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Wild chimpanzees (Pan troglodytes) are caught in snares set for other animals and sometimes injure or lose body parts. Snaring can compromise the health, growth, survival, and behavior of chimpanzees and, thus, represents a threat for the conservation of this endangered species. During a long-term study of chimpanzees at Ngogo in Kibale National Park, Uganda, we started a project to remove snares in and around their territory. We compared the number of times chimpanzees were snared during the 12.75 years after the start of this project with the number of times individuals were snared during the previous 14 years. Only one chimpanzee was snared after we began removing snares compared with 12 individuals caught during the period before. This represents a clear reduction in the risk created by snaring at this site and suggests that removing snares can be employed to protect chimpanzees. [ABSTRACT FROM AUTHOR]

Many anthropogenic-driven changes, such as hunting, have clear and immediate negative impacts on wild primate populations, but others, like climate change, may take generations to become evident. Thus, informed conservation plans will require decades of population monitoring. Here, we expand the duration of monitoring of the diurnal primates at Ngogo in Kibale National Park, Uganda, from 32.9 to 47 years. Over the 3531 censuses that covered 15,340 km, we encountered 2767 primate groups. Correlation analyses using blocks of 25 census walks indicate that encounters with groups of black and white colobus, blue monkeys, and baboons neither increased nor decreased significantly over time, while encounters with groups of redtail monkeys and chimpanzees marginally increased. Encounters with mangabeys and L'Hoesti monkeys increased significantly, while red colobus encounters dramatically decreased. Detailed studies of specific groups at Ngogo document changes in abundances that were not always well represented in the censuses because these groups expanded into areas away from the transect, such as nearby regenerating forest. For example, the chimpanzee population increased steadily over the last 2 + decades but this increase is not revealed by our census data because the chimpanzees expanded, mainly to the west of the transect. This highlights that extrapolating population trends to large areas based on censuses at single locations should be done with extreme caution, as forests change over time and space, and primates adapt to these changes in several ways. [ABSTRACT FROM AUTHOR]

For energetically limited organisms, life‐history theory predicts trade‐offs between reproductive effort and somatic maintenance. This is especially true of female mammals, for whom reproduction presents multifarious energetic and physiological demands.Here, we examine longitudinal changes in the gut virome (viral community) with respect to reproductive status in wild mature female chimpanzees Pan troglodytes schweinfurthii from two communities, Kanyawara and Ngogo, in Kibale National Park, Uganda.We used metagenomic methods to characterize viromes of individual chimpanzees while they were cycling, pregnant and lactating.Females from Kanyawara, whose territory abuts the park's boundary, had higher viral richness and loads (relative quantity of viral sequences) than females from Ngogo, whose territory is more energetically rich and located farther from large human settlements. Viral richness (total number of distinct viruses per sample) was higher when females were lactating than when cycling or pregnant. In pregnant females, viral richness increased with estimated day of gestation. Richness did not vary with age, in contrast to prior research showing increased viral abundance in older males from these same communities.Our results provide evidence of short‐term physiological trade‐offs between reproduction and infection, which are often hypothesized to constrain health in long‐lived species. [ABSTRACT FROM AUTHOR]

The original article can be found online at https://doi.org/10.1007/s10329-023-01087-4.Correction: Primates (2023) 64:609–620https://doi.org/10.1007/s10329-023-01087-4In the original version of this article Fig. 2 appeared with an error; the figure should have appeared as shown below.GraphPublisher's NoteSpringer Nature remains neutral with regard to jurisdictional claims in published maps and institutional affiliations.By Colin A. Chapman; Samuel Angedakin; Thomas M. Butynski; Jan F. Gogarten; John C. Mitani and Thomas T. StruhsakerReported by Author; Author; Author; Author; Author; Author [Extracted from the article]

Objectives: This study examines the sensitivity of the development of the dorsal metacarpal ridge (DMR) to the frequency of knuckle‐walking during ontogeny in chimpanzees, and compares the prevalence and prominence of the DMR in hominoids and baboons. Materials and Methods: We characterized the type and frequency of quadrupedalism of wild chimpanzees of different ages from the Ngogo community, Uganda. Using museum specimens, we quantified third metacarpal DMR angle and height in Pan individuals of different developmental stages, and in adult Gorilla, Pongo, Hylobates, and Papio. Results: When terrestrial, all quadrupedalism consisted of knuckle‐walking, although older individuals knuckle‐walked more than younger. The total amount of quadrupedalism engaged in by different age groups while arboreal was consistent; however, knuckle usage while arboreally quadrupedal increased with age. The DMR ridge did not emerge in Pan until older infancy, after the onset of knuckle‐walking, and reached prominence by the juvenile stage. The DMR angle increased significantly after juvenility but relative height did not. Adult Pan and Gorilla have more prevalent and prominent DMRs than Papio; the DMR was rare in Pongo and absent in Hylobates. Discussion: The timing of development of the DMR in chimpanzees supports it as a knuckle walking character, as does its occurrence in gorillas, and virtual absence in suspensory orangutans and gibbons. The DMR in baboons differs in appearance and frequency from DMRs in African apes, and so differentiates knuckle‐walking from digitigrady in catarrhines. The relationship between the DMR and loading of a vertical hand while terrestrial deserves further study. [ABSTRACT FROM AUTHOR]

