Future coexistence with great apes will require major changes to policy and practice.

The great apes-bonobos, chimpanzees, gorillas and orangutans-are critically threatened by human activities. We have destroyed their habitats, hunted them and transmitted fatal diseases to them. Yet we also conduct research on them, try to protect them and live alongside them. They are endangered, and time is running out. Here we outline what must be done to ensure that future generations continue to share this planet with great apes. We urge dialogue with those who live with great apes and interact with them often. We advocate conservation plans that acknowledge the realities of climate change, economic drivers and population growth. We encourage researchers to use technology to minimize risks to great apes. Our proposals will require substantial investment, and we identify ways to generate these funds. We conclude with a discussion of how field researchers might alter their work to protect our closest living relatives more effectively.

The primate gut mycobiome-bacteriome interface is impacted by environmental and subsistence factors.

The gut microbiome of primates is known to be influenced by both host genetic background and subsistence strategy. However, these inferences have been made mainly based on adaptations in bacterial composition - the bacteriome and have commonly overlooked the fungal fraction - the mycobiome. To further understand the factors that shape the gut mycobiome of primates and mycobiome-bacteriome interactions, we sequenced 16 S rRNA and ITS2 markers in fecal samples of four different nonhuman primate species and three human groups under different subsistence patterns (n = 149). The results show that gut mycobiome composition in primates is still largely unknown but highly plastic and weakly structured by primate phylogeny, compared with the bacteriome. We find significant gut mycobiome overlap between captive apes and human populations living under industrialized subsistence contexts; this is in contrast with contemporary hunter-gatherers and agriculturalists, who share more mycobiome traits with diverse wild-ranging nonhuman primates. In addition, mycobiome-bacteriome interactions were specific to each population, revealing that individual, lifestyle and intrinsic ecological factors affect structural correspondence, number, and kind of interactions between gut bacteria and fungi in primates. Our findings indicate a dominant effect of ecological niche, environmental factors, and diet over the phylogenetic background of the host, in shaping gut mycobiome composition and mycobiome-bacteriome interactions in primates.

Chimpanzees (Pan troglodytes) adapt their nesting behavior after large-scale forest clearance and community decline.

Chimpanzees (Pan troglodytes) build nests at night for sleeping and occasionally during daytime for resting. Over the course of seven years, forest fragments in Bulindi, Uganda, were reduced in size by about 80% when landowners converted forest to agricultural land. However, unlike other studies on nesting behavior in response to habitat disturbance, chimpanzees at Bulindi had no opportunity to retreat into nearby undisturbed forest. To understand behavioral adaptations to forest clearance, we compared Bulindi chimpanzees' nesting characteristics before and after this period of major deforestation. After deforestation, chimpanzees built nests at lower heights in shorter trees, and reused a larger proportion of their nests. Additionally, average nest group size increased after deforestation, even though community size declined by approximately 20% over the same period. The substantial decrease in available forest habitat may have caused the chimpanzees to aggregate for nesting. However, more cohesive nesting may also have been influenced by dietary shifts (increased reliance on agricultural crops) and a need for enhanced safety with increased human encroachment. Conversely, the chimpanzees selected similar tree species for nesting after deforestation, apparently reflecting a strong preference for particular species, nested less often in exotic species, and built integrated nests (constructed using multiple trees) at a similar frequency as before fragment clearance. Chimpanzees living in unprotected habitat in Uganda, as at Bulindi, face mounting anthropogenic pressures that threaten their survival. Nevertheless, our study shows that chimpanzees can adjust their nesting behavior flexibly in response to rapid, extensive habitat change. While behavioral flexibility may enable them to cope with deforestation, at least to a certain point, the long-term survival of chimpanzees in fast-changing human-modified landscapes requires intensive conservation efforts.

Sparse Evidence for Giardia intestinalis, Cryptosporidium spp. and Microsporidia Infections in Humans, Domesticated Animals and Wild Nonhuman Primates Sharing a Farm-Forest Mosaic Landscape in Western Uganda.

