A survey of nocturnality and risk for savanna chimpanzees at Assirik, Senegal.

Chimpanzees (Pan troglodytes) in Senegal may use nocturnality to mitigate hyperthermia risk in semi-arid environments but the degree of nocturnality for such chimpanzees also in sympatry with large carnivores remains uncertain. We compared diel activity among chimpanzees and their potential predators at Assirik in Niokolo-Koba National Park and contextualized these findings relative to other unit-groups in savanna landscapes. From 2015-2018, we generated a predator inventory using multi-modal methods and monitored the diel activity of chimpanzees and predators with camera traps [ N = 2092 camera trap (CT) days]. From 2015-2023, we also surveyed for evidence of predation during recce walks. Six potential nonhuman predators occur at Assirik, including lions (Panthera leo), leopards (Panthera pardus), spotted hyenas (Crocuta crocuta), African wild dogs (Lycaon pictus), Nile crocodiles (Crocodylus niloticus), and rock pythons (Python sebae). We documented one suspected case of a predator killing a chimpanzee. Nocturnality comprised 12.7% of CT events for chimpanzees and these events were more concentrated at twilight. Chimpanzees were more active during the day, predators were more active at night, and there was substantial temporal overlap among chimpanzees and potential predators during twilight intervals. Our findings support the hypothesis that savanna chimpanzees in Senegal are active at night in response to the extremely hot environment. We hypothesize that Assirik chimpanzees experience a tension between decreasing hyperthermia and increasing predation risk during nocturnality.

Chimpanzees (Pan troglodytes) in savanna landscapes.

Chimpanzees (Pan troglodytes) are the only great apes that inhabit hot, dry, and open savannas. We review the environmental pressures of savannas on chimpanzees, such as food and water scarcity, and the evidence for chimpanzees' behavioral responses to these landscapes. In our analysis, savannas were generally associated with low chimpanzee population densities and large home ranges. In addition, thermoregulatory behaviors that likely reduce hyperthermia risk, such as cave use, were frequently observed in the hottest and driest savanna landscapes. We hypothesize that such responses are evidence of a "savanna landscape effect" in chimpanzees and offer pathways for future research to understand its evolutionary processes and mechanisms. We conclude by discussing the significance of research on savanna chimpanzees to modeling the evolution of early hominin traits and informing conservation programs for these endangered apes.

Western chimpanzees (Pan troglodytes verus) access a nutritionally balanced, high energy, and abundant food, baobab (Adansonia digitata) fruit, with extractive foraging and reingestion.

Intrinsic to several hypotheses explaining the evolution of foraging behavior complexity, such as proto-tool use, is the assumption that more complex ingestive behaviors are adaptations allowing individuals to access difficult to procure but nutritionally or energetically rewarding foods. However, nutritional approaches to understanding this complexity have been underutilized. The goal of this study was to evaluate potential nutritional determinants of two unusual foraging behaviors, fruit cracking with anvils and seed reingestion, by adult male western chimpanzees (Pan troglodytes verus) at Fongoli, Senegal during the baobab (Adansonia digitata) fruit season. We examined these behaviors in relation to nutrient and energy intake, and compared macronutrient and energy concentrations found in baobab fruits to other plant foods. Adult males ingested at least 31 distinct foods from 23 plant species. Baobab fruit comprised the majority of daily energy intake (68 ± 34%, range: 0%-98%). The energetic concentration of baobab fruit varied by phenophase and part ingested, with ripe and semi-ripe fruit ranking high in energy return rate. Males preferred ripe and semi-ripe baobab fruit but unripe fruit intake was higher overall. The seed kernels were high in protein and fat relative to fruit pulp, and these kernels were easier to access during the unripe stage. During the ripe stage, seed kernels were accessible by reingestion, after the seed coat was softened during gut passage. In addition to providing macronutrients and energy, baobab fruit was a relatively abundant food source. We conclude that baobab pulp and seed are high quality foods at Fongoli during the baobab season because they are nutritionally balanced, high in energy, and relatively abundant in the environment. These nutritional and abundance characteristics may explain, in part, why these chimpanzees use anvils and reingestion to access a mechanically challenging food.

Informing Protection Efforts for Critically Endangered Chimpanzees (Pan troglodytes verus) and Sympatric Mammals amidst Rapid Growth of Extractive Industries in Senegal.

Updated information on Critically Endangered western chimpanzees (Pan troglodytes verus) in Senegal is urgently needed, given that gold mining is rapidly transforming landscapes and livelihoods. Specifically, biodiversity assessments will better elucidate the chimpanzee extinction risk here and yield baselines for monitoring. We compared mammal species richness between Fongoli (unprotected) and Assirik in Niokolo-Koba National Park to assess the efficacy of the only nationally protected area where chimpanzees range in this country. The primary habitat difference between these sites was the degree of human activity. Although Assirik and Fongoli had similar mammal assemblages and were equivalent in primate species richness, the protected area was higher in species richness overall, particularly for ungulates and carnivores. The protected status and management plan of Niokolo-Koba almost certainly resulted in fewer mammal extinctions. In unprotected areas, the flexible behavioural responses of chimpanzees to human-driven landscape dynamics is likely essential to their long-term survival. Furthermore, the near absence of chimpanzee killings in this country greatly facilitates conservation efforts. Given that mineral extraction is a key development strategy in Senegal, we encourage primatologists to collaborate with mining stakeholders to ensure that sustainability programming includes contributions from experts in savannah chimpanzee behavior and ecology.

