Declarative referential gesturing in a wild chimpanzee (Pan troglodytes).

Humans are argued to be unique in their ability and motivation to share attention with others about external entities-sharing attention for sharing's sake. Indeed, in humans, using referential gestures declaratively to direct the attention of others toward external objects and events emerges in the first year of life. In contrast, wild great apes seldom use referential gestures, and when they do, it seems to be exclusively for imperative purposes. This apparent species difference has fueled the argument that the motivation and ability to share attention with others is a human-specific trait with important downstream consequences for the evolution of our complex cognition [M. Tomasello, Becoming Human (2019)]. Here, we report evidence of a wild ape showing a conspecific an item of interest. We provide video evidence of an adult female chimpanzee, Fiona, showing a leaf to her mother, Sutherland, in the context of leaf grooming in Kibale Forest, Uganda. We use a dataset of 84 similar leaf-grooming events to explore alternative explanations for the behavior, including food sharing and initiating dyadic grooming or playing. Our observations suggest that in highly specific social conditions, wild chimpanzees, like humans, may use referential showing gestures to direct others' attention to objects simply for the sake of sharing. The difference between humans and our closest living relatives in this regard may be quantitative rather than qualitative, with ramifications for our understanding of the evolution of human social cognition.

The use of chimpanzee-modified faunal assemblages to investigate early hominin carnivory.

Chimpanzees regularly hunt and consume prey smaller than themselves. It seems therefore likely that early hominins also consumed small vertebrate meat before they started using and producing stone tools. Research has focused on cut marks and large ungulates, but there is a small body of work that has investigated the range of bone modifications produced on small prey by chimpanzee mastication that, by analogy, can be used to identify carnivory in pre-stone tool hominins. Here, we review these works along with behavioral observations and other neo-taphonomic research. Despite some equifinality with bone modifications produced by baboons and the fact that prey species used in experiments seldom are similar to the natural prey of chimpanzees, we suggest that traces of chimpanzee mastication are sufficiently distinct from those of other predators that they can be used to investigate mastication of vertebrate prey by early hominins.

Investigating infant feeding development in wild chimpanzees using stable isotopes of naturally shed hair.

Measuring the relative contributions of milk and non-milk foods in the diets of primate infants is difficult from observations. Stable carbon (δ13 C) and nitrogen (δ15 N) isotopes in hair can be used to physiologically track infant feeding through development, but few wild studies have done so, likely due to the difficulty in collecting hair non-invasively. We assessed infant feeding at different ages in wild chimpanzees (Pan troglodytes) at Ngogo, Uganda using δ13 C and δ15 N of keratin in 164 naturally shed hairs from 29 infants (61 hairs), 6 juveniles (7 hairs), 28 mothers (67 hairs) and 14 adult males (29 hairs). Hairs were collected when they stuck to feces during defecation or from the ground after chimpanzees groomed or rested. We could not distinguish between the hairs of infants and mothers using strand length and diameter. Infants 1-2 years old were most enriched in 13 C and 15 N and showed means of 1.1‰ in δ13 C and 2.1‰ in δ15 N above their mothers. Infants at 2 years had hair δ13 C values like those of their mothers, which suggests infants began relying more heavily on plants around this age. While mother-infant δ13 C and δ15 N differences generally decreased with offspring age, as is expected when infants rely increasingly more on independent foraging through development, milk seemed to remain an important dietary component for infants older than 2.5 years, as evidenced by continuing elevated δ15 N. We showed that stable carbon and nitrogen isotopes in naturally shed hairs can feasibly detect trophic level differences between chimpanzee infants and mothers. Since it can mitigate some of the limitations associated with behavioral and fecal stable isotope data, the use of hair stable isotopes is a useful, non-invasive tool for assessing infant feeding development in wild primates.

Male chimpanzee sexual coercion and mating success at Ngogo.