Zoonotic pathogen transmission is considered a leading threat to the survival of non-human primates and public health in shared landscapes. Giardia spp., Cryptosporidium spp. and Microsporidia are unicellular parasites spread by the fecal-oral route by environmentally resistant stages and can infect humans, livestock, and wildlife including non-human primates. Using immunoassay diagnostic kits and amplification/sequencing of the region of the triosephosphate isomerase, small ribosomal subunit rRNA and the internal transcribed spacer genes, we investigated Giardia, Cryptosporidium, and microsporidia infections, respectively, among humans, domesticated animals (livestock, poultry, and dogs), and wild nonhuman primates (eastern chimpanzees and black and white colobus monkeys) in Bulindi, Uganda, an area of remarkably high human-animal contact and spatial overlap. We analyzed 137 fecal samples and revealed the presence of G. intestinalis assemblage B in two human isolates, G. intestinalis assemblage E in one cow isolate, and Encephalitozoon cuniculi genotype II in two humans and one goat isolate. None of the chimpanzee and colobus monkey samples were positive for any of the screened parasites. Regular distribution of antiparasitic treatment in both humans and domestic animals in Bulindi could have reduced the occurrence of the screened parasites and decreased potential circulation of these pathogens among host species.

Notes on Morphology and Life History of Probstmayria gombensis (Nematoda: Cosmocercoidea: Atractidae), Parasitic in Eastern Chimpanzees, Pan troglodytes schweinfurthii, in Bulindi, Uganda.

Probstmayria gombensis File, 1976 (Nematoda: Cosmocercoidea: Atractidae) individuals discharged in the feces of eastern chimpanzees, Pan troglodytes schweinfurthii, in Bulindi, Uganda, were examined morphologically. Adults and fourth-stage larvae, all females, found in the feces, and the third-stage larvae excised from the uterus of the gravid females were described. By close observation of the molting worms, it was considered that the uterine third-stage larvae attain molting phase, and then are laid to become fourth-stage larvae. Nutrients required for larval development in the uterus seem to be supplied by the mother after the eggshell is formed. After some growth in the host intestine, the fourth-stage larvae undergo the final molt to the adult stage. The genital primordium was very small in the early fourth-stage larvae but rapidly developed with embryonation in the pre-molt and molting phases. Such precocity suggests parthenogenetic reproduction without insemination by males. This style may enhance rapid autoinfection in the host intestine under the condition of male worm scarcity. Several ellipsoidal pseudocoelomocytes with granules of unknown function were found ventral to the intestine of the adults, fourth-stage larvae, and third-stage larvae.

Atypically High Reproductive Skew in a Small Wild Chimpanzee Community in a Human-Dominated Landscape.

Social rank is positively correlated with reproductive success in numerous species, albeit demographic factors often influence those patterns. In multimale primate species, reproductive skew tends to decrease with increasing numbers of males and sexually receptive females. Alpha male chimpanzees (Pan troglodytes) often sire a disproportionate, though somewhat variable, percentage of offspring compared to other males. In a small community of eastern chimpanzees inhabiting a human-dominated landscape in Bulindi, Uganda, we found extraordinarily high levels of alpha male reproductive success over a 5-year period (7/8 offspring = 88%), despite the presence of multiple subordinate males. The skew exceeds that reported in other studies of chimpanzees as well as closely related bonobos (Pan paniscus). Our findings underscore the role of demographic and social factors in male reproductive success and also suggest that conclusions about species differences may be premature. The interaction of small community size, dispersal limitations, and male reproductive strategies like those found here may decrease genetic diversity and increase the risk of concomitant inbreeding in chimpanzee communities under strong anthropogenic pressure.

Forest fragments become farmland: Dietary Response of wild chimpanzees (Pan troglodytes) to fast-changing anthropogenic landscapes.

Behavioral flexibility, including an ability to modify feeding behavior, is a key trait enabling primates to survive in forest fragments. In human-dominated landscapes, unprotected forest fragments can become progressively degraded, and may be cleared entirely, challenging the capacity of primates to adjust to the changes. We examined responses of wild chimpanzees (Pan troglodytes schweinfurthii) to major habitat change: that is, clearance of forest fragments for agriculture. Over 7 years, fragments in Bulindi, Uganda, were reduced in size by 80%. We compared the chimpanzees' diet at the start and end of this period of rapid deforestation, using data derived mainly from fecal analysis. Similar to other long-term study populations, chimpanzees in Bulindi have a diverse diet comprising over 169 plant foods. However, extensive deforestation seemed to impact their feeding ecology. Dietary changes after fragment clearance included reduced overall frugivory, reduced intake of figs (Ficus spp.; formerly a dietary "staple" for these chimpanzees), and reduced variety of fruits in fecal samples. Nevertheless, the magnitude of most changes was remarkably minor given the extent of forest loss. Agricultural fruits increased in dietary importance, with crops accounting for a greater proportion of fruits in fecal samples after deforestation. In particular, cultivated jackfruit (Artocarpus heterophyllus) became a "staple" food for the chimpanzees but was scarcely eaten before fragment clearance. Crops offer some nutritional benefits for primates, being high in carbohydrate energy and low in hard-to-digest fiber. Thus, crop feeding may have offset foraging costs associated with loss of wild foods and reduced overall frugivory for the chimpanzees. The adaptability of many primates offers hope for their conservation in fragmented, rural landscapes. However, long-term data are needed to establish whether potential benefits (i.e. energetic, reproductive) of foraging in agricultural matrix habitats outweigh fitness costs from anthropogenic mortality risk for chimpanzees and other adaptable primates.

