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In order to achieve a better understanding of the factors that might have led our hominin ancestors to transition to a more terrestrial niche, including sleeping on the ground, we have conducted a study on the ground nesting behavior of chimpanzees (Pan troglodytes). Chimpanzees, like all other species of great apes, build nests in which to sleep each night, but little is known about regional differences in their nesting habits. Previously, nesting on the ground was considered typical of gorillas, but rare in most populations of chimpanzees. Using data acquired during our extensive chimpanzee nesting survey conducted between 2004 and 2013 across a > 50 000 km2 region in northern Democratic Republic of the Congo, we report a distinctive ground nesting behaviour of eastern chimpanzees (P. t. schweinfurthii). We have mapped the geographical distribution of ground nesting and compared its frequency at 20 survey areas on both sides of a large river, the Uele. We found that ground nests made up more than 1% of total nests at 15 of the 20 survey regions. For a subset of 16 of these regions, we utilized statistical models to investigate whether forest type and structure, as well as the abundance of carnivores and large herbivores, and the activities of humans impacted the frequency of ground nesting and nest height. We predicted that higher encounter rates of human and dangerous animal signs would be associated with lower rates of ground nesting as well as increased nest height. Overall, 10.4% of the Bili-Uéré chimpanzee nests were terrestrial, but the frequency of ground nesting varied extensively between the survey areas (0-29% of nests). The occurrence of ground nests was positively associated with denser forests (p = 0.004), herb patches (p < 0.001), and light gaps (p < 0.001). Light gaps (p < 0.001), herb patches (p = 0.044), and vine tangles (p = 0.016) also had a strong negative effect on nest height. Hunting by humans had a negative effect on the probability of the occurrence of ground nests (p = 0.001) and a positive one on nest height (p = 0.013), with a similar but likely marginal effect of large herbivores on nest height (p = 0.023). In addition, the chimpanzees nested at significantly lower heights with increasing distance from roads and settlements (p < 0.001). Carnivore encounter rates, however, had no significant impact on ground nest frequency or nest height. Our results indicate that ground nesting can no longer be considered a rare and patchily-occurring phenomenon in Pan troglodytes, but is instead a major component of the chimpanzee behavioural repertoire across a considerable fraction of the range of the Eastern subspecies. Our study highlights that neither the large body size of gorillas nor the taming of fire are necessary conditions for hominids to sleep overnight on the ground, even in areas inhabited by multiple species of large carnivore. Human hunting, however, appears to reduce the probability of ground nesting, or eliminate the behavior altogether.

This paper documents a community of eastern chimpanzees (Pan troglodytes schweinfurthii Giglioli, 1872) inhabiting three relict forest fragments situated on the Lake Albert escarpment, down the Ituri highlands, of eastern Democratic Republic of the Congo (DRC). The area explored had a combined forested surface of ±18.15 km2 in 2017, shrinking by 1.2% per year between 2010 and 2015. Between 2015 and 2017, we found 160 chimpanzee nests along 37.6 km of pilot walks, some up to 2,000 m altitude. Another 123 nests logged along 6.7 km transects led to an estimate of chimpanzee density of 4.62 weaned individuals per square kilometer of forest habitat. Camera‐trap images and direct observations revealed that this community is comprised of a minimum of 42 weaned individuals, which translates into an estimated density of 2.3 chimpanzees per square kilometer. The increasing rate of forest degradation threatens to erode the cultural and genetic diversity of nonhuman primates in eastern DRC; the local people however exhibit willingness to establish a community managed reserve. We hope that this report will lead to the recognition of this site as a Chimpanzee Conservation Unit, facilitating further research in these “Relict Altitude Forests Fragments of the Albert Lake Escarpment” (or RAFALE landscape) and the Ituri highlands where other undocumented chimpanzee communities occur. peerReviewed