Viral infection is a major cause of ill health in wild chimpanzees (Pan troglodytes), but most evidence to date has come from conspicuous disease outbreaks with high morbidity and mortality. To examine the relationship between viral infection and ill health during periods not associated with disease outbreaks, we conducted a longitudinal study of wild eastern chimpanzees (P. t. schweinfurthii) in the Kanyawara and Ngogo communities of Kibale National Park, Uganda. We collected standardized, observational health data for 4 years and then used metagenomics to characterize gastrointestinal viromes (i.e., all viruses recovered from fecal samples) in individual chimpanzees before and during episodes of clinical disease. We restricted our analyses to viruses thought to infect mammals or primarily associated with mammals, discarding viruses associated with nonmammalian hosts. We found 18 viruses (nine of which were previously identified in this population) from at least five viral families. Viral richness (number of viruses per sample) did not vary by health status. By contrast, total viral load (normalized proportion of sequences mapping to viruses) was significantly higher in ill individuals compared with healthy individuals. Furthermore, when ill, Kanyawara chimpanzees exhibited higher viral loads than Ngogo chimpanzees, and males, but not females, exhibited higher infection rates with certain viruses and higher total viral loads as they aged. Post‐hoc analyses, including the use of a machine‐learning classification method, indicated that one virus, salivirus (Picornaviridae), was the main contributor to health‐related and community‐level variation in viral loads. Another virus, chimpanzee stool‐associated virus (chisavirus; unclassified Picornavirales), was associated with ill health at Ngogo but not at Kanyawara. Chisavirus, chimpanzee adenovirus (Adenoviridae), and bufavirus (Parvoviridae) were also associated with increased age in males. Associations with sex and age are consistent with the hypothesis that nonlethal viral infections cumulatively reflect or contribute to senescence in long‐lived species such as chimpanzees. Research Highlights: Some viruses of wild primates are relatively benign but may be useful as biomarkers of physiology and health.We used metagenomic methods to characterize viral communities in feces of wild chimpanzees in two long‐term study communities (Kanyawara and Ngogo) in Kibale National Park, Uganda.Chimpanzees exhibited higher viral loads (normalized abundances of viral genetic sequences) when they were ill than when those same individuals were previously healthy.When ill, Kanyawara chimpanzees exhibited higher viral loads than Ngogo chimpanzees, and males, but not females, exhibited higher viral loads as they aged. [ABSTRACT FROM AUTHOR]

I have spent over 40 years studying the behavior of our closest living relatives, the apes. In this paper, I review my research on the spacing, mating, and vocal behavior of gibbons and orangutans (Pongo pygmaeus) and the vocal and social behavior of chimpanzees (Pan troglodytes). I devote special attention to results derived from a 25‐year‐long study of a remarkable and extraordinarily large group of chimpanzees that has recently fissioned at Ngogo in Kibale National Park, Uganda. I conclude with some advice for the next generation of field primatologists. Research Highlights: I review the results of my field research on gibbons, orangutans, and chimpanzees.My findings provide insights into the spacing, mating, and vocal behavior of gibbons and orangutans and the vocal and social behavior of chimpanzees. [ABSTRACT FROM AUTHOR]

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. [ABSTRACT FROM AUTHOR]

Respiratory pathogens are expected to spread through social contacts, but outbreaks often occur quickly and unpredictably, making it challenging to simultaneously record social contact and disease incidence data, especially in wildlife. Thus, the role of social contacts in the spread of infectious disease is often treated as an assumption in disease simulation studies, and few studies have empirically demonstrated how pathogens spread through social networks. In July–August 2015, an outbreak of respiratory disease was observed in a wild chimpanzee community in Kibale National Park, Uganda, during an ongoing behavioral study of male chimpanzees, offering a rare opportunity to evaluate how social behavior affects individual exposure to socially transmissible diseases. From May to August 2015, we identified adult and adolescent male chimpanzees displaying coughs and rhinorrhea and recorded 5-m proximity data on males (N = 40). Using the network k-test, we found significant relationships between male network connectivity and the distribution of cases within the network, supporting the importance of short-distance contacts for the spread of the respiratory outbreak. Additionally, chimpanzees central to the network were more likely to display clinical signs than those with fewer connections. Although our analyses were limited to male chimpanzees, these findings underscore the value of social connectivity data in predicting disease outcomes and elucidate a potential evolutionary cost of being social. [ABSTRACT FROM AUTHOR]

In humans, senescence increases susceptibility to viral infection. However, comparative data on viral infection in free-living non-human primates—even in our closest living relatives, chimpanzees and bonobos (Pan troglodytes and P. paniscus)—are relatively scarce, thereby constraining an evolutionary understanding of age-related patterns of viral infection. We investigated a population of wild eastern chimpanzees (P. t. schweinfurthii), using metagenomics to characterize viromes (full viral communities) in the faeces of 42 sexually mature chimpanzees (22 males, 20 females) from the Kanyawara and Ngogo communities of Kibale National Park, Uganda. We identified 12 viruses from at least four viral families possessing genomes of both single-stranded RNA and single-stranded DNA. Faecal viromes of both sexes varied with chimpanzee age, but viral richness increased with age only in males. This effect was largely due to three viruses, salivirus, porprismacovirus and chimpanzee stool-associated RNA virus (chisavirus), which occurred most frequently in samples from older males. This finding is consistent with the hypothesis that selection on males for early-life reproduction compromises investment in somatic maintenance, which has delayed consequences for health later in life, in this case reflected in viral infection and/or shedding. Faecal viromes are therefore useful for studying processes related to the divergent reproductive strategies of males and females, ageing, and sex differences in longevity. This article is part of the theme issue 'Evolution of the primate ageing process'. [ABSTRACT FROM AUTHOR]