Infant carrying by a wild chimpanzee father at Bulindi, Uganda.

Although infanticide by wild adult male chimpanzees has been reported from multiple sites, affiliative infant carrying by males is rare. We observed infant carrying by an alpha male chimpanzee at Bulindi (Uganda) on two consecutive mornings and collected faecal samples from the newborn infant female, her mother and all candidate fathers to determine whether the alpha male was the infant's father using a likelihood-based method of paternity assignment. In contrast to previous observations of male care of orphans, in this case the mother was present during observations. Further, unlike reports of male aggression towards infants, the infant was reunited with her mother on the third morning, and survived. Neither mother nor infant presented visible injuries. The alpha male never directed aggression towards the infant. Rather, he displayed attentive behaviours, for example by holding the infant to his chest, supporting her while moving, grooming her, and 'cuddling' and 'rocking' her. Paternity results revealed with a high degree of certainty that the alpha male was the infant's father. There are several alternative explanations for the male's behaviour, but this unusual case also highlights the need for further studies to determine under what circumstances adult male chimpanzees can recognise their own offspring.

Breaking Through Disciplinary Barriers: Human-Wildlife Interactions and Multispecies Ethnography.

One of the main challenges when integrating biological and social perspectives in primatology is overcoming interdisciplinary barriers. Unfamiliarity with subject-specific theory and language, distinct disciplinary-bound approaches to research, and academic boundaries aimed at "preserving the integrity" of subject disciplines can hinder developments in interdisciplinary research. With growing interest in how humans and other primates share landscapes, and recognition of the importance of combining biological and social information to do this effectively, the disparate use of terminology is becoming more evident. To tackle this problem, we dissect the meaning of what the biological sciences term studies in "human-wildlife conflict" or more recently "human-wildlife interactions" and compare it to what anthropology terms "multispecies ethnography." In the biological sciences, human-wildlife interactions are the actions resulting from people and wild animals sharing landscapes and resources, with outcomes ranging from being beneficial or harmful to one or both species. In the social sciences, human-nonhuman relationships have been explored on a philosophical, analytical, and empirical level. Building on previous work, we advocate viewing landscapes through an interdisciplinary "multispecies lens" in which humans are observed as one of multiple organisms that interact with other species to shape and create environments. To illustrate these interconnections we use the case study of coexistence between people of the Nalu ethnic group and Critically Endangered western chimpanzees (Pan troglodytes verus) at Cantanhez National Park in Guinea-Bissau, to demonstrate how biological and social research approaches can be complementary and can inform conservation initiatives at the human-primate interface. Finally, we discuss how combining perspectives from ethnoprimatology with those from multispecies ethnography can advance the study of ethnoprimatology to aid productive discourse and enhance future interdisciplinary research.

Tie one on: 'nest tying' by wild chimpanzees at Bulindi-a variant of a universal great ape behavior?