To improve our understanding of the evolutionary origins of culture and technology in humans, it is vital that we document the full extent of behavioural diversity in our great ape relatives. About half of the world’s remaining chimpanzees (Pan spp.) live in the Democratic Republic of Congo (DRC), yet until now we have known almost nothing about their behaviour. Here we describe the insect-related tool technology of Bili-Uéré chimpanzees (Pan troglodytes schweinfurthii) inhabiting an area of at least a 50,000-km2 area in northern DRC, as well as their percussive technology associated with food processing. Over a 12-year period, we documented chimpanzee tools and artefacts at 20 survey areas and gathered data on dung, feeding remains and sleeping nests. We describe a new chimpanzee tool kit: long probes used to harvest epigaeic driver ants (Dorylus spp.), short probes used to extract ponerine ants and the arboreal nests of stingless bees, wands to dip for D. kohli and stout digging sticks used to access underground meliponine bee nests. Epigaeic Dorylus tools were significantly longer than the other tool types, and D. kohli tools were significantly thinner. Tools classified as terrestrial honey-digging sticks were a significant predictor for brushed and blunted tool ends, consistent with their presumed use. We describe two potential new tool types, an “ant scoop” and a “fruit hammer.” We document an extensive percussive technology used to process termite mounds of Cubitermes sp. and Thoracotermes macrothorax and hard-shelled fruits such as Strychnos, along with evidence of the pounding open of African giant snails and tortoises. We encountered some geographic variation in behaviour: we found honey-digging tools, long driver ant probes and fruit-pounding sites only to the north of the Uele River; there were more epigaeic Dorylus tools to the north and more ponerine ant tools to the south. We found no evidence of termite-fishing, despite the availability of Macrotermes muelleri mounds throughout the region. This lack of evidence is consistent with the results of dung washes, which revealed a substantial proportion of driver ants, but no evidence of Macrotermes or other termites. Our results allow us to describe a new chimpanzee behavioural complex, characterised by a general similarity of multiple behaviours across a large, ecologically diverse region but with subtle differences in prey choice and techniques. We propose that this widespread and related suite of behaviours be referred to as the Bili-Uéré Chimpanzee Behavioural Realm. Possible explanations for this pattern are a recent chimpanzee expansion across the region and the interconnectedness of this population across at least the entirety of northern DRC.

A key feature of human behavioral diversity is that it can be constrained by cultural preference (“cultural override”); that is, population-specific preferences can override resource availability. Here we investigate whether a similar phenomenon can be found in one of our closest relatives, as well as the potential impacts of ecological differences on feeding behavior. Our study subjects were different subpopulations of Eastern chimpanzees (Pan troglodytes schweinfurthii) occupying two very different habitats, moist tropical lowland forests vs. moist tropical forest–savanna mosaic on opposite sides of a major river. Given differences in encounter rates of different kinds of tool sites on both sides of the Uele River, we predicted that these subpopulations would differ in their likelihood of using tools to prey on two insect species despite similar availability. In surveys conducted over a 9-year period at 19 different survey regions in northern Democratic Republic of Congo (10 in lowland forest and 9 in mosaic), we collected and analyzed data on chimpanzee tool-assisted exploitation of insects. To determine the availability of insect species eaten by the chimpanzees, we counted insects and their mounds on transects and recces at 12 of these sites. For stick tools used to harvest epigaeic Dorylus and ponerine ants, we evaluated seasonal, geographical, and prey-availability factors that might influence their occurrence, using nest encounter rate as a proxy to control for chimpanzee abundance. Across the 19 survey regions spanning both sides of the Uele, we found little difference in the availability of epigaeic Dorylus and ponerine ants. Despite this, tool encounter rates for epigaeic Dorylus, but not ponerine, ants were significantly higher in the mosaic to the north of the Uele. Furthermore, we found no evidence for termite fishing anywhere, despite the availability of Macrotermes mounds throughout the region and the fact that chimpanzees at a number of other study sites use tools to harvest these termites. Instead, the chimpanzees of this region used a novel percussive technique to harvest two other types of termites, Cubitermes sp. and Thoracotermes macrothorax. This mismatch between prey availability and predation is consistent with cultural override, but given the different habitats on the two sides of the Uele River, we cannot fully rule out the influence of ecological factors. Comparing our findings with those of similar studies of other chimpanzee populations promises to contribute to our understanding of the evolution of behavioral diversity in humans and our closest cousins.