Fruit production in tropical forests varies considerably in space and time, with important implications for frugivorous consumers. Characterizing temporal variation in forest productivity is thus critical for understanding adaptations of tropical forest frugivores, yet long‐term phenology data from the tropics, in particular from African forests, are still scarce. Similarly, as the abiotic factors driving phenology in the tropics are predicted to change with a warming climate, studies documenting the relationship between climatic variables and fruit production are increasingly important. Here, we present data from 19 years of monitoring the phenology of 20 tree species at Ngogo in Kibale National Park, Uganda. Our aims were to characterize short‐ and long‐term trends in productivity and to understand the abiotic factors driving temporal variability in fruit production. Short‐term (month‐to‐month) variability in fruiting was relatively low at Ngogo, and overall fruit production increased significantly through the first half of the study. Among the abiotic variables, we expected to influence phenology patterns (including rainfall, solar irradiance, and average temperature), only average temperature was a significant predictor of monthly fruit production. We discuss these findings as they relate to the resource base of the frugivorous vertebrate community inhabiting Ngogo. [ABSTRACT FROM AUTHOR]

When their mothers die, chimpanzees often adopt younger vulnerable siblings who survive with their care. This phenomenon has been widely reported, but few studies provide details regarding how sibling relationships change immediately following the deaths of their mothers. A disease outbreak that killed several females at Ngogo in Kibale National Park, Uganda, furnished an opportunity to document how maternal death influenced the social relationships of siblings. We describe social interactions between four adolescent and young adult males and their younger immature maternal siblings 9 months before and 8 months after their mothers died. We also show how the behavior of individuals in the four recently orphaned sibling pairs contrasts to the behavior displayed by chimpanzees in 30 sibling pairs whose mothers were alive. Following the death of their mothers, siblings increased the amount of time they associated, maintained spatial proximity, groomed, reassured, and consoled each other. During travel, younger orphans followed their older siblings, who frequently looked back and waited for them. Both siblings showed distress when separated, and older siblings demonstrated heightened vigilance in dangerous situations. Chimpanzees who were recently orphaned interacted in the preceding ways considerably more than did siblings whose mothers were alive. These findings suggest that siblings provide each other support after maternal loss. Further research is needed to determine whether this support buffers grief and trauma in the immediate aftermath of maternal loss and whether sibling support decreases the probability that orphans will suffer long-term consequences of losing a mother if they survive. [ABSTRACT FROM AUTHOR]

How can collective action evolve when individuals benefit from cooperation regardless of whether they pay its participation costs? According to one influential perspective, collective action problems are common, especially when groups are large, but may be solved when individuals who have more to gain from the collective good or can produce it at low costs provide it to others as a byproduct. Several results from a 20-y study of one of the most striking examples of collective action in nonhuman animals, territorial boundary patrolling by male chimpanzees, are consistent with these ideas. Individuals were more likely to patrol when (i) they had more to gain because they had many offspring in the group; (ii) they incurred relatively low costs because of their high dominance rank and superior physical condition; and (iii) the group size was relatively small. However, several other findings were better explained by group augmentation theory, which proposes that individuals should bear the short-term costs of collective action even when they have little to gain immediately if such action leads to increases in group size and long-term increases in reproductive success. In support of this theory, (i) individual patrolling effort was higher and less variable than participation in intergroup aggression in other primate species; (ii) males often patrolled when they had no offspring or maternal relatives in the group; and (iii) the aggregate patrolling effort of the group did not decrease with group size. We propose that group augmentation theory deserves more consideration in research on collective action. [ABSTRACT FROM AUTHOR]

J. C. Mitani: Associate Editor, Primates. Meanwhile, Itani's first student, Takayoshi Kano, conducted fieldwork on chimpanzees and other primates for his PhD thesis (Kano [9], [10]). [Extracted from the article]

ABSTRACT Objectives Recent studies indicate that the locomotor behavior of wild chimpanzees changes during development. Before transitioning to quadrupedal knuckle-walking in adulthood, young chimpanzees engage in a significant amount of upper limb loading suspensory behavior. We investigated whether these dramatic changes in locomotion influence the strength and shape of chimpanzee long bones. Materials and Methods We examined changes in chimpanzee arboreal locomotion over the course of development using behavioral data collected on wild chimpanzees. We measured the midshaft geometric properties of femora and humeri of wild-caught individuals housed in museum collections using micro computed tomographic scans. Results Chimpanzees spent less time moving arboreally as they aged. Femoral/humeral strength ratios also increased with age, as predicted by the changing loading environment during development. Additional analyses revealed that femoral shape, but not humeral shape, varied across chimpanzee age classes. Adult femora were more elliptical compared with those of infants. This change in adult femora is consistent with the observation that adult chimpanzees spend most of their time moving terrestrially and consequently experience a less variable loading environment than do infants. Discussion Taken together, these findings contribute to our understanding of how ontogenetic changes in function affect form. As similar changes may have characterized the behavioral and skeletal ontogeny of extinct hominoids including hominins, these findings furnish a potential means to make inferences about the behavior of fossil taxa based on the structural properties of their bones. Am J Phys Anthropol 160:16-29, 2016. © 2016 Wiley Periodicals, Inc. [ABSTRACT FROM AUTHOR]

