the study.     

Ant-dipping among the chimpanzees of Bossou,Guinea, and some comparisons with other sites

We present a detailed study of ant‐dipping among the wild chimpanzees (Pan troglodytes verus) of Bossou, in southeastern Guinea, West Africa. Observations suggest a strong influence of prey (Dorylusspp.) characteristics, including aggressiveness and/or gregariousness, on tool length and technique employed by the chimpanzees. Bossou chimpanzees exhibit two ant‐dipping techniques: 1) direct mouthing, and 2) pull‐through. In addition, they were observed dipping for several species of Dorylus ants, classed into two categories: Red and Black. Tool length was longer when dipping in higher‐risk contexts, i.e., at the ants' nest site or on Black ants. The pull‐through technique was almost exclusively associated with dipping at the nest site. This latter technique was associated with tools over 50cm long, whereas direct mouthing was the only technique observed with tools <50cm long. Our experimental findings, together with our observations on the behavior of the chimpanzees, suggest that at the nest, the pull‐through technique was a more efficient technique than direct mouthing. We review our results in the context of ant‐dipping observed at two other long‐term chimpanzee study sites, i.e., Gombe (Tanzania) and Taï (Côte d'Ivoire), where differences in tool length, technique used, and focal Dorylus ant species have been reported. Finally, we urge similar detailed studies of this tool‐use behavior in both Gombe and Taï to shed further light upon our results and their implications. Am. J. Primatol. 58:133–148, 2002. © 2002 Wiley‐Liss, Inc.     

Noninvasive paternity assignment in Gombe chimpanzees

The relative success of chimpanzee male mating strategies, the role of male dominance rank and the success of inbreeding avoidance behaviour can only be assessed when paternities are known. We report the probable paternities of 14 chimpanzees included in a long‐term behavioural study of chimpanzees (Pan troglodytes schweinfurthii) at Gombe National Park, Tanzania. DNA samples were collected noninvasively from shed hair and faeces and genotyped using 13–16 microsatellite loci characterized in humans. All 14 offspring could be assigned to fathers within the community. While there is a positive relationship between male rank and reproductive success, we demonstrate that a range of male mating strategies (possessiveness, opportunistic mating and consortships) can lead to paternity across all male ranks. Several adult females were at risk of breeding with close male relatives. Most successfully avoided close inbreeding but in one case a high‐ranking male in the community mated with his mother and produced an offspring. In contrast to recent data on chimpanzees (P. t. verus) from the Taï forest, Côte d’Ivoire, no evidence of extra‐group paternity was observed in our study. Reanalysis of Taï data using a likelihood approach casts doubt on the occurrence of extra‐group paternity in that community as well.     

Morphological variation in adult chimpanzees (Pan troglodytes verus) of the Taï National Park, Côte d'Ivoire

Twenty five adult chimpanzee skeletons (Pan troglodytes verus) of known age and sex (15 females, 10 males) from a long‐term study site in Taï National Park, Cote d'Ivoire present new data on variation. These skeletons provide a rare opportunity to measure the cranium and postcranium from the same individuals. We compare measurements and indices of the Taï sample with those of relatively complete Pan troglodytes schweinfurthii skeletons from Gombe National Park, Tanzania. Measurements of Pan paniscus are included as an outside comparison. The Taï and Gombe samples are analyzed by sex; combined sex samples are compared between the two groups, and the two sexes to each other. Taï females and males do not differ in most long bone lengths or in pelvic dimensions, but do differ significantly in cranial capacity, facial measurements, clavicle length, scapular breadth, and femur length. Gombe females and males differ significantly in some facial measurements and in scapular breadth. In combined sex samples, Taï individuals have lower cranial capacity, longer palate and mandible, and greater dimensions in the trunk and limb lengths. Taï females account for most of the variation; males differ from each other only in greater length of humerus and femur. The Taï skeletons provide new data for assessing individual variation and sexual dimorphism within and between populations and species. The combination of cranial and postcranial data provides a clearer picture of chimpanzee intraspecific and interspecific variation than can be gained from either data set alone. Am J Phys Anthropol, 2008. © 2007 Wiley‐Liss, Inc.     