Additional Data relating to Hicks et al. 2019.

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., Chimpanzees in Comoé National Park use specialized stick tools to collect water from tree holes (WDTs). WDTs are significantly different from tools used to gather honey. The use of WDTs is widespread across neighboring communities in the park.

With great ape populations in decline across much of their range, it is crucial to obtain a global picture of their distribution and abundance, in order to guide conservation activities and to provide baseline data against which to monitor their trends. Although great apes are popular, charismatic species, we still do not possess a complete understanding of their distribution and abundance, which hinders their long-term protection. We highlight this problem by providing information on the distribution and abundance of the Eastern chimpanzee (Pan troglodytes schweinfurthii) in the northern Democratic Republic of the Congo (DRC), a region which has until now received little attention. We conducted a standing crop nest survey in the Bili area in 2005 and exploratory reconnaissance walks (recces) across the Bas-Uele region between 2004 and 2009. At Bili, the nest encounter rate in the remote forest was 4.84 nests per km (CI = 2.78–8.55) and in the area closer to the road it was 1.92 nests per km (CI = 1.08–3.43). In 2012, we repeated a part of the original transect survey and found that the nest encounter rate had remained stable over that period. On our recce walks across the region, we encountered chimpanzee nests in all forests surveyed, and within 13 km of the largest population centers. Our results suggest that the Central Uele landscape and neighboring regions are home to one of the largest remaining continuous populations of Eastern chimpanzees, that extends across at least 50,000 km2, likely representing thousands of individuals, but which is falling under increasing pressure from habitat destruction, mining and the bushmeat trade. This population has until now remained hidden from researchers and is not protected. Our results reflect gaps in our current understanding of ape distribution and abundance, and highlight the importance of obtaining more sound and complete data before assessing species status and making recommendations to guide conservation efforts.

Aim: To predict the distribution of suitable environmental conditions (SEC) for eight African great ape taxa for a first time period, the 1990s and then project it to a second time period, the 2000s; to assess the relative importance of factors influencing SEC distribution and to estimate rates of SEC loss, isolation and fragmentation over the last two decades. Location: Twenty-two African great ape range countries. Methods: We extracted 15,051 presence localities collected between 1995 and 2010 from 68 different areas surveyed across the African ape range. We combined a maximum entropy algorithm and logistic regression to relate ape presence information to environmental and human impact variables from the 1990s with a resolution of 5 9 5 km across the entire ape range. We then made SEC projections for the 2000s using updated human impact variables. Results: Total SEC area was approximately 2,015,480 and 1,807,653 km2 in the 1990s and 2000s, respectively. Loss of predicted SEC appeared highest for Cross River gorillas (_59%), followed by eastern gorillas (_52%), western gorillas (_32%), bonobos (_29%), central chimpanzees (_17%) and western chimpanzees (_11%). SEC for Nigeria-Cameroon chimpanzees and eastern chimpanzees was not greatly reduced. Except for Cross River and eastern gorillas, the number of SEC patches did not change significantly, suggesting that SEC loss was caused mainly by patch size reduction. Main conclusions: The first continent-wide perspective of African ape SEC distribution shows dramatic declines in recent years. The model has clear limitations for use at small geographic scales, given the quality of available data and the coarse resolution of predictions. However, at the large scale it has potential for informing international policymaking, mitigation of resource extraction and infrastructure development, as well as for spatial prioritization of conservation effort and evaluating conservation effectiveness.

In order to assess the impact of human activities on chimpanzee behavior, we compared reactions to humans of Eastern chimpanzees (Pan troglodytes schweinfurthii) living in proximity to and at a distance from roads and settlements in the Bili-Uele landscape in Northern Democratic Republic of the Congo (DRC). We found that chimpanzees in the remote Gangu Forest were more likely to show curious or neutral reactions to us and were less likely to flee than those living closer to roads. In addition, arboreal contact durations with Gangu chimpanzees lasted significantly longer than elsewhere. The most likely explanation for this phenomenon is that with increasing distance from roads, chimpanzees have in the recent past had fewer negative encounters with humans, and thus never learned to fear them. The discovery of this population of naive chimpanzees presents us with an important research and conservation opportunity that may result in the installation of a long-term research site and increased protection of the population.

