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Socioecological variables influence the rate and nature of encounters between conspecific primate groups. Both social (i.e., type of encounter) and ecological factors (i.e., fruit availability) affect rates and outcomes of intergroup encounters in eastern gorillas. However, the roles of individual factors underlying these events among western lowland gorillas (Gorilla gorilla gorilla) remain relatively understudied. We assessed whether group composition (the number of subordinate males and females in the group) and fruit availability influenced the rate of intergroup encounters in three groups of western lowland gorillas in the Republic of Congo. From April 2014 to October 2018, we monitored focal gorilla groups daily in the Goualougo and Djeke Triangles. To assess the ecological similarity between sites, we compared botanical surveys of trees and herbs across them and found overall herb densities were similar between sites. We observed 294 intergroup encounters over 2797 observation days, with a mean rate of 2.20 ± SD 0.93 encounters/month (range: 1.59–3.94) across groups, and these interactions ranged from tolerant to aggressive. The number of subordinate males in the group correlated negatively with the occurrence of intergroup encounters, and groups were more likely to avoid interaction or react aggressively when they encountered a solitary male than when they encountered another group. Some differences between the groups may be attributed to familiarity with extragroup members and the status of the dominant silverback. We found no relationship between the rate of encounters and fruit availability. Our findings indicate that defense of mates, rather than of food resources, may be the most important driver of between-group competition in western lowland gorillas. These findings, which are similar to those for mountain gorillas, suggest that primate social systems may be more variable than previously anticipated within taxa. Therefore, further longitudinal studies are needed to determine the generality of our findings and assess the direct impacts of intergroup encounters on fitness.

Paleoclimate reconstructions have enhanced our understanding of how past climates have shaped present-day biodiversity. We hypothesize that the geographic extent of Pleistocene forest refugia and suitable habitat fluctuated significantly in time during the late Quaternary for chimpanzees (Pan troglodytes). Using bioclimatic variables representing monthly temperature and precipitation estimates, past human population density data, and an extensive database of georeferenced presence points, we built a model of changing habitat suitability for chimpanzees at fine spatio-temporal scales dating back to the Last Interglacial (120,000 BP). Our models cover a spatial resolution of 0.0467 degrees (approximately 5.19 km(2) grid cells) and a temporal resolution of between 1000 and 4000 years. Using our model, we mapped habitat stability over time using three approaches, comparing our modeled stability estimates to existing knowledge of Afrotropical refugia, as well as contemporary patterns of major keystone tropical food resources used by chimpanzees, figs (Moraceae), and palms (Arecacae). Results show habitat stability congruent with known glacial refugia across Africa, suggesting their extents may have been underestimated for chimpanzees, with potentially up to approximately 60,000 km(2) of previously unrecognized glacial refugia. The refugia we highlight coincide with higher species richness for figs and palms. Our results provide spatio-temporally explicit insights into the role of refugia across the chimpanzee range, forming the empirical foundation for developing and testing hypotheses about behavioral, ecological, and genetic diversity with additional data. This methodology can be applied to other species and geographic areas when sufficient data are available.

Funder: Max-Planck-Gesellschaft (Max Planck Society); doi: https://doi.org/10.13039/501100004189, Funder: Max Planck Society Innovation Fund Heinz L. Krekeler Foundation, Much like humans, chimpanzees occupy diverse habitats and exhibit extensive behavioural variability. However, chimpanzees are recognized as a discontinuous species, with four subspecies separated by historical geographic barriers. Nevertheless, their range-wide degree of genetic connectivity remains poorly resolved, mainly due to sampling limitations. By analyzing a geographically comprehensive sample set amplified at microsatellite markers that inform recent population history, we found that isolation by distance explains most of the range-wide genetic structure of chimpanzees. Furthermore, we did not identify spatial discontinuities corresponding with the recognized subspecies, suggesting that some of the subspecies-delineating geographic barriers were recently permeable to gene flow. Substantial range-wide genetic connectivity is consistent with the hypothesis that behavioural flexibility is a salient driver of chimpanzee responses to changing environmental conditions. Finally, our observation of strong local differentiation associated with recent anthropogenic pressures portends future loss of critical genetic diversity if habitat fragmentation and population isolation continue unabated.

Large brains and behavioural innovation are positively correlated, species-specific traits, associated with the behavioural flexibility animals need for adapting to seasonal and unpredictable habitats. Similar ecological challenges would have been important drivers throughout human evolution. However, studies examining the influence of environmental variability on within-species behavioural diversity are lacking despite the critical assumption that population diversification precedes genetic divergence and speciation. Here, using a dataset of 144 wild chimpanzee (Pan troglodytes) communities, we show that chimpanzees exhibit greater behavioural diversity in environments with more variability - in both recent and historical timescales. Notably, distance from Pleistocene forest refugia is associated with the presence of a larger number of behavioural traits, including both tool and non-tool use behaviours. Since more than half of the behaviours investigated are also likely to be cultural, we suggest that environmental variability was a critical evolutionary force promoting the behavioural, as well as cultural diversification of great apes.