Chimpanzees (Pan troglodytes) have a complex mating system in which both sexes use multiple tactics. Most copulations occur in group contexts, and high-ranking males can gain high mating and reproductive success, but females typically mate with multiple males and the potential for sperm competition is high. Also, male-female dyads sometimes form temporary exclusive mating associations (consortships). Male aggression to receptive females is common. Several studies have supported the hypothesis that this is sexual coercion, but debate exists regarding the importance of coercion relative to that of female choice. The number of adult males in a community can influence the balance between these processes. In the large Ngogo community, male dominance ranks and rates of aggression to fully-swollen females were positively related to mating success as estimated by copulation rates and by proportions of copulations achieved. Aggression rates were higher than at other sites, overall and per male, especially during periovulatory periods, and increased with the number of males associating with a female. Aggression impaired female foraging efficiency. Males initiated most copulations and females rarely refused mating attempts. Male-to-female grooming was positively associated with male mating success and with the proportion of copulations that females initiated, but the amount of grooming was typically small and whether grooming-for-mating trading occurs is uncertain. These results are consistent with the hypothesis that sexual coercion is an important component of male chimpanzee mating strategies in many sociodemographic circumstances, but also show that male tactics vary both in response to and independently of those circumstances.

Meat eating by nonhuman primates: A review and synthesis.

Most nonhuman primates prey on vertebrates. Meat-eating, defined as ingestion of vertebrate tissue, occurs in 12 families, ≥39 genera, and ≥89 species. It is most common in capuchins (Cebus and Sapajus spp.), baboons (Papio spp.), bonobos (Pan paniscus), and chimpanzees (Pan troglodytes) and modestly common in blue monkeys (Cercopithecus mitis), callitrichids (Callithrix spp. and Saguinus spp.), and squirrel monkeys (Saimiri spp.). It is uncommon in other cercopithecines, rare in other haplorhines and in lemurs, and virtually absent in colobines. Birds are the prey class eaten by the most species (≥53), followed by reptiles (≥48), amphibians (≥38), mammals (≥35), and fish (≥7). Major hypotheses for the importance of meat eating are that it is (1) mainly an energy source, especially (1a) when plant-source foods (PSFs) with high energy return rates are scarce (energy shortfall hypothesis); (2) mainly a protein source; and (3) mainly a source of micronutrients scarce in PSFs. Meat eating bouts sometimes provide substantial energy and protein, and some chimpanzees gain substantial protein from meat monthly or annually. However, meat typically accounts for only small proportions of feeding time and of total energy and protein intake, and quantitative data are inconsistent with the energy shortfall hypothesis. PSFs and/or invertebrates are presumably the main protein sources, even for chimpanzees. Support is strongest for the micronutrient hypothesis. Most chimpanzees eat far less meat than recorded for hunter-gatherers, but the highest chimpanzee estimates approach the lowest for African hunter-gatherers. In fundamental contrast to the human predatory pattern, other primates only eat vertebrates much smaller than they are, tool-assisted predation is rare except in some capuchins and chimpanzees, and tool use in carcass processing is virtually absent. However, harvesting of small prey deserves more attention with reference to the archaeological and ethnographic record.

Premasticated food transfer by wild chimpanzee mothers with their infants: Effects of maternal parity, infant age and sex, and food properties.

Premasticated food transfer, when an individual partially breaks down food through chewing and feeds it to another individual, usually mouth-to-mouth, is described widely across human cultures. This behavior plays an important role in modern humans' strategy of complementary feeding, which involves supplementing maternal milk in infant diets with processed, easily digestible versions of adult foods. The extent to which other primates engage in premasticated food transfer with infants is unclear, as premasticated food transfers have been only occasionally reported in other ape species. We investigated premasticated food transfers in 62 mother-infant pairs of wild chimpanzees at Ngogo, Uganda, as well as unresisted food taking, when mothers passively allow infants to seize food. We evaluated the presence or absence and rates of premasticated food transfer and unresisted food taking relative to maternal parity and infant age and sex and assessed the food species and part used. We found that chimpanzee mothers regularly shared premasticated food with their infants aged between 6 months and 4 years, but they were more likely to share, and more frequently shared, with younger infants. The frequency with which females shared premasticated food may relate to maternal experience, as multiparous females shared premasticated food more often than did first-time mothers, which we did not find with unresisted food taking. Both easy-to-chew, commonly eaten foods and tougher, rarely eaten foods were shared. Premasticated food transfer and unresisted food taking may be infant-rearing strategies to facilitate the transition from a diet of exclusive maternal milk to solid food during early infancy. Premasticated food transfer in particular may provide energetic, immune, or growth benefits to infants through reduced chewing effort and maternal saliva. Given our findings in chimpanzees and earlier reports in other ape species, we suggest that the foundation of complementary feeding, a uniquely hominin strategy, might have been present in a common ancestor shared with the other great apes in the form of premasticated, mouth-to-mouth food transfer by mothers with their offspring.

