The archaeological visibility of chimpanzee (Pan troglodytes) nut-cracking.

The earliest evidence for complex tool use in the archaeological record dates to 3.3 Ma. While wooden tools may have been used by our earliest ancestors, the evidence is absent due to poor preservation. However, insights into possible early hominin wooden tool use can be gained from observing the tool-use practices of our closest living relatives, chimpanzees (Pan troglodytes). By using stone hammers used to crack various nuts, chimpanzees leave a durable material signature comprised of formal tools and associated diagnostic fragments. While the archaeological evidence of chimpanzee wooden tool use is temporary, the combination of stone hammers and wooden anvils can create a more enduring lithic record. This study explores the lithic assemblages associated with wooden and stone anvil use at nut-cracking sites in Taï National Park, Côte d'Ivoire, using technological and use-wear analyses. Our results indicate clear differences in density, fracture patterns, and use-wear in the lithic records between wooden anvil and stone anvil sites. New archaeological excavations at six chimpanzee nut-cracking sites reveal that the anvils' material directly influences the visibility of nut-cracking evidence in the archaeological record. By examining the nature of the lithic signatures associated with wooden anvil and stone anvil use by chimpanzees, we can formulate hypotheses about the probability of such behaviors being preserved and identifiable in the Plio-Pleistocene hominin archaeological record. The variability in material signatures from nut-cracking on different anvils suggests that stone anvils leave a clear archaeological record. Evidence for wooden anvil use is likely underrepresented due to the more ephemeral nature of the associated percussive damage and material signature. It may, however, still be possible, albeit challenging, to identify wooden anvil use in the archaeological record.

Does Oxygen Stable Isotope Composition in Primates Vary as a Function of Vertical Stratification or Folivorous Behaviour?

Stable isotopes of oxygen often vary within a community of primates. For example, folivorous monkeys that forage in the upper reaches of the forest canopy tend to evince high δ18O values, whereas those that prefer the understory tend to have lower δ18O values. Given that leaves also have high δ18O values, particularly higher in the canopy, there is uncertainty as to which behavioural variable - vertical stratification or folivory - is the primary determinant of variation in δ18O values. Here, we explore further δ18O values from the Taï Forest monkeys (n = 7 species; n = 33 individuals) by examining the interaction between diet and vertical stratification, thereby allowing us to differentiate the effects of each covariate. We found that δ18O values varied as a function of mean canopy height, but not folivory, resolving uncertainty about the primary cause of δ18O variation. This outcome revolves largely, but not entirely, on the behaviours of Procolobus verus, a highly folivorous but understory forager. Relatively elevated values in Cercopithecus diana, a frugivorous but middle-to-high canopy forager, raises the possibility that plant reproductive tissues (e.g., fruits, flowers) may be increasingly sensitive to evaporative fractionation at higher forest canopy levels. Overall, our results further affirm the value of using δ18O values to estimate the vertical behaviour of primate species in a fossil assemblage.

Dietary Variation in Diana Monkeys (Cercopithecus diana): The Effects of Polyspecific Associations.

Establishing dietary central tendencies and extremes remains an important goal of primate research. While habitat differences and spatial discontinuity are well-documented contributors to dietary variation, other factors including polyspecific associations may significantly impact diet through changes in strata use and/or increased feeding competition. Here, we examine polyspecific association with closely related species as a source of dietary variation in a rain forest primate. Diana monkeys (Cercopithecus diana) in Côte d'Ivoire's Taï Forest frequently participate in mixed-species groups. We use data collected over a 5-year period on 4 Diana monkey groups to examine how association affects group diets. Groups exhibited significant differences in association rates with other guenon species, but this minimally influenced diet when food categories (fruit, invertebrates, leaves) were compared: diet overlap of 4 groups across the study period ranges from 90.8 to 98.1%. Examination of species composition within food categories is more revealing: intergroup dietary overlap decreases to 69.8-79.4% across the study period when comparing species contributing to total frugivory and folivory. These data support earlier findings that Diana monkeys maintain fruit-rich diets by competitively excluding sympatric congeners while highlighting that even selective frugivores such as Diana monkeys may exhibit remarkable dietary flexibility. Our study underscores the fact that broad categorical labels can obscure significant dietary differences.

Fatal monkeypox in wild-living sooty mangabey, Côte d'Ivoire, 2012.

We isolated a monkeypox virus from a wild-living monkey, a sooty mangabey, found dead in Taï National Park, Côte d'Ivoire, in March 2012. The whole-genome sequence obtained from this isolate and directly from clinical specimens showed its close relationship to monkeypox viruses from Western Africa.

