Although wild chimpanzees (Pan troglodytes) have been studied intensely for more than 50 years, there are still many aspects of their ecology and behavior that are not well understood. Every time that a new population of chimpanzees has been studied, new behaviors and unknown aspects of their ecology have been discovered. All this accumulated knowledge is helping us to piece together a model of how could last human and chimpanzee common ancestors have lived and behaved between seven and five million years ago. Comoé chimpanzees had never been studied in depth, until we started our research in October 2014, only a few censuses had been realized. The last surveys prior our work, stated that the population was so decimated that was probably functionally extinct. When we started this research, we had to begin with a new intensive survey, using new methods, to ascertain the real status and distribution of the chimpanzees living in Comoé National Park (CNP). During the last five years, we have realized a deep study aiming to know more about their ecology and behavior. We combined transects and reconnaissance marches (recces) with the use of camera traps, for the first time in CNP, obtaining a wealth of data that is not fully comprised in this dissertation. With this research, we determined that there is a sustainable continuous population of Western chimpanzees (Pan troglodytes verus) in CNP and the adjacent area of Mont Tingui, to the West, with a minimum of 127 weaned chimpanzees living in our main 900 km2 study area, SW of CNP. We found that this population is formed by a minimum of eight different chimpanzee communities, of which we studied seven, four of them more in detail. These chimpanzees spent much more time in the forest than in the savanna habitats. We also found that Comoé chimpanzees consumed at least 58 different food items in their dit, which they obtained both from forest and savanna habitats. Another finding was that insectivory had an important role in their diet, with at least four species of ants, three of termites and some beetle larvae. These chimpanzees also hunted at least three species of monkeys and maybe rodents and duikers and occasionally consumed the big land snails of genus Achatina. We found that, during the fruit scarcity period in the late rainy season, they intensely consumed the cambium of Ceiba pentandra, as fallback food, much more than the bark or cambium of any other tree species. Another interesting finding was that all the chimpanzees in the studied area realized this particular bark-peeling behavior and had been repeatedly peeling the trees of this species for years. This did not increase tree mortality and the damage caused to the trees was healed in two years, not reducing the growth, thus being a sustainable use of the trees. We found that Comoé chimpanzees produced and used a great variety of tools, mainly from wooden materials, but also from stone and herbaceous vegetation. Their tool repertory included stick tools to dip for Dorylus burmeisteri ants, to fish for Camponotus and Crematogaster ants, to dip for honey, mainly from Meliponini stingless bees, but sometimes from honey bees (Apis mellifera). It also included the use of stick tools to fish termites of Macrotermes subhyalinus and Odontotermes majus (TFTs), to dip for water from tree holes and investigatory probes for multiple purposes. Additionally, these chimpanzees used leaf-sponges to drink from tree holes and to collect clayish water from salt-licks. They also used stones to hit the buttresses of trees during displays, the so called accumulative stone throwing behavior and probably used stones as hammers, to crack open hard-shelled Strichnos spinosa and Afraegle paniculata fruits and Achatina snails. The chimpanzees also used objects that are not generally accepted as animal tools, for being attached to the substrate, with different purposes: they drummed buttresses of trees with hands and/or feet to produce sound during male displays and they pounded open hard-shelled fruits, Achatina snails and Cubitermes termite mounds on stone or root anvils. We finally measured the stick tools and found significant differences between them suggesting that they were specialized tools made specifically for every purpose. We studied more in detail the differences between apparently similar tools, the honey dipping tools and the water dipping tools, often with brushes made at their tips to collect the fluids. These last tools were exclusive from Comoé and have not been described at any other site. We found that total length, diameter and brush length were significantly different, suggesting that they were specialized tools. We concluded that Comoé chimpanzees had a particular culture, different from those of other populations of Western chimpanzees across Africa. Efficient protection, further research and permanent presence of research teams are required to avoid that this unique population and its culture disappears by the poaching pressure and maybe by the collateral effects of climate change.