Lethal aggression in Pan is better explained by adaptive strategies than human impacts.

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.

Long-term trends in fruit production in a tropical forest at Ngogo, Kibale National Park, Uganda.

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.

Group augmentation, collective action, and territorial boundary patrols by male chimpanzees.

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.

Favorable ecological circumstances promote life expectancy in chimpanzees similar to that of human hunter-gatherers.

Demographic data on wild chimpanzees are crucial for understanding the evolution of chimpanzee and hominin life histories, but most data come from populations affected by disease outbreaks and anthropogenic disturbance. We present survivorship data from a relatively undisturbed and exceptionally large community of eastern chimpanzees (Pan troglodytes schweinfurthii) at Ngogo, Kibale National Park, Uganda. We monitored births, deaths, immigrations, and emigrations in the community between 1995 and 2016. Using known and estimated ages, we calculated survivorship curves for the whole community, for males and females separately, and for individuals ≤2 years old when identified. We used a novel method to address age estimation error by calculating stochastic survivorship curves. We compared Ngogo life expectancy, survivorship, and mortality rates to those from other chimpanzee communities and human hunter-gatherers. Life expectancy at birth for both sexes combined was 32.8 years, far exceeding estimates of chimpanzee life expectancy in other communities, and falling within the range of human hunter-gatherers (i.e., 27-37 years). Overall, the pattern of survivorship at Ngogo was more similar to that of human hunter-gatherers than to other chimpanzee communities. Maximum lifespan for the Ngogo chimpanzees, however, was similar to that reported at other chimpanzee research sites and was less than that of human-hunter gatherers. The absence of predation by large carnivores may contribute to some of the higher survivorship at Ngogo, but this cannot explain the much higher survivorship at Ngogo than at Kanyawara, another chimpanzee community in the same forest, which also lacks large carnivores. Higher survivorship at Ngogo appears to be an adaptive response to a food supply that is more abundant and varies less than that of Kanyawara. Future analyses of hominin life history evolution should take these results into account.

A novel fecal stable isotope approach to determine the timing of age-related feeding transitions in wild infant chimpanzees.

OBJECTIVES: Determining nutritional development in wild primates is difficult through observations because confirming dietary intake is challenging. Physiological measures are needed to determine the relative contributions of maternal milk and other foods at different ages, and time of weaning. We used fecal stable carbon and nitrogen isotopes (δ13 C, δ15 N) and fecal nitrogen concentrations (%N) from wild chimpanzees at Ngogo, Uganda, to derive physiological dietary indicators during the transition from total reliance on maternal milk to adult foods after weaning. MATERIALS AND METHODS: We analyzed 560 fecal samples collected non-invasively from 48 infants, their mothers, and 6 juvenile siblings. Most infant and juvenile samples (90%) were matched to samples collected from mothers on the same day. Isotopic assessments were compared with observations of nursing and feeding. RESULTS: Infants ≤1 year old showed average δ15 N, δ13 C and %N ratios that were 2.0‰, 0.8‰ and 1.3% greater than their mothers, respectively, interpreted as trophic level effects. Although data collected on newborns were few, results suggest that solid foods were consumed within 2-5 months after birth. Trophic level differences decreased steadily after 1 year, which indicates a decreasing relative contribution of milk to the diet. Isotopic results indicated infants were weaned by 4.5 years old-more than a year earlier than observations of nipple contacts ceased, which revealed the occurrence of "comfort nursing." Juvenile isotopic signatures indicate no nursing overlap between siblings. DISCUSSION: Our results resemble the stable isotope differences of human babies. This study contributes to a model of chimpanzee nutritional development required to understand early life history patterns in hominins.