A network of resource management areas (RMAs) exists across tropical Africa to protect natural resources. However, many are poorly managed and weakly protected. We evaluated how the lack of conservation effort influences the extinction risk of African great apes. We compiled information on presence/absence of primary (law enforcement guards) and secondary (tourism, research) conservation activities and nongovernmental conservation organizations (NGOs) support for 109 RMAs over the last 20 years. Along with these data, we collected environmental and anthropogenic variables, including recent records of ape presence/absence for all RMAs. As expected, law enforcement as a primary activity was the best predictor of ape survival rather than tourism or research as secondary activities. Furthermore, long-term NGO support had a significant positive influence on ape persistence. Our study demonstrates the feasibility of evaluating the relative importance of different conservation activities, an important step towards more evidence-based approaches in ape conservation.

African forest elephants- taxonomically and functionally unique-are being poached at accelerating rates, but we lack range-wide information on the repercussions. Analysis of the largest survey dataset ever assembled for forest elephants (80 foot-surveys; covering 13,000 km; 91,600 person-days of fieldwork) revealed that population size declined by ca. 62% between 2002-2011, and the taxon lost 30% of its geographical range. The population is now less than 10% of its potential size, occupying less than 25% of its potential range. High human population density, hunting intensity, absence of law enforcement, poor governance, and proximity to expanding infrastructure are the strongest predictors of decline. To save the remaining African forest elephants, illegal poaching for ivory and encroachment into core elephant habitat must be stopped. In addition, the international demand for ivory, which fuels illegal trade, must be dramatically reduced.

In order to assess the impact of human activities on chimpanzee behavior, we compared reactions to humans of Eastern chimpanzees (Pan troglodytes schweinfurthii) living in proximity to and at a distance from roads and settlements in the Bili-Uele landscape in Northern Democratic Republic of the Congo (DRC). We found that chimpanzees in the remote Gangu Forest were more likely to show curious or neutral reactions to us and were less likely to flee than those living closer to roads. In addition, arboreal contact durations with Gangu chimpanzees lasted significantly longer than elsewhere. The most likely explanation for this phenomenon is that with increasing distance from roads, chimpanzees have in the recent past had fewer negative encounters with humans, and thus never learned to fear them. The discovery of this population of "naïve chimpanzees" presents us with an important research and conservation opportunity that may result in the installation of a long-term research site and increased protection of the population.

Currently, the timber company Industrie Forestiere du Batalimo is selectively logging the Ngotto Forest in the Central African Republic. The forest is home to a population of chimpanzees (Pan troglodytes troglodytes) and gorillas (Gorilla gorilla gorilla) and provides the Bofi-speaking people of Grima with food, medicine, housing material, and other commodities. Over a 7-month period, the research team conducted a line-transect survey of the great ape population in the forests to the south of Grima to document their distribution. For comparison purposes, the team also surveyed a section of adjacent forest that had already been logged. Ape nests were significantly rarer in the logged forest than in the unlogged forest, and ape nests were most common in the more pristine forests to the south. This report further discusses the effects of logging and other human activities.

Over a 7-month period, stick tools constructed by chimpanzees were collected and measured at the Ngotto Forest site in Central African Republic. The chimpanzees were found to use tools to dip for ants and to probe for honey. The basic descriptions of these tools and the contexts in which they were found are presented. The lengths of two of the tool types were compared with the use of a t-test for independent groups. It was found that the lengths of the tools differed significantly depending upon their function. The location and habitat type of each tool site were plotted on a map. The tool types were distributed throughout the southern part of the study area, and with one exception all tool sites were found in the same type of habitat. Two tool sites with two other types of tools (honey hammer/club and ant club) were found. The tool types at Ngotto are compared with those found at other chimpanzee field sites, and the implications for diversity in chimpanzee material culture are discussed.

The authors would like to thank the Max Planck Society and Krekeler Foundation for generous funding of the Pan African Programme. 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. Publisher PDF