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Studies of the evolutionary relationships among gorilla populations using autosomal and mitochondrial sequences suggest that male-mediated gene flow may have been important in the past, but data on the Y-chromosomal relationships among the gorilla subspecies are limited. Here, we genotyped blood and noninvasively collected fecal samples from 12 captives and 257 wild male gorillas of known origin representing all four subspecies (Gorilla gorilla gorilla, G. g. diehli, G. beringei beringei, and G. b. graueri) at 10 Y-linked microsatellite loci resulting in 102 unique Y-haplotypes for 224 individuals. We found that western lowland gorilla (G. g. gorilla) haplotypes were consistently more diverse than any other subspecies for all measures of diversity and comprised several genetically distinct groups. However, these did not correspond to geographical proximity and some closely related haplotypes were found several hundred kilometers apart. Similarly, our broad sampling of eastern gorillas revealed that mountain (G. b. beringei) and Grauer's (G. b. graueri) gorilla Y-chromosomal haplotypes did not form distinct clusters. These observations suggest structure in the ancestral population with subsequent mixing of differentiated haplotypes by male dispersal for western lowland gorillas, and postisolation migration or incomplete lineage sorting due to short divergence times for eastern gorillas.

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.

Aim: Modelling African great ape distribution has until now focused on current or past conditions, while future scenarios remain scarcely explored. Using an ensemble forecasting approach, we predicted changes in taxon-specific distribution under future scenarios of climate, land use and human populations for (1) areas outside protected areas (PAs) only (assuming complete management effectiveness of PAs), (2) the entire study region and (3) interspecies range overlap. Location: Tropical Africa. Methods: We compiled occurrence data (n = 5,203) on African apes from the IUCN A.P.E.S. database and extracted relevant climate-, habitat- and human-related predictors representing current and future (2050) conditions to predict taxon-specific range change under a best- and a worst-case scenario, using ensemble forecasting. Results: The predictive performance of the models varied across taxa. Synergistic interactions between predictors are shaping African ape distribution, particularly human-related variables. On average across taxa, a range decline of 50% is expected outside PAs under the best scenario if no dispersal occurs (61% in worst scenario). Otherwise, an 85% range reduction is predicted to occur across study regions (94% worst). However, range gains are predicted outside PAs if dispersal occurs (52% best, 21% worst), with a slight increase in gains expected across study regions (66% best, 24% worst). Moreover, more than half of range losses and gains are predicted to occur outside PAs where interspecific ranges overlap. Main Conclusions: Massive range decline is expected by 2050, but range gain is uncertain as African apes will not be able to occupy these new areas immediately due to their limited dispersal capacity, migration lag and ecological constraints. Given that most future range changes are predicted outside PAs, Africa's current PA network is likely to be insufficient for preserving suitable habitats and maintaining connected ape populations. Thus, conservation planners urgently need to integrate land use planning and climate change mitigation measures at all decision-making levels both in range countries and abroad.

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.

AimModelling African great ape distribution has until now focused on current or past conditions, whilst future scenarios remain scarcely explored. Using an ensemble forecasting approach, we predicted changes in taxon-specific distribution under future scenarios of climate, land-use and human population changes.LocationSub-Saharan AfricaMethodsWe compiled occurrence data on African ape populations from the IUCN A.P.E.S. database and extracted relevant human-, climate- and habitat-related predictors representing current and future (2050) conditions to predict taxon-specific distribution under a best- and a worst-case scenario, using ensemble forecasting. Given the large effect on model predictions, we further tested algorithm sensitivity by considering default and non-default modelling options. The latter included interactions between predictors and polynomial terms in correlative algorithms.ResultsThe future distributions of gorilla and bonobo populations are likely to be directly determined by climate-related variables. In contrast, future chimpanzee distribution is influenced mostly by anthropogenic variables. Both our modelling approaches produced similar model accuracy, although a slight difference in the magnitude of range change was found for Gorilla beringei beringei, G. gorilla diehli, and Pan troglodytes schweinfurthii. On average, a decline of 50% of the geographic range (non-default; or 55% default) is expected under the best scenario if no dispersal occurs (57% non-default or 58% default in worst scenario). However, new areas of suitable habitat are predicted to become available for most taxa if dispersal occurs (81% or 103% best, 93% or 91% worst, non-default and default, respectively), except for G. b. beringei.Main ConclusionsDespite the uncertainty in predicting the precise proportion of suitable habitat by 2050, both modelling approaches predict large range losses for all African apes. Thus, conservation planners urgently need to integrate land-use planning and simultaneously support conservation and climate change mitigation measures at all decision-making levels both in range countries and abroad.