Ecological factors influencing habitat use by chimpanzees at Ngogo, Kibale National Park, Uganda.

Although numerous ecological and social factors influence range use in vertebrates, the general assumption is that ranging patterns typically accord with principles of optimal foraging theory. However, given temporal variability in resource abundance, animals can more easily meet nutritional needs at some times than at others. For species in which sociality is particularly important for fitness, such as chimpanzees (Pan troglodytes) and other group-living primates, the influences of social factors can be particularly strong, and likely interact closely with ecological factors. We investigated home range use by a community of chimpanzees at Ngogo, Kibale National Park, Uganda, to determine whether range use corresponded to energy-based optimality principles. Chimpanzees were particularly attracted to areas of the home range where individuals of Ficus mucuso (a large but low-density resource) were found, but only if those areas also offered other preferred or important resource classes. The aggregation of large foraging parties at F. mucuso crowns (frequently seen year-round) facilitates a number of socially beneficial activities for both males and females. Because chimpanzees apparently seek out F. mucuso in areas where other high-quality feeding opportunities exist, these social benefits likely do not come at the expense of fitness benefits accrued from feeding on high-quality resources. Am. J. Primatol. 78:432-440, 2016. © 2015 Wiley Periodicals, Inc.

Two Cases of Cleft Lip and Other Congenital Anomalies in Wild Chimpanzees Living in Kibale National Park, Uganda.

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.

Removing snares is an effective conservation intervention: a case study involving chimpanzees.

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.

Chimpanzee-red colobus encounter rates show a red colobus population decline associated with predation by chimpanzees at Ngogo.

Chimpanzees (Pan troglodytes) hunt various primates, but concentrate on red colobus monkeys (Piliocolobus spp.) wherever the two species are sympatric. The extraordinarily large Ngogo chimpanzee community in Kibale National Park, Uganda, preys heavily on the local population of red colobus (P. tephrosceles). Census data showed a steep decline in this population in the center of the chimpanzees' home range between 1975 and 2007 [Lwanga et al., 2011; Teelen, 2007b]. Given no obvious change in food availability, predation by chimpanzees was the most likely cause [ibid.; Teelen, 2008]. However, census data from other parts of the home range raised the possibility that the decline was restricted to this central area [Teelen, 2007a] We present data from 1998 to 2012 on the rate of encounters between chimpanzees and red colobus that provide a chimpanzee-centered estimate of red colobus density, thus of predation opportunities, throughout the home range. These corroborate census data by showing a long-term decline in encounters near the center. They also show that encounters become relatively more common at increasing distances from the center, but encounter rates have decreased even in peripheral areas and, by implication, the red colobus population has declined throughout the study area. These data corroborate Teelen's [2008] conclusion that chimpanzee predation on red colobus during the 1990s and early 2000s was unsustainable. Hunting rates and prey offtake rates have also declined markedly; whether this will allow the red colobus population to recover is unknown. In contrast, rates at which chimpanzees encountered redtail monkeys (Cercopithecus ascanius) and grey-cheeked mangabeys (Lophocebus albigena) did not decrease. Neither did they increase, however, contrary to long-term census data from the center of the study area [Lwanga et al., 2011].

Demographic and hormonal evidence for menopause in wild chimpanzees.