Design: Observations of wild chimpanzees (Pan troglodytes schweinfurthii) were conducted in Kibale National Park, Uganda, at the sites of Sebitoli and Ngogo. Results: We report the first two cases of cleft lip in wild chimpanzees. Additionally, some other chimpanzees in the Sebitoli community show facial dysplasia and congenital anomalies, such as patches of depigmented hairs and limb defects. Conclusions: Cleft lip has been documented in several species of nonhuman primates, but much remains unknown about the occurrence of cleft lip and cleft palate in great apes, probably because such malformations are rare, wild apes are difficult to monitor and observe, and severe cases associated with cleft palates render suckling impossible and lead to early death of infants. The genetic basis of such defects in great apes warrants investigation, as does the possibility that environmental toxins contribute to their etiology in Kibale in ways that could affect humans as well. [ABSTRACT FROM AUTHOR]

Coalitions and alliances are ubiquitous in humans and many other mammals, being part of the fabric of complex social systems. Field biologists and ethologists have accumulated a vast amount of data on coalition and alliance formation, while theoretical biologists have developed modelling approaches. With the accumulation of empirical data and sophisticated theory, we are now potentially able to answer a host of questions about how coalitions emerge and are maintained in a population over time, and how the psychology of this type of cooperation evolved. Progress can only be achieved, however, by effectively bridging the communication gap that currently exists between empiricists and theoreticians. In this paper, we aim to do so by asking three questions: (I) What are the primary questions addressed by theoreticians interested in coalition formation, and what are the main building blocks of their models? (2) Do empirical observations support the assumptions of current models, and if not, how can we improve this situation? (3) Has theoretical work led to a better understanding of coalition formation, and what are the most profitable lines of inquiry for the future? Our overarching goal is to promote the integration of theoretical and field biology by motivating empirical scientists to collect data on aspects of coalition formation that are currently poorly quantified and to encourage theoreticians to develop a comprehensive theory of coalition formation that is testable under real-world conditions. [ABSTRACT FROM AUTHOR]

Observations of chimpanzees (Pan troglodytes) and bonobos (Pan paniscus) provide valuable comparative data for understanding the significance of conspecific killing. Two kinds of hypothesis have been proposed. Lethal violence is sometimes concluded to be the result of adaptive strategies, such that killers ultimately gain fitness benefits by increasing their access to resources such as food or mates. Alternatively, it could be a non-adaptive result of human impacts, such as habitat change or food provisioning. To discriminate between these hypotheses we compiled information from 18 chimpanzee communities and 4 bonobo communities studied over five decades. Our data include 152 killings (n = 58 observed, 41 inferred, and 53 suspected killings) by chimpanzees in 15 communities and one suspected killing by bonobos. We found that males were the most frequent attackers (92% of participants) and victims (73%); most killings (66%) involved intercommunity attacks; and attackers greatly outnumbered their victims (median 8:1 ratio). Variation in killing rates was unrelated to measures of human impacts. Our results are compatible with previously proposed adaptive explanations for killing by chimpanzees, whereas the human impact hypothesis is not supported. [ABSTRACT FROM AUTHOR]

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. [ABSTRACT FROM AUTHOR]

Fossils and molecular data are two independent sources of information that should in principle provide consistent inferences of when evolutionary lineages diverged. Here we use an alternative approach to genetic inference of species split times in recent human and ape evolution that is independent of the fossil record. We first use genetic parentage information on a large number of wild chimpanzees and mountain gorillas to directly infer their average generation times. We then compare these generation time estimates with those of humans and apply recent estimates of the human mutation rate per generation to derive estimates of split times of great apes and humans that are independent of fossil calibration. We date the human-chimpanzee split to at least 7-8 million years and the population split between Neanderthals and modern humans to 400,000-800,000 y ago. This suggests that molecular divergence dates may not be in conflict with the attribution of 6- to 7-million-y-old fossils to the human lineage and 400,000-y-old fossils to the Neanderthal lineage. [ABSTRACT FROM AUTHOR]

Highly frugivorous primates like chimpanzees ( Pan trogolodytes) must contend with temporal variation in food abundance and quality by tracking fruit crops and relying more on alternative foods, some of them fallbacks, when fruit is scarce. We used behavioral data from 122 months between 1995 and 2009 plus 12 years of phenology records to investigate temporal dietary variation and use of fallback foods by chimpanzees at Ngogo, Kibale National Park, Uganda. Fruit, including figs, comprised most of the diet. Fruit and fig availability varied seasonally, but the exact timing of fruit production and the amount of fruit produced varied extensively from year to year, both overall and within and among species. Feeding time devoted to all major fruit and fig species was positively associated with availability, reinforcing the argument that chimpanzees are ripe fruit specialists. Feeding time devoted to figs-particularly Ficus mucuso (the top food)-varied inversely with the abundance of nonfig fruits and with foraging effort devoted to such fruit. However, figs contributed much of the diet for most of the year and are best seen as staples available most of the time and eaten in proportion to availability. Leaves also contributed much of the diet and served as fallbacks when nonfig fruits were scarce. In contrast to the nearby Kanywara study site in Kibale, pith and stems contributed little of the diet and were not fallbacks. Fruit seasons (periods of at least 2 months when nonfig fruits account for at least 40% of feeding time; Gilby & Wrangham., Behavioral Ecology and Sociobiology 61:1771-1779, 2007) were more common at Ngogo than Kanyawara, consistent with an earlier report that fruit availability varies less at Ngogo [Chapman et al., African Journal of Ecology 35:287-302, 1997]. F. mucuso is absent at Kanyawara; its high density at Ngogo, combined with lower variation in fruit availability, probably helps to explain why chimpanzee population density is much higher at Ngogo. [ABSTRACT FROM AUTHOR]