Nest-Building Behaviour by Chimpanzees (Pan troglodytes verus) in the Non-Protected Area of Diaguiri (Kedougou, Senegal): Implications for Conservation.

Diaguiri is a non-protected area situated in south-eastern Senegal. Chimpanzees (Pan troglodytes verus) have been living here since 1999 but information on the characteristics of their nest sites have been lacking until now. Furthermore, we implemented nest survey methods recently standardised to arid landscapes during a national chimpanzee survey. Variations in nest decay rates are poorly understood in savanna-woodland mosaic habitats. This study aims to describe in Diaguiri: (1) the distribution of chimpanzee nests and the location of their core sleeping area, (2) the tree species used for nest building, (3) the association between nests and nesting tree heights, and (4) differences in nest decay rates between habitats. We surveyed 43 km of chimpanzee nesting habitats and recorded 871 nests in gallery forest and woodland habitats between March 2015 and June 2016. Diaguiri chimpanzees used at least 17 tree species for nest building, and 58.8% of nests were in only 3 tree species: Pterocarpus erinaceus, Anogeissus leiocarpus and Diospyros mespiliformis. Nest decay rate was faster in gallery forests than in woodlands. These data are of great value for understanding habitat use by a Critically Endangered subspecies of savanna chimpanzee, for population density estimates of the species and for conservation programmes in this region and in savanna biomes.

Feeding in fear? How adult male western chimpanzees (Pan troglodytes verus) adjust to predation and savanna habitat pressures.

OBJECTIVES: We evaluated risk-sensitive foraging in adult male western chimpanzees (Pan troglodytes verus) occupying a savanna environment at Fongoli, Senegal. The aim of this study was to determine how the risks of predation and heat stress influenced their behavior while feeding on a key food, fruit of the baobab tree (Adansonia digitata). MATERIALS AND METHODS: Proximity of fruiting baobab trees to anthropogenic landmarks were compared to food intake, feeding rate, and behavioral indicators of fear in adult males (N = 11) at Fongoli. Additionally, we compared foraging to vegetative habitats, baobab ripe fruit nutritive quality, surface water availability, and foraging party composition. RESULTS: Fruit abundance increased with proximity to anthropogenic landmarks, and chimpanzees exhibited higher frequencies of antipredator behaviors as they approached these risky areas. However, predation risk did not deter adult males from visiting these fruiting trees; instead, risky foraging bouts were associated with higher food intakes and longer feeding times. Additionally, higher feeding rates were observed in open-canopy habitats, and this behavior may have minimized their risk of heat stress. CONCLUSIONS: Adaptations that minimize predation risk are widespread in mammalian prey species, but these traits are poorly understood in chimpanzees. Great apes encounter few nonhuman predators capable of successfully capturing and killing them; thus, such events are rarely observed. Although people rarely hunt chimpanzees in Senegal, we found that adult males perceived humans as predators and adjusted their behavior while foraging in risky habitats. From an applied perspective, risk-taking behavior is important for understanding and mitigating the problem of crop-feeding in locations where chimpanzees and humans live in sympatry.

Updating Chimpanzee Nesting Data at Mount Assirik (Niokolo Koba National Park, Senegal): Implications for Conservation.

The Niokolo Koba National Park (NKNP) is the largest protected area in Senegal and lies at the northern limit of the chimpanzee's range in West Africa. Recent information on nesting behavior and factors influencing nesting behavior is available for several sites outside NKNP. However, the information available for NKNP is obsolete. Considering that the adequate management of chimpanzee populations cannot be achieved without strong scientific knowledge, it is essential to update data on chimpanzee, Pan troglodytes verus, nesting behavior in NKNP. For this reason, we surveyed their habitat in Mt. Assirik and recorded 626 chimpanzee nests. The results of the study showed that chimpanzees nest more often in closed-canopy habitats such as gallery forests. The average nest height observed in this study was 8.07 ± 0.36 m, varying between 2 and 20 m, which is well below the heights described in most sites where chimpanzees cohabit with large carnivores. Botanical surveys confirmed that chimpanzees select tree species bearing their nests. In Assirik, 12 of the 37 tree species bearing nests are the most used. The nest decay rate (or the time it takes for a nest to go from the fresh to the skeletal stage) at Assirik averaged 68.8 ± 5.8 days.

Reciprocity and beyond: Explaining meat transfers in savanna-dwelling chimpanzees at Fongoli, Senegal.