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.

Noninvasive samples as a source of DNA are gaining interest in genomic studies of endangered species. However, their complex nature and low endogenous DNA content hamper the recovery of good quality data. Target capture has become a productive method to enrich the endogenous fraction of noninvasive samples, such as faeces, but its sensitivity has not yet been extensively studied. Coping with faecal samples with an endogenous DNA content below 1% is a common problem when prior selection of samples from a large collection is not possible. However, samples classified as unfavourable for target capture sequencing might be the only representatives of unique specific geographical locations, or to answer the question of interest. To explore how library complexity may be increased without repeating DNA extractions and generating new libraries, in this study we captured the exome of 60 chimpanzees (Pan troglodytes) using faecal samples with very low proportions of endogenous content (, C.F. is supported by the “La Caixa” doctoral fellowship program. M.A.E. is supported by an FPI (Formación de Personal Investigador) PRE2018-083966 from Ministerio de Ciencia, Universidades e Investigación. The Pan African Programme: The Cultured Chimpanzee (PanAf) is generously funded by the Max Planck Society, the Max Planck Society Innovation Fund and the Heinz L. Krekeler Foundation. E.L is supported by CGL2017-82654-P (MINECO/FEDER,UE). M.K. is supported by “la Caixa” Foundation (ID 100010434), fellowship code LCF/BQ/PR19/11700002. T.M.-B is supported by funding from the European Research Council (ERC) under the European Union’s Horizon 2020 research and innovation programme (grant agreement No. 864203), BFU2017-86471-P (MINECO/FEDER, UE), “Unidad de Excelencia María de Maeztu”, funded by the AEI (CEX2018-000792-M), Howard Hughes International Early Career, Obra Social "La Caixa" and Secretaria d’Universitats i Recerca and CERCA Programme del Departament d’Economia i Coneixement de la Generalitat de Catalunya (GRC 2017 SGR 880).

Ecological complexity has been proposed to play a crucial role in primate brain-size evolution. However, detailed quantification of ecological complexity is still limited. Here we assess the spatio-temporal distribution of tropical fruits and young leaves, two primary chimpanzee (Pan troglodytes) foods, focusing on the predictability of their availability in individual trees. Using up to 20 years of information on monthly availability of young leaf, unripe and ripe fruit in plant species consumed by chimpanzees from tropical forests in East, Central, and West Africa, we estimated: (1) the forest-wide frequency of occurrence of each food type and (2) the predictability of finding ripe fruit-bearing trees, focusing on the timing, frequency, and amount of ripe fruit present. In all three forests, at least half of all encountered trees belonged to species that chimpanzees were known to feed on. However, the proportion of these trees bearing young leaves and fruit fluctuated widely between months. Ripe fruit was the most ephemeral food source, and trees that had more than half of their crown filled were at least nine times scarcer than other trees. In old growth forests only one large ripe fruit crop was on average encountered per 10 km. High levels of inter-individual variation in the number of months that fruit was present existed, and in some extreme cases individuals bore ripe fruit more than seven times as often as conspecifics. Some species showed substantially less variation in such ripe fruit production frequencies and fruit quantity than others. We hypothesize that chimpanzees employ a suite of cognitive mechanisms, including abilities to: (1) generalize or classify food trees; (2) remember the relative metrics of quantity and frequency of fruit production across years; and (3) flexibly plan return times to feeding trees to optimize high-energy food consumption in individual trees, and efficient travel between them. Am. J. Primatol. 78:626-645, 2016. © 2016 Wiley Periodicals, Inc.