Chimpanzees ( Pan troglodytes) are ecologically flexible omnivores with broad diets comprising many plant and animal foods, although they mostly eat fruit (including figs). Like other ecologically flexible nonhuman primates (e.g., baboons, Papio spp.) with broad diets, their diets vary across habitats. Much data on diets come from short studies that may not capture the range of variation, however, and data are scant on variation within habitats and populations. We present data on diet composition and diversity for chimpanzees at Ngogo, in Kibale National Park, Uganda, collected over a 15-year period, with a focus on the plant components of the diet. We compare Ngogo data to those on chimpanzees at the nearby Kibale site of Kanyawara, on other chimpanzee populations, and on some other frugivorous-omnivorous primates. Results support the argument that chimpanzees are ripe fruit specialists: Ngogo chimpanzees ate a broad, mostly fruit-based diet, feeding time devoted to fruit varied positively with fruit availability, and diet diversity varied inversely with fruit availability. Comparison of Ngogo and Kanyawara shows much similarity, but also pronounced within-population dietary variation. Chimpanzees fed much more on leaves, and much less on pith and stems, at Ngogo. Figs accounted for somewhat less feeding time at Ngogo, but those of Ficus mucuso were quantitatively the most important food. This species is essentially absent at Kanayawara; its abundance and high productivity at Ngogo, along with much higher abundance of several other important food species, help explain why chimpanzee community size and population density are over three times higher at Ngogo. High inter-annual variation at Ngogo highlights the value of long-term data for documenting the extent of ecological variation among chimpanzee populations and understanding how such variation might affect population biology and social dynamics. Am. J. Primatol. 73:1-16, 2011. © 2011 Wiley Periodicals, Inc. [ABSTRACT FROM AUTHOR]

The complex cooperative behavior exhibited by wild chimpanzees generates considerable theoretical and empirical interest, yet we know very little about the mechanisms responsible for its evolution. Here, we investigate the influence of kinship on the cooperative behavior of male chimpanzees living in an unusually large community at Ngogo in Kibale National Park, Uganda. Using long-term field observations and molecular genetic techniques to identify kin relations between individuals, we show that male chimpanzees clearly prefer to affiliate and cooperate with their maternal brothers in several behavioral contexts. Despite these results, additional analyses reveal that the impact of kinship is limited; paternal brothers do not selectively affiliate and cooperate, probably because they cannot be reliably recognized, and the majority of highly affiliative and cooperative dyads are actually unrelated or distantly related. These findings add to a growing body of research that indicates that animals cooperate with each other to obtain both direct and indirect fitness benefits and that complex cooperation can occur between kin and nonkin alike. [ABSTRACT FROM AUTHOR]

Although estrous synchrony has been reported in a number of mammalian species, most often among primates, methodological and analytical problems make it difficult to interpret these results. We developed a novel estrous synchrony index and employed a randomization procedure to analyze long-term observations of female chimpanzee ( Pan troglodytes schweinfurthii) estrous cycles at the Mahale Mountains National Park, Tanzania. Our results revealed that female chimpanzees at Mahale avoid synchronizing their estrous periods with each other. We also found that birthrates decreased as the breeding sex ratio increased. We suggest that estrous asynchrony decreases female–female competition for mates. Asynchrony may also reduce the potential for male sexual coercion by nonpreferred mating partners. [ABSTRACT FROM AUTHOR]

Chimpanzees (Pan troglodytes) have hostile intergroup relations throughout most or all of their geographic range. Hostilities include aggressive encounters between members of neighboring communities during foraging and during patrols in which members of one community search for neighbors near territory boundaries. Attacks on neighbors involve coalitions of adult males, and are sometimes fatal. Targets include members of all age/sex classes, but the risk of lethal intergroup coalitionary aggression is highest for adult males and infants, and lowest for sexually swollen females. The best-supported adaptive explanation for such behavior is that fission-fusion sociality allows opportunities for low-cost attacks that, when successful, enhance the food supply for members of the attackers' community, improve survivorship, and increase female fertility. We add to the database on intergroup coalitionary aggression in chimpanzees by describing three fatal attacks on adult males, plus a fourth attack on an adult male and an attack on a juvenile that were almost certainly fatal. Observers saw four of these attacks and inferred the fifth from forensic and behavioral evidence. The attackers were males in two habituated, unprovisioned communities (Ngogo and Kanyawara) in Kibale National Park, Uganda. We also summarize data on other intercommunity attacks at Ngogo. Our observations are consistent with the “imbalance of power” hypothesis [Manson & Wrangham, Current Anthropology 32:369–390, 1991] and support the argument that lethal coalitionary intergroup aggression by male chimpanzees is part of an evolved behavioral strategy. Am. J. Primatol. 68:161-180, 2006. © 2006 Wiley-Liss, Inc. [ABSTRACT FROM AUTHOR]

The article offers a look at the contributions of Professor Toshisada Nishida to the study of primate behavior and primatology. Nishida pioneered the study of chimpanzee behavior, its culture and its conservation starting in 1965. Nishida retires as the head of Kyoto University's Laboratory of Human Evolution Studies in 2004.