With data accumulating from a growing pool of chimpanzee field studies, new behaviors as well as novel variants on common behaviors continue to be described. Nest construction is a universal behavior in wild great apes. Among chimpanzee populations, reported variation in nest building behavior mostly reflects environmental constraints. Despite the ubiquity of nest making by chimpanzees, only ground nesting has been recognized as a behavioral variant, potentially determined by both environmental and social factors. In a study of nests made by chimpanzees (Pan troglodytes schweinfurthii) in Bulindi, Uganda, I identified a hitherto undescribed nest construction technique, termed 'nest tying'. Five observed nests lacked strong weight-bearing structures beneath them, such as large branches or a supporting trunk. Instead, the nests appeared 'tied' (or 'tethered') to an adjacent trunk by looping leafy stems or palm fronds around it and interweaving these into the nest mattress, securing the nest against the trunk; thus, nest tying presumably functions to provide added stability and support. This preliminary report presents a description of the observed nests. Irrespective of whether nest tying constitutes true knot making-commonly considered absent in wild great apes-this nest construction technique would seem to require advanced dexterity and a sophisticated understanding of the mechanical properties of the plants used. Forest fragments in Bulindi are highly degraded. Thus, nest tying-and construction of integrated nests (i.e., utilizing multiple plants, often small trees and shrubs) generally-may be promoted by a relative paucity of suitable nesting trees at this site. Still, insofar as nest building is learned in chimpanzees, different construction techniques including nest tying are potentially acquired through social learning. Further investigation is required to ascertain the prevalence and acquisition of this nest construction technique at Bulindi, and to verify its presence or absence in other habitats.

Zoonotic Enterobacterial Pathogens Detected in Wild Chimpanzees.

Infectious diseases including those acquired through direct or indirect contact with people and livestock threaten the survival of wild great apes. Few studies have reported enterobacterial pathogens in chimpanzees. We used multiplex PCR to screen faeces of chimpanzees sharing a landscape with villagers and livestock in Bulindi, Uganda for Salmonella spp., enterohemorrhagic Escherichia coli (E. coli) and Shigella spp./enteroinvasive E. coli. All three potentially zoonotic pathogens were detected. Individual prevalence ranged between 7 and 20%, with most infections observed in mature male chimpanzees. These preliminary findings suggest detailed investigation of enterobacterial infections in people, primates and livestock in this ecosystem is warranted.

Low Levels of Fruit Nitrogen as Drivers for the Evolution of Madagascar's Primate Communities.

The uneven representation of frugivorous mammals and birds across tropical regions - high in the New World, low in Madagascar and intermediate in Africa and Asia - represents a long-standing enigma in ecology. Several hypotheses have been proposed to explain these differences but the ultimate drivers remain unclear. Here, we tested the hypothesis that fruits in Madagascar contain insufficient nitrogen to meet primate metabolic requirements, thus constraining the evolution of frugivory. We performed a global analysis of nitrogen in fruits consumed by primates, as collated from 79 studies. Our results showed that average frugivory among lemur communities was lower compared to New World and Asian-African primate communities. Fruits in Madagascar contain lower average nitrogen than those in the New World and Old World. Nitrogen content in the overall diets of primate species did not differ significantly between major taxonomic radiations. There is no relationship between fruit protein and the degree of frugivory among primates either globally or within regions, with the exception of Madagascar. This suggests that low protein availability in fruits influences current lemur communities to select for protein from other sources, whereas in the New World and Old World other factors are more significant in shaping primate communities.

Gastrointestinal parasite infections and self-medication in wild chimpanzees surviving in degraded forest fragments within an agricultural landscape mosaic in Uganda.

Monitoring health in wild great apes is integral to their conservation and is especially important where they share habitats with humans, given the potential for zoonotic pathogen exchange. We studied the intestinal parasites of wild chimpanzees (Pan troglodytes schweinfurthii) inhabiting degraded forest fragments amid farmland and villages in Bulindi, Uganda. We first identified protozoan and helminth parasites infecting this population. Sixteen taxa were demonstrated microscopically (9 protozoa, 5 nematodes, 1 cestode, and 1 trematode). DNA sequence analysis enabled more precise identification of larval nematodes (e.g. Oesophagostomum stephanostomum, O. bifurcum, Strongyloides fuelleborni, Necator sp. Type II) and tapeworm proglottids (genus Bertiella). To better understand the ecology of infections, we used multidimensional scaling analysis to reveal general patterns of association among parasites, climate, and whole leaf swallowing-a prevalent self-medicative behaviour at Bulindi linked to control of nodular worms (Oesophagostomum spp.). Prevalence of parasites varied with climate in diverse ways. For example, Oesophagostomum sp. was detected in faeces at higher frequencies with increasing rainfall but was most clearly associated with periods of low temperature. Certain parasites occurred together within chimpanzee hosts more or less frequently than expected by chance. For example, the commensal ciliate Troglodytella abrassarti was negatively associated with Balantidium coli and Oesophagostomum sp., possibly because the latter taxa make the large intestine less suitable for T. abrassarti. Whole leaves in faeces showed independent associations with the prevalence of Oesophagostomum sp., Strongyloides sp., and hookworm by microscopic examination, and with egestion of adult O. stephanostomum by macroscopic inspection. All parasites identified to species or genus have been reported in wild chimpanzees inhabiting less-disturbed environments than Bulindi. Nevertheless, several disease-causing taxa infecting these chimpanzees are potentially transmissible between apes and humans (e.g. rhabditoid and strongyle nematodes), underscoring the importance of identifying and reducing risks of pathogen exchange in shared landscapes.