How feeding competition determines female chimpanzee gregariousness and ranging in the Taï National Park, Côte d'Ivoire

Socioecological theory suggests that feeding competition shapes female social relationships. Chimpanzees (Pan troglodytes) live in fission–fusion societies that allow them to react flexibly to increased feeding competition by forming smaller foraging parties when food is scarce. In chimpanzees at Gombe and Kibale, female dominance rank can crucially influence feeding competition and reproductive success as high‐ranking females monopolize core areas of relatively high quality, are more gregarious, and have higher body mass and reproductive success than low‐ranking females. Chimpanzee females in Taï National Park do not monopolize core areas; they use the entire territory as do the males of their community and are highly gregarious. Although female chimpanzees in Taï generally exhibit a linear dominance hierarchy benefits of high rank are currently not well understood. We used a multivariate analysis of long‐term data from two Taï chimpanzee communities to test whether high‐ranking females (1) increase gregariousness and (2) minimize their travel costs. We found that high‐ranking females were more gregarious than low‐rankers only when food was scarce. During periods of food scarcity, high rank allowed females to enjoy benefits of gregariousness, while low‐ranking females strongly decreased their gregariousness. High‐ranking females traveled more than low‐ranking females, suggesting that low‐rankers might follow a strategy to minimize energy expenditure. Our results suggest that, in contrast to other chimpanzee populations and depending on the prevailing ecological conditions, female chimpanzees at Taï respond differently to varying levels of feeding competition. Care needs to be taken before generalizing results found in any one chimpanzee population to the species level. Am. J. Primatol. 73:305–313, 2011. © 2010 Wiley‐Liss, Inc.     

Role of Nonbehavioral Factors in Adjusting Long Bone Diaphyseal Structure in Free-ranging <Emphasis Type="Italic">Pan troglodytes</Emphasis>

Limb bones deform during locomotion and can resist the deformations by adjusting their shapes. For example, a tubular-shaped diaphysis best resists variably-oriented deformations. As behavioral profiles change during adulthood, patterns of bone deformation may exhibit age trends. Habitat characteristics, e.g., annual rainfall, tree density, and elevation changes, may influence bone deformations by eliciting individual components of behavioral repertoires and suppressing others, or by influencing movements during particular components. Habituated chimpanzee communities provide a unique opportunity to examine these factors because of the availability of morphological data and behavioral observations from known-age individuals inhabiting natural habitats. We evaluated adult femora and humeri of 18 female and 10 male free-ranging chimpanzees (Pan troglodytes) from communities in Gombe (Tanzania), Mahale Mountains (Tanzania), and Taï Forest (Côte d’Ivoire) National Parks. We compare cross sections at several locations (35%, 50%, 65% diaphyseal lengths). Community comparisons highlight different diaphyseal shapes of Taï females relative to Mahale and Gombe females, particularly in humeral diaphyses. Age trends in diaphyseal shapes are consistent with reduced activity levels in general, not only reduced arboreal activity. Age-related bone loss is apparent among community females, but is less striking among males. Community trends in diaphyseal shape are qualitatively consistent with ranked annual rainfall at localities, tree density, and elevation change or ruggedness of terrain. Habitat characteristics may contribute to variation in diaphyseal shape among chimpanzee communities, much like among modern human groups, but verification awaits further rigorous experimental and comparative analyses.     

Symbolic communication in wild chimpanzees?

The language abilities of captive chimpanzees give rise to the question of the existence and use of similar capabilities in wild chimpanzees. In Taï forest, wild chimpanzees seem to use drumming on buttressed trees to convey information an changes of travel direction, resting periods or both information combined. This communication system is iconic and relies on some arbitrariness. Emergence of symbol-like communication in wild chimpanzees seems mainly dependent on a low visibility environment, a high predation pressure and a large group of males.     