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

Dispersal and grouping patterns form the foundations of social interactions in group-living mammals and are the outcomes of a complex interplay between inbreeding avoidance, kin cooperation and competition, predation pressure and food resource distribution. In species where both sexes disperse, the potential for kin-biased associations would seem limited. In one such species, the western lowland gorilla (WLG), short-term data suggest that female kin associations may be present due to directed local dispersal decisions, but monitoring of groups over longer timescales is needed to better elucidate this pattern. Using autosomal genotyping of 419 faecal samples representing 85 unhabituated gorillas collected non-invasively over 5 years in a 132 km2 section of Loango National Park, Gabon, we investigated the dynamics of WLG group composition, social structure and patterns of dispersal. By revealing two group dissolutions, one group formation and the movement of 13 gorillas between groups, this study demonstrates the utility of genetic analysis as a way to track individuals, groups and population dynamics on a larger scale than when monitoring the behaviour of a limited number of habituated groups or through one-time genetic sampling. Furthermore, we find that females are found in groups containing their female kin more often than expected by chance, suggesting that dispersal may not impede female kin associations in WLGs.

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.

Summary Temperament and personality research in humans and nonhuman animals measures behavioral variation in individual, population, or species-specific traits with implications for survival and fitness, such as social status, foraging, and mating success [ 1 , 2 , 3 , 4 , 5 ]. Curiosity and risk-taking tendencies have been studied extensively across taxa by measuring boldness and exploration responses to experimental novelty exposure [ 3 , 4 , 6 , 7 , 8 , 9 , 10 , 11 , 12 , 13 , 14 , 15 ]. Here, we conduct a natural field experiment using wildlife monitoring technology to test variation in the reaction of wild great apes (43 groups of naive chimpanzees, bonobos, and western gorillas across 14 field sites in Africa) to a novel object, the camera trap. Bonobo and gorilla groups demonstrated a stronger looking impulse toward the camera trap device compared to chimpanzees, suggesting higher visual attention and curiosity. Bonobos were also more likely to show alarm and other fearful behaviors, although such neophobic (and conversely, neophilic) responses were generally rare. Among all three species, individuals looked at cameras longer when they were young, were associating with fewer individuals, and did not live near a long-term research site. Overall, these findings partially validate results from great ape novelty paradigms in captivity [ 7 , 8 ]. We further suggest that species-typical leadership styles [ 16 ] and social and environmental effects, including familiarity with humans, best explain novelty responses of wild great apes. In sum, this study illustrates the feasibility of large-scale field experiments and the importance of both intrinsic and extrinsic factors in shaping animal curiosity. Video Abstract Download : Download video (48MB)

Species commonly exist in sympatry, yet ecological studies are often based on a single species approach while ignoring the impact of sympatric competitors. Over 13 mo we used 24 remote video-camera traps to monitor habitat use of sympatric chimpanzee, gorilla and elephant in four different habitat types in Loango National Park, Gabon. Habitat use by each species was predicted to vary according to seasonal changes in food availability and precipitation. Increased interspecific competition between the three species was expected at times of reduced resource availability, leading to exclusion of the inferior competitor. Supporting the predictions, species abundance per habitat showed seasonal variation: all three species responded positively to increased fruit availability in all habitats, but the response was only significant for gorilla in mature forest and elephant in coastal forest. Responses to rainfall varied, with the chimpanzee responding negatively to rainfall in swamp forest, the gorilla responding positively to rainfall in coastal and secondary forest, and the elephant responding positively to rainfall in mature forest. Elephant presence resulted in competitive exclusion of the apes under certain conditions: the chimpanzee was excluded by the elephant where fruit availability was low, whereas the gorilla was excluded by the elephant in areas of low herb density despite high fruit availability. Our results emphasize the value of applying a multi-species, longer-term approach to studying variation in habitat use among sympatric species and highlight the impact competitors can exert on one another's distribution.

Pygmy populations occupy a vast territory extending west-to-east along the central African belt from the Congo Basin to Lake Victoria. However, their numbers and actual distribution is not known precisely. Here, we undertake this task by using locational data and population sizes for an unprecedented number of known Pygmy camps and settlements (n = 654) in five of the nine countries where currently distributed. With these data we develop spatial distribution models based on the favourability function, which distinguish areas with favourable environmental conditions from those less suitable for Pygmy presence. Highly favourable areas were significantly explained by presence of tropical forests, and by lower human pressure variables. For documented Pygmy settlements, we use the relationship between observed population sizes and predicted favourability values to estimate the total Pygmy population throughout Central Africa. We estimate that around 920,000 Pygmies (over 60% in DRC) is possible within favourable forest areas in Central Africa. We argue that fragmentation of the existing Pygmy populations, alongside pressure from extractive industries and sometimes conflict with conservation areas, endanger their future. There is an urgent need to inform policies that can mitigate against future external threats to these indigenous peoples’ culture and lifestyles.