Recent research has revealed substantial diversity in the behavior of wild chimpanzees. Understanding the sources of this variation has become a central focus of investigation. While genetic, ecological, and cultural factors are often invoked to explain behavioral variation in chimpanzees, the demographic context is sometimes overlooked as a contributing factor. Observations of chimpanzees at Ngogo, Kibale National Park, Uganda, reveal that the size and structure of the unit group or community can both facilitate and constrain the manifestation of behavior. With approximately 150 individuals, the Ngogo community is much larger than others that have been studied in the wild. We have taken advantage of the unusual demographic structure of this community to document new and intriguing patterns of chimpanzee behavior with respect to hunting, territoriality, and male social relationships. Chimpanzees at Ngogo hunt often and with a considerable degree of success. In addition, male chimpanzees there frequently patrol the boundary of their territory and engage in repeated bouts of lethal intergroup aggression. By forming two distinct subgroups, male chimpanzees at Ngogo also develop social bonds above the level of dyadic pairs. While the sheer number of chimpanzees contributes to differences in hunting, patrolling, mating, and subgrouping at Ngogo, the demographic situation may also constrain behavioral interactions. At Ngogo, male chimpanzees who are closely related genetically through the maternal line do not appear to affiliate or cooperate with each other. Demographic constraints may be responsible for this finding. In this paper, I use these examples to illustrate how the demographic context affects the possible range of behavioral options open to individuals and ultimately contributes to the explanation of behavioral diversity in chimpanzees. [ABSTRACT FROM AUTHOR]

Strong social bonds typically develop between dyadic pairs of male chimpanzees. These bonds are manifest in several contexts, including association, grooming, and proximity. Here we demonstrate that social bonds exist at a higher level of organization among males living in an extremely large community at Ngogo, Kibale National Park, Uganda. An analysis of over 2,500 hours of observation of 35 individuals revealed two distinct subgroups of male chimpanzees. Males that composed each subgroup can be identified on the basis of their tendency to associate in temporary parties. Matrix permutation tests indicated that subgroup members tended to maintain spatial proximity to each other and participate together in territorial boundary patrols. Subgroups formed along the lines of age and rank; members of a small subgroup were younger and lower ranking than individuals in a larger subgroup. Despite this social clustering of males, community integrity remained intact with low levels of aggression between individuals of different subgroups. After controlling for the effect of association, significantly more aggression occurred within compared to between subgroups. In addition, males of the different subgroups displayed significant overlap in their use of the community territory and thus showed no tendency to divide spatially. We compare our findings with those from other animal species and chimpanzee populations and discuss them in the context of the unusual demography of the Ngogo community. [ABSTRACT FROM AUTHOR]

Demography provides critical data to increase our understanding of the evolution, ecology, and conservation of primate populations. The chimpanzees of the Mahale Mountains National Park, Tanzania, have been studied for more than 34 yr on the basis of individual identification and standardized attendance records. From this long-term study, we derived the following demographic data: The major cause of death was disease (48%), followed by senescence (24%) and within-species aggression (16%). Fifty percent of Mahale chimpanzees died before weaning. The median ages of female life history variables were: first maximal swelling, 10.0 yr (n=5); emigration, 11.0 yr (n=11); and first birth, 13.1 yr (n=5). The median period of adolescent infertility was 2.8 yr (n=4) when calculated from the age at immigration to that at first birth. Female fecundity was highest between 20 and 35 yr of age, with an annual birth rate of 0.2. Twenty-six females that were observed from a young age (10–13 yr) to death at various ages (15–40 yr) gave birth to an average of 3.9 and weaned an average of 1.4 offspring. Twenty-five females that were observed from middle age (18–33 yr) to death in older age (31–48) gave birth to an average of 2.7 and weaned an average of 2.0 offspring. The post-reproductive lifespan for female chimpanzees was defined as the number of years that passed from the year when the last offspring was born to the year when the female died, minus 5. Twenty-five percent of old females had a post-reproductive lifespan. The interbirth interval after the birth of a son (&xmacr; = 72 mo) tended to be longer than that after the birth of a daughter (&xmacr; = 66 mo). The extent of female transfer, which is a rule in chimpanzees, is influenced by the size and composition of the unit group and size of the overall local community. Am. J. Primatol. 59:99–121, 2003. © 2003 Wiley-Liss, Inc. [ABSTRACT FROM AUTHOR]