Variability in molar crown morphology and cusp wear in two Western chimpanzee populations.

OBJECTIVES: Chimpanzees (Pan troglodytes) possess a relatively generalized molar morphology allowing them to access a wide range of foods. Comparisons of crown and cusp morphology among the four subspecies have suggested relatively large intraspecific variability. Here, we compare molar crown traits and cusp wear of two geographically close populations of Western chimpanzees, P. t. verus, to provide further information on intraspecific dental variability. MATERIALS AND METHODS: Micro-CT reconstructions of high-resolution replicas of first and second molars of two Western chimpanzee populations from Ivory Coast (Taï National Park) and Liberia, respectively were used for this study. First, we analyzed projected tooth and cusp 2D areas as well as the occurrence of cusp six (C6) on lower molars. Second, we quantified the molar cusp wear three-dimensionally to infer how the individual cusps alter with advancing wear. RESULTS: Both populations are similar in their molar crown morphology, except for a higher appearance rate of a C6 in Taï chimpanzees. In Taï chimpanzees, lingual cusps of upper molars and buccal cusps of lower molars possess an advanced wear pattern compared to the remaining cusps, while in Liberian chimpanzees this wear gradient is less pronounced. DISCUSSION: The similar crown morphology between both populations fits with previous descriptions for Western chimpanzees and provides additional data on dental variation within this subspecies. The wear pattern of the Taï chimpanzees are in concordance with their observed tool rather than tooth use to open nuts/seeds, while the Liberian chimpanzees may have consumed hard food items crushed between their molars.

Pheidoleklaman sp. nov.: a new addition from Ivory Coast to the Afrotropical pulchella species group (Hymenoptera, Formicidae, Myrmicinae).

In this study the taxonomy of the Pheidolepulchella species group is updated for the Afrotropical region and the new species P.klaman sp. nov. described. It is integrated into the existing taxonomic system by an updated identification key for the whole group and an update of the known distribution ranges of its members. High quality focus stacking images are provided, with X-ray micro-CT scanned digital 3D representations, of major and minor worker type specimens.

Animal reactivity to camera traps and its effects on abundance estimate using distance sampling in the Taï National Park, Côte d'Ivoire.

The use of camera traps (CTs) has become an increasingly popular method of studying wildlife, as CTs are able to detect rare, nocturnal, and elusive species in remote and difficult-to-access areas. It thus makes them suited to estimate animal density and abundance, identify activity patterns and new behaviours of animals. However, animals can react when they see the CTs and this can lead to bias in the animal population estimates. While CTs may provide many advantages, an improved understanding of their impacts on individual's behaviour is necessary to avoid erroneous density estimates. Yet, the impact of CTs on detected individuals, such as human odour near the device and the environment, or the infrared illumination, has received relatively little attention. To date, there is no clear procedure to remove this potential bias. Here, we use camera trap distance sampling (CTDS) to (1) quantify the bias resulting from the different animal responses to the CTs when determining animal density and abundance, and (2) test if olfactory, visual and auditory signals have an influence on the animals' reaction to CTs. Between March 2019 and March 2020, we deployed CTs at 267 locations distributed systematically over the entire Taï National Park. We obtained 58,947 videos from which we analysed four medium- to-large-bodied species (Maxwell's duiker (Philantomba maxwellii), Jentink's duiker (Cephalophus jentinki), pygmy hippopotamus (Choeropsis liberiensis) and Western chimpanzee (Pan troglodytes verus)) displaying different behaviours towards the CTs. We then established species-specific ethograms describing the behavioural responses to the CTs. Using these species-specific responses, we observed that the Maxwell's duiker reacted weakly to CTs (about 0.11% of the distance data), contrary to Jentink's duiker, pygmy hippopotamus and Western chimpanzee which reacted with relatively high frequencies, representing 32.82%, 52.96% and 16.14% of the distance data, respectively. Not taking into account the species-specific responses to the CTs can lead to an artificial doubling or tripling of the populations' sizes. All species reacted more to the CTs at close distances. Besides, the Jentink's duiker and the pygmy hippopotamus reacted significantly more to the CTs at night than during the day. Finally, as for olfactory signals, the probability of reaction to the CTs during the first days after CTs installation was weak in Maxwell's duiker, but concerned 18% of the video captures in Western chimpanzees which decreasing with time, but they remained high in pygmy hippopotamus and Jentink's duiker (65% and 70% of the video captures respectively). Careful consideration should be given to animal's response to CTs during the analysis and in the field, by reducing human's impact around the CTs installation.