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.

Adenoviruses (AdVs) broadly infect vertebrate hosts, including a variety of nonhuman primates (NHPs). In the present study, we identified AdVs in NHPs living in their natural habitats, and through the combination of phylogenetic analyses and information on the habitats and epidemiological settings, we detected possible horizontal transmission events between NHPs and humans. Wild NHPs were analyzed with a pan-primate AdV-specific PCR using a degenerate nested primer set that targets the highly conserved adenovirus DNA polymerase gene. A plethora of novel AdV sequences were identified, representing at least 45 distinct AdVs. From the AdV-positive individuals, 29 nearly complete hexon genes were amplified and, based on phylogenetic analysis, tentatively allocated to all known human AdV species ( Human adenovirus A to Human adenovirus G [HAdV-A to -G]) as well as to the only simian AdV species ( Simian adenovirus A [SAdV-A]). Interestingly, five of the AdVs detected in great apes grouped into the HAdV-A, HAdV-D, HAdV-F, or SAdV-A clade. Furthermore, we report the first detection of AdVs in New World monkeys, clustering at the base of the primate AdV evolutionary tree. Most notably, six chimpanzee AdVs of species HAdV-A to HAdV-F revealed a remarkably close relationship to human AdVs, possibly indicating recent interspecies transmission events.

Dietary overlap of sympatric apes is complex and understudied, but its examination is essential to further our understanding of species distribution, abundance, and community ecology. Over 3 yr we studied food availability and dietary composition of central chimpanzees (Pan troglodytes troglodytes) and western gorillas (Gorilla gorilla gorilla) in Loango National Park, Gabon. We predicted that living in a habitat dominated by mature forest with sparse terrestrial herbaceous vegetation would lead to an increase in frugivory by gorillas, resulting in increased dietary overlap between the 2 ape species vs. other sites, but that chimpanzees would remain more frugivorous than gorillas. Through fecal analysis we measured overlap in fruit consumption between the 2 species on a bimonthly basis using the Renkonens method. Mean overlap was 27.5% but varied greatly seasonally, ranging between 0.3% and 69%, indicating that when examined on a finer scale, the degree of overlap appears much lower than at other study sites. In contrast to studies elsewhere, there was not a positive correlation between rainfall and fruit availability in Loango, and the long dry season was a period of high fruit production. As observed elsewhere, we found a positive correlation between fruit consumption and fruit availability for both chimpanzees and gorillas and we found that chimpanzees were more frugivorous than gorillas. A very low availability of herbs did not lead to increased frugivory by gorillas nor increased overlap between the 2 ape species vs. other field sites. We conclude that forest composition, fruit availability, and dietary variability of sympatric species can vary greatly between locations, and that chimpanzees and gorillas can adapt to primary forest with little undergrowth, where they concentrate their diet on fruit and leaves.

Obtaining reliable population size or abundance estimates of endangered species is critical for their conservation and management. Genotyping non-invasively collected samples is an effective way to gain insights into the number and grouping patterns of rare or elusive animals. In this study we used genetic capture–recapture estimators to obtain a precise estimate of the size of a western gorilla population inhabiting an intensely sampled 101 km 2 area in Loango National Park, Gabon. Using 394 putative gorilla samples collected opportunistically over a 3 year period, we identified 83 unique genotypes. We used a rarefaction curve, Bayesian estimator and two maximum-likelihood methods to estimate that between 87 and 107 individuals used the study area between February 2005 and September 2007. The confidence interval surrounding the genetic estimate was smaller than that obtained using traditional ape survey methods. In addition, genetic analysis showed that gorilla and chimpanzee faeces were identified with 98% and 95% accuracy in the field, respectively. Patterns of co-occurrence of individual gorillas suggest that at least 11 gorilla social groups and five lone silverback males lived in the study area and that several individuals transferred between groups during the 3-year study period. When properly designed and implemented as part of a long-term biomonitoring program, genetic capture-recapture should prove an invaluable tool for evaluating, even on a large-scale, the population size and dynamics of apes and other elusive species.