Among mammals, post-reproductive life spans are currently documented only in humans and a few species of toothed whales. Here we show that a post-reproductive life span exists among wild chimpanzees in the Ngogo community of Kibale National Park, Uganda. Post-reproductive representation was 0.195, indicating that a female who reached adulthood could expect to live about one-fifth of her adult life in a post-reproductive state, around half as long as human hunter-gatherers. Post-reproductive females exhibited hormonal signatures of menopause, including sharply increasing gonadotropins after age 50. We discuss whether post-reproductive life spans in wild chimpanzees occur only rarely, as a short-term response to favorable ecological conditions, or instead are an evolved species-typical trait as well as the implications of these alternatives for our understanding of the evolution of post-reproductive life spans.

Diet of chimpanzees (Pan troglodytes schweinfurthii) at Ngogo, Kibale National Park, Uganda, 1. Diet composition and diversity.

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.

Diet of chimpanzees (Pan troglodytes schweinfurthii) at Ngogo, Kibale National Park, Uganda, 2. Temporal variation and fallback foods.

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.

Effects of infant age and sex, and maternal parity on the interaction of lactation with infant feeding development in chimpanzees.

The interaction between infant feeding and maternal lactational physiology influences female inter-birth intervals and mediates maternal reproductive trade-offs. We investigated variation in feeding development in 72 immature wild chimpanzees (Pan troglodytes schweinfurthii) at Ngogo, Kibale National Park, Uganda, and made inferences about maternal lactation over the course of infancy. We compared the percentage (%) of time that mothers nursed infants as a function of infant age and assessed how hourly rates and bout durations of nursing and foraging varied in association with differences in offspring age, sex, and maternal parity. Nursing % times, rates and durations were highest for infants ≤ 6 months old but did not change significantly from 6 months to 5 years old. Nursing continued at a decreasing rate for some 5- to 7-year-olds. Infants ≤ 6 months old foraged little. Foraging rates did not change after 1 year old, but foraging durations and the % time devoted to foraging increased with age. Independent foraging probably became a dietary requirement for infants at 1 year old, when their energy needs may have surpassed the available milk energy. Infants spent as much time foraging by the time they were 4 to 5 years old as adults did. No sex effect on infant nursing or foraging was apparent, but infants of primiparous females had higher foraging rates and spent more time foraging than the infants of multiparous females did. Although no data on milk composition were collected, these findings are consistent with a working hypothesis that like other hominoids, chimpanzee mothers maintained a fixed level of lactation effort over several years as infants increasingly supplemented their growing energy, micronutrient and hydration needs via independent foraging. Plateauing lactation may be a more widespread adaptation that allows hominoid infants time to attain the physiology and skills necessary for independent feeding, while also providing them with a steady dietary base on which they could rely consistently through infancy, and enabling mothers to maintain a fixed, predictable level of lactation effort.

Lethal Coalitionary Aggression Associated with a Community Fission in Chimpanzees (Pan troglodytes) at Ngogo, Kibale National Park, Uganda.

Many animals engage in aggression, but chimpanzees stand out in terms of fatal attacks against adults of their own species. Most lethal aggression occurs between groups, where coalitions of male chimpanzees occasionally kill members of neighboring communities that are strangers. However, the first observed cases of lethal violence in chimpanzees, which occurred at Gombe, Tanzania in the 1970s, involved chimpanzees that once knew each other. They followed the only observed case of a permanent community fission in chimpanzees. A second permanent fission recently transpired at Ngogo, Kibale National Park, Uganda. Members of a large western subgroup gradually ceased associating peacefully with the rest of the community and started behaving antagonistically toward them. Affiliation effectively ended by 2017. Here, we describe two subsequent lethal coalitionary attacks by chimpanzees of the new western community on males of the now separate central community, one in 2018 and the second in 2019. The first victim was a young adult male that never had strong social ties with his attackers. The second was a high-ranking male that had often associated with the western subgroup before 2017; he groomed regularly with males there and formed coalitions with several. Other central males present at the start of the second attack fled, and others nearby did not come to the scene. Several western females joined in the second attack; we suggest that female-female competition contributed to the fission. This event highlighted the limits on protection afforded by long-term familiarity and the constraints on costly cooperation among male chimpanzees.