Molecular features of hookworm larvae (Necator spp.) raised by coproculture from Ugandan chimpanzees and Gabonese gorillas and humans.

Species composition of Necator hookworms was surveyed in (i) Ugandan chimpanzees living around farms and villages at Bulindi, (ii) Gabonese gorillas under habituation in Moukalaba-Doudou National Park (MDNP), and (iii) Gabonese villagers living adjacent to MDNP. Internal transcribed spacers (ITS) of rDNA and partial cytochrome c oxidase subunit 1 (Cox1) gene of mtDNA were analyzed from larvae obtained by coproculture. Three ITS types (I, II and III) and three Cox1 haplotype groups (A, B and C) were demonstrated. ITS type I and Cox1 haplotype group A, representing Necator americanus, were demonstrated in the hookworm larvae from Gabonese gorillas and humans, but not from Ugandan chimpanzees. Type II and haplotype groups B and C, presumably representing N. gorillae, were found in larvae from Ugandan chimpanzees and Gabonese gorillas and humans. These features were overall similar with those found previously in the Central African Republic. Meanwhile, type III was proven in a larva from a Gabonese gorilla as the first demonstration from a non-human primate. Cox1 haplotypes obtained from Ugandan chimpanzees formed a subgroup within group B, presumably reflecting dispersal and diversification processes of the apes.

Strongyloides infections of humans and great apes in Dzanga-Sangha Protected Areas, Central African Republic and in degraded forest fragments in Bulindi, Uganda.

DNA sequence analysis was carried out on Strongyloides spp. larvae obtained from fecal samples of local humans, a wild western lowland gorilla (Gorilla gorilla gorilla) and a central chimpanzee (Pan troglodytes troglodytes) inhabiting Dzanga-Sangha Protected Areas (DSPA), Central African Republic, and eastern chimpanzees (Pan troglodytes schweinfurthii) living in degraded forest fragments on farmland in Bulindi, Uganda. From humans, both Strongyloides fuelleborni and Strongyloides stercoralis were recorded, though the former was predominant. Only S. fuelleborni was present in the great apes in both areas. Phylogenetic analysis of partial mtDNA cytochrome c oxidase subunit 1 gene (Cox1) and comparison of 18S rDNA hyper variable region IV (HVR-IV) sequences implied that in DSPA S. fuelleborni populations in humans differ from those in the nonhuman great apes.

Cars kill chimpanzees: case report of a wild chimpanzee killed on a road at Bulindi, Uganda.

Roads have broadly adverse impacts on wildlife, including nonhuman primates. One direct effect is mortality from collisions with vehicles. While highly undesirable, roadkills provide valuable information on the health and condition of endangered species. We present a case report of a wild chimpanzee (Pan troglodytes schweinfurthii) killed crossing a road in Bulindi, Uganda, where chimpanzees inhabit forest fragments amid farmland. Details of the collision are constructed from eyewitness accounts of pedestrians. Physical examination of the cadaver indicated good overall body condition; at 40 kg, the deceased female was heavier than usual for an adult female East African chimpanzee. No external wounds or fractures were noted. Coprological assessment demonstrated infection by several gastrointestinal parasites commonly reported in living wild chimpanzees. Histopathology revealed eosinophilic enteritis and biliary hyperplasia potentially caused by parasite infection. However, eosinophilia was not widely spread into the submucosa, while egg/cyst counts suggested low-intensity parasite infections compared to healthy female chimpanzees of similar age in nearby Budongo Forest. No behavioral indicators of ill health were noted in the deceased female in the month prior to the accident. We conclude that cause of death was acute, i.e., shock from the collision, and was probably unrelated to parasite infection or any other underlying health condition. Notably, this female had asymmetrical polythelia, and, while nursing at the time of her death, had one functioning mammary gland only. In Uganda, where primates often inhabit human-dominated landscapes, human population growth and economic development has given rise to increasing motor traffic, while road development is enabling motorists to travel at greater speeds. Thus, the danger of roads to apes and other wildlife is rising, necessitating urgent strategies to reduce risks. Installation of simple speed-bumps-common on Ugandan roads-would be effective in reducing risks to wildlife, and would also make roads safer for human pedestrians.