OBJECTIVES: To understand the function of food sharing among our early hominin ancestors, we can turn to our nonhuman primate relatives for insight. Here, we examined the function of meat sharing by Fongoli chimpanzees, a community of western chimpanzees (Pan troglodytes verus) in southeastern Sénégal. MATERIALS AND METHODS: We tested three non-mutually exclusive hypotheses that have been used to explain patterns of food sharing: kin selection, generalized reciprocity, and meat-for-mating opportunities. We analyzed meat sharing events (n = 484) resulting from hunts, along with data on copulations, age-sex class, and kinship to determine which variables predict the likelihood of meat sharing during this study period (2006-2019). RESULTS: We found full or partial support for kin selection, direct reciprocity, and meat-for-mating-opportunities. However, the analyses reveal that reciprocity and a mother/offspring relationship were the strongest predictors of whether or not an individual shared meat. CONCLUSIONS: The results of this study emphasize the complexity of chimpanzee meat sharing behaviors, especially at a site where social tolerance offers increased opportunities for meat sharing by individuals other than dominant males. These findings can be placed in a referential model to inform hypotheses about the sensitivity of food sharing to environmental pressures, such as resource scarcity in savanna landscapes.

Gastrointestinal parasites of baboons (Papio papio) in Niokolo-Koba National Park, Senegal.

Background: Primates can harbor parasites that could be pathogenic or not for humans and primates themselves. It is necessary to know the parasitological situation of the primates that are under surveillance in the park. Aim: To estimate the prevalence and diversity of gastrointestinal parasites, including zoonotic potential parasites, in baboons in the Niokolo-Koba National Park located in Senegal. Method: Fecal samples (n = 50) from two groups of baboons (A and B) were collected in October 2019. The samples were processed using the flotation technique and the modified Ritchie method. Slides were examined microscopically and the parasite identification was based on morphology, color, and parasite content. Results: A total of seven nematodes (Strongyloides sp., Trichirus sp., Ancylostoma sp., Mammo monogamus, Enterobius sp., Strongyloides stercoralis, Strongyle digestif), one cestode (Bertiella sp.), and one trematode (Fasciolopsis sp.) were identified. The overall prevalence was 78%, while the prevalence of poly-infected samples was 49%. The parasite with zoonotic potential, S. stercoralis, was identified in group B samples. Trichuris sp., which is common and pathogenic to humans and primates, was present with prevalence of 52% and of 32% in groups A and B, respectively. Conclusion: These results suggest that baboons are infested with zoonotic parasites and this situation could expose people working in this park to infection. Contact between humans and these baboons or their feces could expose them to infection with zoonotic parasites.

Tool use in wild spider monkeys (Ateles geoffroyi).

Tool use has been observed in a variety of primate species, including both New and Old World monkeys. However, such reports mainly address the most prodigious tool users and frequently limit discussions of tool-using behavior to a foraging framework. Here, we present observations of novel and spontaneous tool use in wild black-handed spider monkeys (Ateles geoffroyi), where female spider monkeys used detached sticks in a self-directed manner. We introduce factors to explain Ateles tool-using abilities and limitations, and encourage the synthesis of relevant research in order to gain insight into the cognitive abilities of spider monkeys and the evolution of tool-using behaviors in primates.

Plant-food and tool transfer among savanna chimpanzees at Fongoli, Senegal.

Transferring food is considered a defining characteristic of humans, as such behavior is relatively uncommon in other animal species save for kin-based transfer. Chimpanzees (Pan troglodytes) are one exception, as they commonly transfer meat among nonrelatives but rarely transfer other resources. New observations at Fongoli, Senegal, show habitual transfer of wild-plant foods and other non-meat resources among community members beyond transfers from mother to offspring. We explore various explanations for these behaviors with a focus on age- and sex-class patterns in transfer events. In a total of 27 of 41 cases, male chimpanzees at Fongoli transferred wild-plant foods or tools to females. Most other cases involved transfer among males or males taking food from females. In light of male-female transfer patterns at Fongoli, we examine four hypotheses that have been applied to food transfer in apes: (1) testing for male-coercive tendency (van Noordwijk and van Schaik, Behav Ecol Sociobiol 63:883-890, 2009), (2) costly signaling (Hockings et al. PLoS ONE 2:e886, 2007), (3) food-for-sex (Gomes and Boesch, PLoS ONE 4:5116, 2009), and (4) sharing-under-pressure (Gilby, Anim Behav 71:953-963, 2006). We also consider hypotheses posed to explain transfer among callitrichids, where such behavior is more common (Ruiz-Miranda et al. Am J Primatol 48:305-320, 1999). Finally, we examine variables such as patch and food size and food transport. We discuss our findings relative to general patterns of non-meat transfer in Pan and examine them in the context of chimpanzee sociality in particular. We then contrast chimpanzee species and subspecies in terms of non-meat food and tool transfer and address the possibility that a savanna environment contributes to the unusual pattern observed at Fongoli.