Chimpanzees (Pan troglodytes ) prey on a variety of vertebrates, mostly on red colobus (Procolobus spp. ) where the two species are sympatric. Variation across population occurs in hunting frequency and success, in whether hunting is cooperative, i.e., payoffs to individual hunters increase with group size, and in the extent to which hunters coordinate their actions in space and time, and in the impact of hunting on red colobus populations. Also, hunting frequency varies over time within populations, for reasons that are unclear. We present new data on hunting by chimpanzees at Ngogo, Kibale National Park, Uganda, and combine them with earlier data (Mitani and Watts, 1999, Am. J. Phys. Anthropol. 109: 439–454) to examine hunting frequency and success, seasonality, and cooperation. The Ngogo community is the largest and has the most males of any known community. Chimpanzees there mostly hunt red colobus and are much more successful and make many more kills per hunt than at other sites; they kill 6–12% of the red colobus population annually. The number of kills and the offtake of meat per hunt increase with the number of hunters, but per capita meat intake is independent of hunting party size; this suggests that cheating occurs in large parties. Some behavioral cooperation occurs. Hunting success and estimated meat intake vary greatly among males, partly due to dominance rank effects. The high overall success rate leads to relatively high average per capita meat intake despite the large number of consumers. The frequency of hunts and of hunting patrols varies positively with the availability of ripe fruit; this is the first quantitative demonstration of a relationship between hunting frequency and the availability of other food, and implies that the chimpanzees hunt most when they can easily meet energy needs from other sources. We provide the first quantitative support for the argument that variation in canopy structure influences decisions to hunt red colobus because hunts are easier where the canopy is broken. [ABSTRACT FROM AUTHOR]

Chimpanzees are among the few mammals that engage in lethal coalitionary aggression between groups. Most attacks on neighbors occur when parties made up mostly of adult males patrol boundaries of their community's range. Patrols have time, energy, and opportunity costs, and entail some risks despite the tendency of males to attack only when they greatly outnumber their targets. These factors may lead to a collective action problem. Potential benefits include protection of community members, particularly infants; range expansion and increases in the amount and quality of food available; and incorporation of more females into the community. Males may not share these equally; for example, those able to obtain large shares of matings may stand to gain most by participating in patrols and to lose most by refraining. Despite the attention that boundary patrolling has attracted, few relevant quantitative data are available. Here, we present detailed data on boundary patrolling and intergroup aggression in a chimpanzee community at Ngogo, Kibale National Park, Uganda, that is unusually large and has more males than any other known community. Males there patrolled much more often, and patrol parties were much larger on average, than at two other sites for which comparative data exist. Our findings support the argument that male participation varies along with variation in potential gains, in willingness to take risks, and in skill at handling these risks. Both the overall frequency with which individual males patrolled and their willingness to join patrols as others set off on them were positively associated with variation in mating success, in participation in hunts of red colobus monkeys, and in hunting success. Males patrolled relatively often with others with whom they associated often in general, with whom they often groomed, and with whom they formed coalitions in withincommunity agonism. This indicates that they were most willing to take risks associated with patrolling when with others they trusted to take the same risks. [ABSTRACT FROM AUTHOR]

Researchers have documented infanticide by adult males in four wild chimpanzee populations. Males in three of these have killed infants from outside of their own communities, but most infanticides, including one from Kanyawara, in Kibale National Park, Uganda, took place within communities. Here we report two new cases of infanticide by male chimpanzees at a second Kibale site, Ngogo, where the recently habituated chimpanzee community is the largest yet known. Both infanticides happended during boundary patrols, which occur at a high frequency there. Patrolling males attacked solitary females who were unable to defend their infants successfully. The victims were almost certainly not members of the Ngogo community. Males cannibalized both infants and completely consumed their carcasses. These observations show that infanticide by males is widespread in the Kibale population and that between-community infanticide also happens there. We discuss our observations in the context of the sexual selection hypothesis and other proposed explanations for infanticide by male chimpanzees. The observations support the arguments that infanticide has been an important selective force in chimpanzee social evolution and that females with dependent infants can be at great risk near range boundaries, but why male chimpanzees kill infants is still uncertain. [ABSTRACT FROM AUTHOR]

Few data exist regarding long-term changes in primate populations in old-growth, tropical forests. In the absence of this information, it is unclear how to assess population trends efficiently and economically. We addressed these problems by conducting line-transect censuses 23.5 years apart at the Ngogo study area in Kibale National Park, Uganda. We conducted additional censuses over short time intervals to determine the degree to which the temporal distribution of censuses affected estimates of primate numbers. Results indicate that two species, blue monkeys and red colobus, may have experienced significant reductions over the past 23.5 years at Ngogo. In contrast, five other species, baboons, black-and-white colobus, chimpanzees, mangabeys, and red-tailed guenons, have not changed in relative abundance. Additional findings indicate that different observers may vary significantly in their estimates of sighting distances of animals during censuses, thus rendering the use of measures of absolute densities problematic. Moreover, censuses conducted over short periods produce biased estimates of primate numbers. These results provide guidelines for the use of line-transect censuses and underscore the importance of protecting large blocks of forests for primate conservation. [ABSTRACT FROM AUTHOR]