Tailored Enrichment Strategies and Stereotypic Behavior in Captive Individually Housed Macaques (Macaca spp.).

The welfare of nonhuman animals in captivity is widely dependent on the natural psychological, physical, and behavioral needs of the animals and how adequately these needs are met. Inability to engage in natural behaviors can lead to chronic stress and expression of stereotypic behavior. The majority of research on decreasing stereotypic behavior in captivity addresses problems at the group level and does not account for individual variability in each animal's needs, history, and preferences. This study combined physiological and behavioral measures of well being to comprehensively assess the unique needs of individually housed captive macaques (Macaca spp.) with the aim of developing tailored welfare strategies. Behavioral and hormonal data were collected under 2 conditions: baseline and individualized enrichment. The results showed a significant decrease in stereotypic behavior under the enrichment condition. Additionally, 7 out of 9 individuals showed a decrease in fecal glucocorticoid (stress hormone) levels, indicating a reduction in hypothalamic-pituitary-adrenal activity. Addressing welfare on an individual, rather than group, level allows for a better overall characterization of well being and maximizes the probability of improving the welfare of each animal.

Molecular identification of Oesophagostomum spp. from 'village' chimpanzees in Uganda and their phylogenetic relationship with those of other primates.

Oesophagostomum spp. are parasitic nematodes of mammals, including humans and other primates. To identify species and determine phylogeny, we analysed DNA sequences of adult and larval Oesophagostomum from wild chimpanzees in Bulindi, Uganda, which inhabit degraded forest fragments amid villages. Oesophagostome larvae and/or eggs from baboons in Tanzania and South Africa and from a Japanese macaque were also sequenced. Based on the internal transcribed spacer 2 (ITS2) of nuclear ribosomal DNA and partial cytochrome c oxidase subunit 1 gene (Cox1) of mtDNA, O. stephanostomum and O. bifurcum were identified from chimpanzees. Bulindi is the second locality where molecular detection of O. bifurcum in wild chimpanzees has been made. While most O. stephanostomum had ITS2 genotypes recorded previously, three new genotypes were detected. Among four ITS2 genotypes of O. bifurcum from chimpanzees, one was identical to that from various monkey species in Kibale, Uganda, and baboons from Tanzania and South Africa; another was shared by a baboon from Tanzania. No genotype was identical with that of the cryptic species reported from humans and monkeys in Kibale. Phylogeny based on Cox1 sequences of O. stephanostomum showed locality-dependent clades, whereas those of O. bifurcum formed clades composed of worms from different hosts and localities.

Is honey a fallback food for wild chimpanzees or just a sweet treat?

OBJECTIVES: Honey is a highly nutritious resource for any primate able to exploit it. Wild chimpanzees exploit nests of honey-making bees (Apini and Meliponini) for honey and brood, typically using tools to overcome the bees' defences. The universality of honey-gathering among modern human foragers in tropical climates and chimpanzees suggests energy-rich honey, acquired with tools, was likely a regular food for ancestral hominins. However, few studies have assessed its role in seasonal foraging strategies of chimpanzees. This study asks whether honey serves as a high-quality fallback food for chimpanzees at Bulindi, Uganda. MATERIALS AND METHODS: Honey consumption was investigated via fecal analysis over 22 months during two studies (Study 1: 2007-2008; Study 2: 2012-2014). Additionally, flower and fruit phenology was measured during Study 1; peak flowering intensity was expected to facilitate increased honey and/or brood production by bees. RESULTS: Chimpanzees consumed honey (and/or brood) at low frequencies year-round, but bees/beeswax appeared in feces at higher frequencies with decreasing fruit availability (Study 1). Honey consumption was unrelated to flowering and chimpanzees did not consume honey more frequently during the "honey season" when local people harvest beehives. Moreover, consumption was inversely related to fruit intake (both study periods). DISCUSSION: Although honey fits the functional definition of a filler fallback food at Bulindi, the chimpanzees unlikely depend on honey to replace nutrients provided by fruit. Overall, honey best qualifies as an energy-dense "treat" during low fruiting months. The data lend support to the hypothesis that tools can facilitate chimpanzees' access to high-quality fallbacks including insect foods when fruit availability is low.