Buttress drumming by wild chimpanzees: Temporal patterning, phrase integration into loud calls, and preliminary evidence for individual distinctiveness

Wild chimpanzees (Pan troglodytes) generate low-frequency sounds that are audible to humans from a distance of at least 1 km away by hitting the buttresses of trees with their hands and feet. This buttress drumming occurs in discrete bouts of rapidly delivered beats that usually accompany “pant hoots,” the species-specific long-distance vocalization. Individual differences in male chimpanzee (P.t. verus) drumming were investigated during a 6-month field study in the Taï National Park, Ivory Coast. Analysis of drumming bouts recorded from six adult males revealed significant differences between individuals in three acoustic features: (1) mean duration of inter-beat interval; (2) mean number of beats per bout; and (3) mean bout duration. Preliminary analysis indicated that individuals differ in their tendency to deliver drum beats in temporally close pairs separated by longer interbeat intervals. Qualitative examination also suggested that individuals may differ in the temporal integration of drumming into the pant hoot vocalization. These results suggest that there may be acoustic cues available for chimpanzees to recognize unseen males by their drumming performances alone. Drumming by Taï chimpanzees was also compared to drumming by chimpanzees (P.t. schweinfurthii) from the Kanyawara study group in Kibale National Park. Uganda. The Kanyawara chimpanzees appeared to drum more often without vocalizing than did the Taï chimpanzees. When they did drum and vocalize together, the Kanyawara chimpanzees appeared to integrate their drumming later into the associated pant hoots than did the Taï chimpanzees. These results suggest the possibility that interpopulation variation exists in chimpanzee buttress drumming.     

Stable nitrogen isotope analysis of dentine serial sections elucidate sex differences in weaning patterns of wild chimpanzees (Pan troglodytes)

Offspring provisioning is one of the most energetically demanding aspects of reproduction for female mammals. Variation in lactation length and weaning strategies between chimpanzees (Pan troglodytes), our closest living relative, and modern human societies have been reported. When and why these changes occurred is frequently debated. Our study used stable nitrogen isotope data of tooth root dentine from wild Western chimpanzees (Pan troglodytes verus) in Taï National Park, Côte d'Ivoire, to quantify weaning in these chimpanzees and explore if infant sex plays a role in maternal investment. We analyzed serial sections of deciduous lateral incisor root dentine from four Taï chimpanzees to establish the δ¹⁵N signal of nursing infants; we then analyzed serial sections of first permanent mandibular molar root dentine from 12 Taï chimpanzees to provide quantitative δ¹⁵N data on weaning in this population. Up to 2 years of age both sexes exhibited dentine δ¹⁵N values ≈2–3‰ higher than adult female Taï chimpanzees, consistent with a nursing signal. Thereafter a steady decrease in δ¹⁵N values consistent with the onset, and progression, of weaning, was visible. Sex differences were also evident, where male δ¹⁵N values decreased at a significantly slower rate compared to females. Confirmation of sex differences in maternal investment among Taï chimpanzees, demonstrates the viability of using isotope analysis to investigate weaning in non‐human primates. Additionally, assuming that behaviors observed in the Taï chimpanzees are illustrative of the ancestral pattern, our results provide a platform to enable the trajectory of weaning in human evolution to be further explored. Am J Phys Anthropol 153:635–642, 2014. © 2013 Wiley Periodicals, Inc.     

Handedness in wild chimpanzees

The debate over nonhuman primate precursors to human handedness is unsettled mainly due to lack of data, particularly on apes. Handedness in wild chimpanzees at the Taï National Park Côte d'Ivoire, has been monitored in four tasks. For the simple unimanual ones, reaching and grooming, adults use both hands equally (ambidextrous), while for the more complex unimanual wadge-dipping and the complex bimanual nut-cracking, adults are highly lateralized. These results support the hypothesis that lateralization increases with the complexity of the task. The lateralization is constant for years for each task but may vary in an individual with respect to different tasks. For nutcracking females are more lateralized than males. The ontogeny of handedness for nut-cracking shows many variations in the tendency to use one hand and in the side preferred, until at about 10 years of age, the individual achieves her adult handedness. No population bias toward one side exists in Taï chimpanzees. No heritability of handedness between mother and offspring was observed. Human and chimpanzees handedness are compared.     