Wild chimpanzees form temporary parties that vary in size and composition. Previous studies have revealed considerable intraspecific variation in party compositions. We examined patterns of association among age, sex, and reproductive classes of chimpanzees at Ngogo in the Kibale National Park, Uganda. We employed a class-based association index and a randomization procedure to control for confounding factors and to test for differences between classes. Results indicate that males associated with other males significantly more than expected if all classes behaved equivalently, while females generally associated with individuals of the same sex less than expected. To interpret these patterns we used two additional indices that separate associations into two components: general gregariousness and preference for particular classes of associates. Males and estrous females were more gregarious than other classes, while anestrous females were less so. After controlling for general gregariousness, adult males as a class showed no specific preference for associating with each other. Anestrous females preferred each other as party members, and estrous females avoided each other. These results are consistent with previous findings that adult males are more gregarious than females. They diverge from the standard picture of chimpanzee society, however, by suggesting a mutual affinity among anestrous females, but not among adult males as a class. [ABSTRACT FROM AUTHOR]

Presents information on a study which investigated the demographic effects on the hunting behavior of a community of wild chimpanzees at Ngogo in the Kibale National Park, Uganda. Methods; Results; Discussion and comparisons with other studies.

Investigates the factors that facilitate the evolution of individual vocal signatures of wild chimpanzees by examining two components of call repertoire--the pant hoot and the pant grunt. Within-individual acoustic variation; Between individual acoustic variation.

Examines screams of chimpanzees and bonobos to investigate variability in call structure. Differences between and within species; Spectral characteristics; Acoustic differences; Factors for variations.

Responses of mated individuals to playbacks of the songs of solitary females and males permit evaluating the behavioral mechanisms maintaining monogamy in gibbons. When female songs are played back from the centers of their ranges, mated female gibbons typically initiate duets and group approaches toward playback sites. Female songs played back from range boundary locations elicit duetting responses. Responses to solitary female songs do not differ from responses to song duets used by established mated pairs to mediate patterns of intergroup spacing. Mated males lead silent group approaches toward the sites of male song playbacks. These results suggest that range defense by female gibbons forces males into accepting monogamous mating relationships and that monogamy in gibbons is regulated by intersexually-supported, intrasexual aggression. [ABSTRACT FROM AUTHOR]

1. Existing theory suggests that territoriality will evolve when resources are limited and defendable, but defendability has seldom been analyzed quantitatively. 2. Here we argue that defendability depends on the ability of an animal to monitor the boundaries of its range in order to detect potential intruders and introduce an index of defendability (D) which is the ratio of observed daily path length (d) to an area equal to the diameter (d') of a circle with area equal to home range area of the animal. This index is sensitive only to extreme deviation from circular shape. 3. Review of the literature on primate ranging reveals that all territorial groups for which data are available have an index of 1.0 or greater, and that few nonterritorial species have an index of 1.0 or greater. 4. Regression analysis of the relationship of daily path length to feeding group weight and foliage in the diet reveals that both feeding group weight and foliage in the diet account for a large proportion of the variance in daily path length, and that territorial and nonterritorial groups do not differ in day range for a given group weight and diet. [ABSTRACT FROM AUTHOR]

Primate social groups frequently contain multiple males. Male group size has been hypothesized to result from male mating competition, but the selective factors responsible for the evolution of multimale groups are unclear. Short breeding seasons create situations that are not conducive for single males to monopolize mating access to females, and may therefore favor the formation of large male groups. Alternately, since the costs of mate defense increase with the spatial clumping of females, female group size may be a primary determinant of the number of males in a primate group. We used comparative methods designed to control for the potentially confounding effects of hidden third variables associated with phylogeny to test the breeding season and female group size hypotheses for the evolution of multimale groups. Our results revealed no association between breeding season duration and the number of males in groups. In contrast, we provide support for the female group size hypothesis by demonstrating a strong pattern of correlated evolution between female and male group size. [ABSTRACT FROM AUTHOR]

Chimpanzees emit a loud, species-typical long distance call known as the pant hoot. Geographic variation between the pant hoots of chimpanzees living in two neighboring populations, the Mahale Mountains and Gombe Stream National Parks, Tanzania, was examined. Analysis of six acoustic features revealed subtle differences in the way chimpanzees from the two populations called. Individuals from the Mahale study site uttered one section of their pant hoots at a faster rate and with shorter elements than animals from Gombe Stream. In addition, individuals at Mahale delivered broader-band, higher pitched "climax" elements than males from Gombe. While genetic factors, anatomical differences, variations in the use of calls at the two sites, and changes in calling over time may account for the variability between populations, we suggest the additional possibility that differences in pant hooting may be due to learning. [ABSTRACT FROM AUTHOR]

Acoustic analyses and field playback experiments were conducted to investigate interspecific song differences and discrimination in male gibbons. Songs were recorded and analyzed from males of three populations: agile gibbons (Hylobates agilis) in West Kalimantan and South Sumatra and Müller's gibbons (Hylobates muelleri) in East Kalimantan. Male songs from each of the three populations were played back to West Kalimantan agile gibbons. Acoustic analyses indicated that the songs of the two Species differed in spectral features and in the temporal patterning of syllables. Agile gibbon songs from the West Kalimantan and South Sumatran populations varied in song durations and in the maximum frequencies of one song element. Field playbacks showed that West Katimantan agile gibbons respond in a similar manner to local songs and to conspecific songs from South Sumatra. In contrast, the animals responded differentially to conspecific and heterospecific Müller's gibbon songs. These results indicate that gibbons discriminate specific differences in songs, and support the division of agile and Müller's gibbons into distinct species. [ABSTRACT FROM AUTHOR]