Little information is currently available on the occurrence and genetic diversity of pathogenic and commensal protist species in captive and semi-captive non-human primates (NHP) resident in zoological gardens or sanctuaries in low- and medium-income countries. In this molecular-based study, we prospectively collected individual faecal samples from apparently healthy NHP at the Abidjan Zoological Garden (AZG) in Côte d’Ivoire, the Tacugama Sanctuary (TS) in Sierra Leone, and the Quistococha Zoological Garden (QZG) in Peru between November 2018 and February 2020. We evaluated for the presence of pathogenic (Cryptosporidium spp., Entamoeba histolytica, Giardia duodenalis, Blastocystis sp., Enterocytozoon bieneusi, Balantioides coli) and commensal (Entamoeba dispar, Troglodytella abrassarti) protist species using PCR methods and Sanger sequencing. Giardia duodenalis was the most prevalent species found (25.9%, 30/116), followed by Blastocystis sp. (22.4%, 26/116), and E. dispar (18.1%, 21/116). We detected E. bieneusi (4.2%, 1/24) and T. abrassarti (12.5%, 3/24) only on NHP from AZG. Cryptosporidium spp., E. histolytica, and B. coli were undetected at the three sampling sites investigated here. Sequence analyses revealed the presence of zoonotic sub-assemblages BIII (n = 1) in AZG and BIV (n = 1) in TS within G. duodenalis. We identified Blastocystis subtype ST3 (100%, 6/6) in AZG, ST1 (80.0%, 12/15), ST2 (6.7%, 1/15), and ST3 (13.3%, 2/15) in TS, and ST2 (80.0%, 4/5) and ST3 (20.0%, 1/5) in QZG. The only E. bieneusi isolate detected here was identified as zoonotic genotype CAF4. Our PCR-based data indicate that potentially pathogenic protist species including G. duodenalis, Blastocystis sp., E. bieneusi, and B. coli are present at variable rates in the three NHP populations investigated here. The identification of zoonotic genotypes within these species indicates that human-NHP transmission is possible, although the extent and directionality of these events need to be elucidated in future molecular surveys.
The white naped mangabey (Cercocebus lunulatus) is on the IUCN Red List of Critically Endangered Species and is endemic to 3 countries in West Africa. To date, while there are conservation efforts in Ghana, there is no established conservation strategy for this species, particularly in Côte d'Ivoire and Burkina Faso. At the risk of seeing this species disappear in the near future, it is urgent to put in place a conservation strategy that integrates both specific protection policies and specific scientific information in order to better develop this conservation strategy. Even if we note in recent years that the OIPR, the managing institution, is taking over the park with monitoring actions that need additional support, it is necessary to highlight the lack of updated scientific data to develop a more effective protection strategy for the remaining populations in Côte d'Ivoire. It is with this in mind that we conducted the first preliminary studies after the 2011 military-political crisis in Côte d'Ivoire on the structure and distribution of the groups of Cercocebus lunulatus, and the characterization of the habitats they currently occupy. in the savannah-forest mosaic of the Comoé Park. Our first results show that groups have a high percentage of juveniles and groups are distributeded along, watercourses mainly in gallery forests at Cynometra 33% followed by 25% deciduous forest galleries, although they also use extensively the forest islands, not linked to watercourses.. These results provide the base for a larger and more intensive study to provide updated scientific data for the development of a good conservation strategy.
Primates often consume either bark or cambium (inner bark) as a fallback food to complete their diet during periods of food scarcity. Wild chimpanzees exhibit great behavioral diversity across Africa, as studies of new populations frequently reveal. Since 2014, we have been using a combination of camera traps and indirect signs to study the ecology and behavior of wild chimpanzees (Pan troglodytes verus) in Comoé National Park, Ivory Coast, to document and understand the behavioral adaptations that help them to survive in a savanna–forest mosaic landscape. We found that Comoé chimpanzees peel the bark of the buttresses of kapok tree (Ceiba pentandra) trees to eat the cambium underneath. Individuals of all sex/age classes across at least six neighboring communities peeled the bark, but only during the late rainy season and beginning of the dry season, when cambium may represent an important fallback food. Baboons (Papio anubis) also target the same trees but mainly eat the bark itself. Most of the bark-peeling wounds on Ceiba trees healed completely within 2 years, seemingly without any permanent damage. We recorded chimpanzees visiting trees in early stages of wound recovery but leaving them unpeeled. Only 6% of peeled trees (N = 53) were reexploited after a year, suggesting that chimpanzees waited for the rest of the trees to regrow the bark fully before peeling them again, thus using them sustainably. Many human groups of hunter-gatherers and herders exploited cambium sustainably in the past. The observation that similar sustainable bark-peeling behavior evolved in both chimpanzees and humans suggests that it has an important adaptive value in harsh environments when other food sources become seasonally scarce, by avoiding the depletion of the resource and keeping it available for periods of scarcity.