Sexual Differences in Chimpanzee Sociality

Scientists usually attribute sexual differences in sociality to sex-specific dispersal patterns and the availability of kin within the social group. In most primates, the dispersing sex, which has fewer kin around, is the less social sex. Chimpanzees fit well into the pattern, with highly social philopatric males and generally solitary dispersing females. However, researchers in West Africa have long suggested that female chimpanzees can be highly social. We investigated whether chimpanzees in the Taï Forest (Côte d’Ivoire) exhibit the expected sexual differences in 3 social parameters: dyadic association, party composition, and grooming interactions. Though we found a significant sexual difference in each of the 3 parameters, with males being more social than females, the actual values do not reveal striking differences between the sexes and do not support the notion of female chimpanzees as asocial: females had dyadic association indices comparable to mixed-sex dyads, spent ca. 82% of their time together with other adult chimpanzees, and had a comparable number of grooming partners. Further, female associations can be among the strongest bonds within the community, indicating that both sexes can have strongly favored association partners. The findings are in contrast to reports on East African chimpanzees, the females of which are mainly solitary and rarely interact with other females. Our results suggest that researchers cannot generally regard chimpanzee females as asocial and need to redefine models deriving patterns of sociality from dispersal patterns to integrate the possibility of high female sociality in male philopatric systems.     

Anxiety-related behavior of orphan chimpanzees (Pan troglodytes schweinfurthii) at Gombe National Park, Tanzania

This study examined the anxiety levels and social interactions of two orphan and four mother-reared adolescent chimpanzees (Pan troglodytes schweinfurthii) in the Kasekela community at Gombe National Park, Tanzania. We used focal sampling in the field at Gombe to observe these adolescent individuals. Their social interactions and anxious behavior, measured as rough scratching, were recorded. The two orphans differed from others of a similar age by exhibiting higher levels of anxiety and lower levels of play. These results suggest that a mother’s absence, even in naturalistic conditions in which other members of the community are available to the orphan, may have long-lasting impact on an adolescent’s anxiety and its ability to engage in complex social interactions, such as play.     

Hunting behavior of wild chimpanzees in the Taï National Park

Hunting is often considered one of the major behaviors that shaped early hominids' evolution, along with the shift toward a drier and more open habitat. We suggest that a precise comparison of the hunting behavior of a species closely related to man might help us understand which aspects of hunting could be affected by environmental conditions. The hunting behavior of wild chimpanzees is discussed, and new observations on a population living in the tropical rain forest of the Taï National Park, Ivory Coast, are presented. Some of the forest chimpanzees' hunting performances are similar to those of savanna-woodlands populations; others are different. Forest chimpanzees have a more specialized prey image, intentionally search for more adult prey, and hunt in larger groups and with a more elaborate cooperative level than savanna-woodlands chimpanzees. In addition, forest chimpanzees tend to share meat more actively and more frequently. These findings are related to some theories on aspects of hunting behavior in early hominids and discussed in order to understand some factors influencing the hunting behavior of wild chimpanzees. Finally, the hunting behavior of primates is compared with that of social carnivores.     

Social grooming and play behavior of a captive group of chimpanzees

Aspects of the social grooming and play behavior of a group of six adolescent and young adult chimpanzees are contrasted and compared. Eleven months’ data indicate that older chimpanzees groomed more and played less than younger individuals. This transition period occurred earlier for females than males. Grooming behavior appeared to vary with reproductive state. A positive correlation was found between the estrous condition of cycling females and the amount of grooming that they received from the males. A mother of a young infant received particularly high levels of grooming from the other group members. Less variation among individuals was found for frequencies of play as compared to grooming. Play dropped following the death of one individual and was entirely inhibited for three weeks following the group’s transfer to a new environment and the reintroduction of a former group member. Comparison to a free-ranging population indicates important differences in both frequencies and general patterns of play and grooming.     