Funder: Max-Planck-Gesellschaft (Max Planck Society); doi: https://doi.org/10.13039/501100004189, Funder: Heinz L. Krekeler Foundation, Large brains and behavioural innovation are positively correlated, species-specific traits, associated with the behavioural flexibility animals need for adapting to seasonal and unpredictable habitats. Similar ecological challenges would have been important drivers throughout human evolution. However, studies examining the influence of environmental variability on within-species behavioural diversity are lacking despite the critical assumption that population diversification precedes genetic divergence and speciation. Here, using a dataset of 144 wild chimpanzee (Pan troglodytes) communities, we show that chimpanzees exhibit greater behavioural diversity in environments with more variability — in both recent and historical timescales. Notably, distance from Pleistocene forest refugia is associated with the presence of a larger number of behavioural traits, including both tool and non-tool use behaviours. Since more than half of the behaviours investigated are also likely to be cultural, we suggest that environmental variability was a critical evolutionary force promoting the behavioural, as well as cultural diversification of great apes.

AimPaleoclimate reconstructions have enhanced our understanding of how past climates may have shaped present-day biodiversity. We hypothesize that habitat stability in historical Afrotropical refugia played a major role in the habitat suitability and persistence of chimpanzees (Pan troglodytes) during the late Quaternary. We aimed to build a dynamic model of changing habitat suitability for chimpanzees at fine spatio-temporal scales to provide a new resource for understanding their ecology, behaviour and evolution.LocationAfrotropics.TaxonChimpanzee (Pan troglodytes), including all four subspecies (P. t. verus, P. t. ellioti, P. t. troglodytes, P. t. schweinfurthii).MethodsWe used downscaled bioclimatic variables representing monthly temperature and precipitation estimates, historical human population density data and an extensive database of georeferenced presence points to infer chimpanzee habitat suitability at 62 paleoclimatic time periods across the Afrotropics based on ensemble species distribution models. We mapped habitat stability over time using an approach that accounts for dispersal between time periods, and compared our modelled stability estimates to existing knowledge of Afrotropical refugia. Our models cover a spatial resolution of 0.0467 degrees (approximately 5.19 km2 grid cells) and a temporal resolution of every 1,000–4,000 years dating back to the Last Interglacial (120,000 BP).ResultsOur results show high habitat stability concordant with known historical forest refugia across Africa, but suggest that their extents are underestimated for chimpanzees. We provide the first fine-grained dynamic map of historical chimpanzee habitat suitability since the Last Interglacial which is suspected to have influenced a number of ecological-evolutionary processes, such as the emergence of complex patterns of behavioural and genetic diversity.Main ConclusionsWe provide a novel resource that can be used to reveal spatio-temporally explicit insights into the role of refugia in determining chimpanzee behavioural, ecological and genetic diversity. This methodology can be applied to other taxonomic groups and geographic areas where sufficient data are available.

Funding: Generous funding for the Pan African Programme was provided by the Max Planck Society, Max Planck Society Innovation Fund, Krekeler Foundation, Robert Bosch Foundation, Chimbo Foundation and UCSD-SALK Institute Center for Academic Research and Training in Anthropogeny (CARTA). 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. Postprint

Lions (Panthera leo) are of particular conservation concern due to evidence of recent, widespread population declines in what has hitherto been seen as a common species, robust to anthropogenic disturbance. Here we use non-invasive methods to recover complete mitochondrial genomes from single hair samples collected in the field in order to explore the identity of the Gabonese Plateaux Batéké lion. Comparison of the mitogenomes against a comprehensive dataset of African lion sequences that includes relevant geographically proximate lion populations from both contemporary and ancient sources, enabled us to identify the Plateaux Batéké lion as a close maternal relative to now extirpated populations found in Gabon and nearby Congo during the twentieth century, and to extant populations of Southern Africa. Our study demonstrates the relevance of ancient DNA methods to field conservation work, and the ability of trace field samples to provide copious genetic information about free-ranging animals. Acknowledgements We thank the laboratory technicians of the Centre for GeoGenetics and the staff of the Danish National High-Throughput DNA Sequencing Centre for technical assistance. This project received funding from the European Union�s Seventh Framework Programme for research, technological development and demonstration under grant agreement no. FP7-PEOPLE-2011-IEF-298820, and ERC Consolidator Grant 681396 � Extinction Genomics. Fieldwork in Gabon was funded by Panthera and The Aspinall Foundation. We thank the Staff at Projet Protection Gorilles Gabon and the Gabonese National Park Agency for their assistance in fieldwork. Thanks also to Tony King for helpful comments on the manuscript. Permission to collect and export the samples was granted by the respective Ministries of the Environment (Eaux et Forets) in Gabon and Congo, and by Gabon�s National Park Agency (ANPN). Scopus

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