Seed choice differs by sex in sooty mangabeys (Cercocebus atys).

The sooty mangabey (Cercocebus atys) practices year-round durophagy. A large part of the C. atys diet consists of the oily nut of Sacoglottis gabonensis, which is accessed by post-canine crushing of the hard, protective seed coat. During a typical foraging bout, some seeds are discarded after initial crushing attempts using isometric biting, but the reason mangabeys reject some seeds and break into others is unclear. Although C. atys is sexually dimorphic, little is known about whether differences between males and females affect the selectivity of mechanically protected foods. We studied C. atys feeding on S. gabonensis in the Taï National Park,  Côte d'Ivoire, in July and August 2016. Nuts discarded after an initial crushing attempt were collected and their hardness measured using a Shore D durometer. Measurements were taken in the region of the nut where monkeys attempted to crush it. Hardness values of nuts rejected by adult male (n = 79) and adult female (n = 104) C. atys were compared to those of a control assemblage of nuts collected randomly on the forest floor (n = 69). Nuts rejected by either sex do not differ statistically from the random sample; however, they do differ from each other, with females rejecting harder nuts. This suggests that males are more effective at broaching harder seed husks, and discard seeds based on other factors.

Social rank overrides environmental and community fluctuations in determining meat access by female chimpanzees in the Taï National Park, Côte d'Ivoire.

Meat, long hypothesized as an important food source in human evolution, is still a substantial component of the modern human diet, with some humans relying entirely on meat during certain times of the year. Understanding the socio-ecological context leading to the successful acquisition and consumption of meat by chimpanzees (Pan troglodytes), our closest living relative, can provide insight into the emergence of this trait because humans and chimpanzees are unusual among primates in that they both (i) hunt mammalian prey, (ii) share meat with community members, and (iii) form long-term relationships and complex social hierarchies within their communities. However, females in both human hunter-gatherer societies as well as chimpanzee groups rarely hunt, instead typically accessing meat via males that share meat with group members. In general, female chimpanzee dominance rank affects feeding competition, but so far, the effect of female dominance rank on meat access found different results within and across studied chimpanzee groups. Here we contribute to the debate on how female rank influences meat access while controlling for several socio-ecological variables. Multivariate analyses of 773 separate meat-eating events collected over more than 25 years from two chimpanzee communities located in the Taï National Park, Côte d'Ivoire, were used to test the importance of female dominance rank for being present at, and for acquiring meat, during meat-eating events. We found that high-ranking females were more likely to be present during a meat-eating event and, in addition, were more likely to eat meat compared to the subordinates. These findings were robust to both large demographic changes (decrease of community size) and seasonal ecological changes (fruit abundance dynamics). In addition to social rank, we found that other female properties had a positive influence on presence to meat-eating events and access to meat given presence, including oestrus status, nursing of a small infant, and age. Similar to findings in other chimpanzee populations, our results suggest that females reliably acquire meat over their lifetime despite rarely being active hunters. The implication of this study supports the hypothesis that dominance rank is an important female chimpanzee property conferring benefits for the high-ranking females.

Cytomegaloviruses in a Community of Wild Nonhuman Primates in Taï National Park, Côte D'Ivoire.

Cytomegaloviruses (CMVs) are known to infect many mammals, including a number of nonhuman primates (NHPs). However, most data available arose from studies led on captive individuals and little is known about CMV diversity in wild NHPs. Here, we analyzed a community of wild nonhuman primates (seven species) in Taï National Park (TNP), Côte d'Ivoire, with two PCR systems targeting betaherpesviruses. CMV DNA was detected in 17/87 primates (4/7 species). Six novel CMVs were identified in sooty mangabeys, Campbell's monkeys and Diana monkeys, respectively. In 3/17 positive individuals (from three NHP species), different CMVs were co-detected. A major part of the glycoprotein B coding sequences of the novel viruses was amplified and sequenced, and phylogenetic analyses were performed that included three previously discovered CMVs of western red colobus from TNP and published CMVs from other NHP species and geographic locations. We find that, despite this locally intensified sampling, NHP CMVs from TNP are completely host-specific, pinpointing the absence or rarity of cross-species transmission. We also show that on longer timescales the evolution of CMVs is characterized by frequent co-divergence with their hosts, although other processes, including lineage duplication and host switching, also have to be invoked to fully explain their evolutionary relationships.