The West African chimpanzee is critically endangered (CR). From 1990 to 2007, Ivory Coast lost 90% of its population to habitat destruction and poaching. In order to effectively implement conservation measures, we need to determine the status of any remaining populations in the country. The chimpanzee population of Comoé National Park (CNP) was assumed to have been severely depleted following the politico-military crisis of 2002–2011. Surveys in 2007, 2010, 2012, and 2014 failed to find significant evidence of chimpanzees in the park, leading managers to believe that no sustainable population was left. To evaluate status and distribution of chimpanzees in CNP, in 2015 we conducted a stratified survey in our study area in the southwest of the park. Over the next 3 years, we conducted recce walks in the north, east, and center of the park, and in 2017 we collected additional data on distribution of chimpanzees during the full park survey for elephants. Additionally, for the first time in northern Ivory Coast, we carried out a local nest decay study. In our main study area, we estimated a density of 0.14 weaned chimpanzees/km2, with an abundance of 127 (92–176) weaned chimpanzees, representing a sustainable population in CNP. We identified 123 individual chimpanzees via parallel camera-trap survey. We discovered a resident chimpanzee population to the east of the Comoé River, an area previously assumed devoid of chimpanzees. This study confirms the viability of a population key for the conservation of Western chimpanzee. We stress the importance of concentrating stratified surveys in potential wildlife habitat to determine the distribution of this and other cryptic threatened species.
The goal of this study was to explain the patterns of diversity and distribution of arboreal social bees nesting in forest habitats of the Comoé National park, within the home-ranges of wild chimpanzees that consume their honey. Investigations were done using a total sixteen plots, one hectare each, established in three habitat types (mature forest island, secondary forest island and gallery forest). The diversity and distribution of arboreal social bees was estimated with visuals searches. The exploitation of the beehives of these bee by the chimpanzees was also evaluated using chimpanzees’ honey dipping tools as indicators. Results revealed five bees’ species belonging to two tribes; Meliponini (Meliponula ferruginea, Meliponula togoensis, Meliponula bocandei, Hypotrigona gribodoi) and Apini (Apis mellifera). Frequent exploitation of the honey of stingless bees by the chimpanzees was observed, except for H. gribodoi. Meliponula ferruginea was the most exploited species by chimpanzees. A total of 114 beehives were found in the overall established plots leading to an estimated density of 2.4 beehives/ha in the study area. Among the surveyed habitats, mature forest island was found to harbor the highest beehive density (4.2 beehives/ha), followed respectively by secondary-forest island (1.9 beehives/ha) and gallery forest (1.1 beehives/ha). Finally, all bee species were found nesting in cavities of trees with a DBH ranging from 15 to 87.3 cm, with a special preference for Dialium guinneense. However, the DBH of nesting trees and beehives’ height, measured from the ground level, did not significantly influence the honey exploitation by chimpanzees. In sum bee species diversity and distribution might be important in the survival of chimpanzees of a forest savanna landscape.
Objectives We investigated occurrences and patterns of terrestrial nocturnal activity in wild chimpanzees (Pan troglodytes) and modelled the influence of various ecological predictors on nocturnal activity. Methods Data were extracted from terrestrial camera‐trap footage and ecological surveys from 22 chimpanzee study sites participating in the Pan African Programme: The Cultured Chimpanzee. We described videos demonstrating nocturnal activity, and we tested the effects of the percentage of forest, abundance of predators (lions, leopards and hyenas), abundance of large mammals (buffalos and elephants), average daily temperature, rainfall, human activity, and percent illumination on the probability of nocturnal activity. Results We found terrestrial nocturnal activity to occur at 18 of the 22 study sites, at an overall average proportion of 1.80% of total chimpanzee activity, and to occur during all hours of the night, but more frequently during twilight hours. We found a higher probability of nocturnal activity with lower levels of human activity, higher average daily temperature, and at sites with a larger percentage of forest. We found no effect of the abundance of predators and large mammals, rainfall, or moon illumination. Discussion Chimpanzee terrestrial nocturnal activity appears widespread yet infrequent, which suggests a consolidated sleeping pattern. Nocturnal activity may be driven by the stress of high daily temperatures and may be enabled at low levels of human activity. Human activity may exert a relatively greater influence on chimpanzee nocturnal behavior than predator presence. We suggest that chimpanzee nocturnal activity is flexible, enabling them to respond to changing environmental factors.