Passive acoustic monitoring reveals group ranging and territory use: a case study of wild chimpanzees (<Emphasis Type="Italic">Pan troglodytes</Emphasis>)

Background

Assessing the range and territories of wild mammals traditionally requires years of data collection and often involves directly following individuals or using tracking devices. Indirect and non-invasive methods of monitoring wildlife have therefore emerged as attractive alternatives due to their ability to collect data at large spatiotemporal scales using standardized remote sensing technologies. Here, we investigate the use of two novel passive acoustic monitoring (PAM) systems used to capture long-distance sounds produced by the same species, wild chimpanzees (Pan troglodytes), living in two different habitats: forest (Taï, Côte d’Ivoire) and savanna-woodland (Issa valley, Tanzania).

Results

Using data collected independently at two field sites, we show that detections of chimpanzee sounds on autonomous recording devices were predicted by direct and indirect indices of chimpanzee presence. At Taï, the number of chimpanzee buttress drums detected on recording devices was positively influenced by the number of hours chimpanzees were seen ranging within a 1 km radius of a device. We observed a similar but weaker relationship within a 500 m radius. At Issa, the number of indirect chimpanzee observations positively predicted detections of chimpanzee loud calls on a recording device within a 500 m but not a 1 km radius. Moreover, using just seven months of PAM data, we could locate two known chimpanzee communities in Taï and observed monthly spatial variation in the center of activity for each group.

Conclusions

Our work shows PAM is a promising new tool for gathering information about the ranging behavior and habitat use of chimpanzees and can be easily adopted for other large territorial mammals, provided they produce long-distance acoustic signals that can be captured by autonomous recording devices (e.g., lions and wolves). With this study we hope to promote more interdisciplinary research in PAM to help overcome its challenges, particularly in data processing, to improve its wider application.

     

Alpha male chimpanzee grooming patterns: implications for dominance “style”

In social primates, individuals use various tactics to compete for dominance rank. Grooming, displays and contact aggression are common components of a male chimpanzee's dominance repertoire. The optimal combination of these behaviors is likely to differ among males with individuals exhibiting a dominance “style” that reflects their tendency to use cooperative and/or agonistic dominance tactics. Here, we examine the grooming behavior of three alpha male chimpanzees at Gombe National Park, Tanzania. We found that (1) these males differed significantly in their tendency to groom with other males; (2) each male's grooming patterns remained consistent before, during and after his tenure as alpha, and (3) the three males tended to groom with high‐ middle‐ and low‐ranking partners equally. We suggest that body mass may be one possible determinant of differences in grooming behavior. The largest male exhibited the lowest overall grooming rates, whereas the smallest male spent the most time grooming others. This is probably because large males are more effective at physically intimidating subordinates. To achieve alpha status, a small male may need to compensate for reduced size by investing more time and energy in grooming, thereby ensuring coalitionary support from others. Rates of contact aggression and charging displays conformed to this prediction, suggesting that each male exhibited a different dominance “style.” Am. J. Primatol. 71:136–144, 2009. © 2008 Wiley‐Liss, Inc.     

Census and distribution of chimpanzees in Côte D’Ivoire

Most methods of estimating chimpanzee population densities rely on nest counts. We tested the most frequently used techniques on a known chimpanzee community living in the rainforest of the Taï National Park, Côte d’Ivoire. The best density estimates are given by counts that assume groups of nests to be distributed randomly and that use the mean group size for homogenous habitat but the median for heterogenous habitats. Correction for real forest cover within the region should be made because chimpanzees make nests only in forested regions. This method gave the exact chimpanzee density for the Taï population, i.e. 1.7 nest builders/km². For the nationwide survey, we first estimated the chimpanzee density for different types of habitat (e.g. intact primary forest: 1.64 chimpanzees/km²; degraded forests: 0.4 chimpanzees/km²; human encroached forests and mosaic habitats: 0.09 chimpanzees/km²). Second, we estimated the total forest cover of the country with satellite pictures. This gave an estimated chimpanzee population in Côte d’Ivoire of about 11,676 ± 1,168 individuals, which equals the number of spectators at a soccer game in an average European town. Sadly, only three National Parks may have chimpanzee populations large enough to be viable, whereas the rest are scattered and isolated small populations that are already threatened in their survival.