Many animals show population specific behavioral variation, with chimpanzees (Pan troglodytes) exhibiting exceptionally high levels of behavioral diversity (1, 2). This diversity has been documented in a variety of contexts, including communication , thermoregulation and extractive foraging (table S1). Chimpanzees are also proficient tool-users, using sticks, leaves and stones to access honey, insects, meat, nuts and algae (table S1). Many of these behaviors are inferred to be socially learned and therefore cultural (2), although the influence of genetic and environmental variation cannot always be ruled out (3). Culture in chimpanzees is supported by the occurrence of local traditions irrespective of resource or tool abundance (1, 2), and controlled experiments demonstrating that naïve chimpanzees can socially learn new behaviors (4, 5). Moreover, new behaviors, or variants, are regularly discovered when observing previously unstudied populations (5) Chimpanzees possess a large number of behavioral and cultural traits among non-human species. The 'disturbance hypothesis' predicts that human impact depletes resources and disrupts social learning processes necessary for behavioral and cultural transmission. We used an unprecedented data set of 144 chimpanzee communities, with information on 31 behaviors, to show that chimpanzees inhabiting areas with high human impact have a mean probability of occurrence reduced by 88%, across all behaviors, compared to low impact areas. This behavioral diversity loss was evident irrespective of the grouping or categorization of behaviors. Therefore, human impact may not only be associated with the loss of populations and genetic diversity, but also affects how animals behave. Our results support the view that 'culturally significant units' should be integrated into wildlife conservation.
The study of the archaeological remains of fossil hominins must rely on reconstructions to elucidate the behaviour that may have resulted in particular stone tools and their accumulation. Comparatively, stone tool use among living primates has illuminated behaviours that are also amenable to archaeological examination, permitting direct observations of the behaviour leading to artefacts and their assemblages to be incorporated. Here, we describe newly discovered stone tool-use behaviour and stone accumulation sites in wild chimpanzees reminiscent of human cairns. In addition to data from 17 mid- to long-term chimpanzee research sites, we sampled a further 34 Pan troglodytes communities. We found four populations in West Africa where chimpanzees habitually bang and throw rocks against trees, or toss them into tree cavities, resulting in conspicuous stone accumulations at these sites. This represents the first record of repeated observations of individual chimpanzees exhibiting stone tool use for a purpose other than extractive foraging at what appear to be targeted trees. The ritualized behavioural display and collection of artefacts at particular locations observed in chimpanzee accumulative stone throwing may have implications for the inferences that can be drawn from archaeological stone assemblages and the origins of ritual sites.
Over a 6 month period during the dry season, from the end of October 2014 to the beginning of May 2015, we studied tool use behavior of previously unstudied and non-habituated savanna chimpanzees (Pan troglodytes verus) living in the Comoé National Park, Ivory Coast (CI). We analyzed all the stick tools and leaf-sponges found that the chimpanzees used to forage for ants, termites, honey, and water. We found a particular behavior to be widespread across different chimpanzee communities in the park, namely, dipping for water from tree holes using sticks with especially long brush-tip modifications, using camera traps, we recorded adults, juveniles, and infants of three communities displaying this behavior. We compared water dipping and honey dipping tools used by Comoé chimpanzees and found significant differences in the total length, diameter, and brush length of the different types of fluid-dipping tools used. We found that water dipping tools had consistently longer and thicker brush-tips than honey dipping tools. Although this behavior was observed only during the late dry season, the chimpanzees always had alternative water sources available, like pools and rivers, in which they drank without the use of a tool. It remains unclear whether the use of a tool increases efficient access to water. This is the first time that water dipping behavior with sticks has been found as a widespread and well-established behavior across different age and sex classes and communities, suggesting the possibility of cultural transmission. It is crucial that we conserve this population of chimpanzees, not only because they may represent the second largest population in the country, but also because of their unique behavioral repertoire.