Homo faber was once proposed as a label for humans specifically to highlight their unique propensity for tool use. However, new observations on complex tool use by the chimpanzees of Loango National Park, Gabon, expand our knowledge about tool-using abilities in Pan troglodytes. Chimpanzees in Loango, when using tools to extract honey from three types of bee nests, were observed to regularly use three- to five-element tool sets. In other words, different types of tools were used sequentially to access a single food source. Such tool sets included multi-function tools that present typical wear for two distinct uses. In addition, chimpanzees exploited underground bee nests and used ground-perforating tools to locate nest chambers that were not visible from the ground surface. These new observations concur with others from Central African chimpanzees to highlight the importance of honey extraction in arguments favoring the emergence of complex tool use in hominoids, including different tool types, expanded tool sets, multifunction tools, and the exploitation of underground resources. This last technique requires sophisticated cognitive abilities concerning unseen objects. A sequential analysis reveals a higher level of complexity in honey extraction than previously proposed for nut cracking or hunting tools, and compares with some technologies attributed to early hominins from the Early and Middle Stone Age. A better understanding of similarities in human and chimpanzee tool use will allow for a greater understanding of tool-using skills that are uniquely human.

In some populations, chimpanzees engage in lethal aggression within and between social units. We report a fatal attack on an adult male chimpanzee at a new research site in Loango National Park, Gabon. We found a fresh corpse of an adult male chimpanzee only a few hundred meters from the research camp, after noting numerous vocalizations and chimpanzee movements the previous evening. Previous contacts with chimpanzees and fresh tracks in the area around the corpse suggest that 2 communities of chimpanzees range where the attack occurred and that members of the neighboring community killed the chimpanzee. To support the conclusion, we conducted genetic analysis for 13 Y-chromosome loci and 9 microsatellite loci of fecal samples from the dead individual, 5 possible attackers, and 2 members of the other community Though we cannot exclude the possibility of an intracommunity killing, the combined observational and genetic evidence suggest an intercommunity attack. The case study adds to the growing evidence that intercommunity killings are a rare but widespread phenomenon among chimpanzees and not an artifact of human provisioning or habituation.

The feeding ecology of sympatric great ape species yields valuable information for palaeodietary reconstructions in sympatric early hominin species. However, no isotopic references on sympatrically living apes and their feeding ecology are currently available. Here we present the first isotopic study on sympatric great apes, namely western lowland gorillas (Gorilla gorilla gorilla) and central chimpanzees (Pan troglodytes troglodytes) from Loango National Park, Gabon. We successfully analyzed the stable carbon and nitrogen isotope ratios in a selection of food plants (n=31) and hair samples (n=30) retrieved from sleeping nests to test whether niche partitioning among sympatric chimpanzees and gorillas is detectable using isotope analysis of hair. Ape hair strands with roots were sectioned into sequential segments (n=100) to investigate temporal isotopic variation related to seasonal variations in food resources. We found significant δ(13)C differences between herbaceous plants and fruits, most likely due to canopy effects. While the δ(13)C values of chimpanzees indicate the consumption of fruit, the low δ(13)C values in gorilla hair indicate folivory, most likely the consumption of (13)C-depleted herbaceous vegetation. Our isotopic data also confirmed dietary overlap between chimpanzees and gorillas, which varied by season. Gorillas showed significant variation in δ(13)C values in response to season due to shifting proportions of herbaceous plants versus fruits. In chimpanzees, significant seasonal variation in δ(15)N was likely related to the seasonal availability of fruit species with particularly high δ(15)N values. In summary, we found isotopic evidence for niche partitioning and seasonal dietary variation among sympatric great apes at Loango. These findings provide a valuable reference for palaeodietary research on fossil hominins using δ(13)C analyses, particularly for studies focusing on sympatric taxa and on temporal isotopic variation within incremental tissues such as tooth enamel.

Adenoviruses (AdV) broadly infect vertebrate hosts including a variety of primates. We identified a novel AdV in the feces of captive gorillas by isolation in cell culture, electron microscopy and PCR. From the supernatants of infected cultures we amplified DNA polymerase (DPOL), preterminal protein (pTP) and hexon gene sequences with generic pan primate AdV PCR assays. The sequences in-between were amplified by long-distance PCRs of 2 - 10 kb length, resulting in a final sequence of 15.6 kb. Phylogenetic analysis placed the novel gorilla AdV into a cluster of primate AdVs belonging to the species Human adenovirus B (HAdV-B). Depending on the analyzed gene, its position within the cluster was variable. To further elucidate its origin, feces samples of wild gorillas were analyzed. AdV hexon sequences were detected which are indicative for three distinct and novel gorilla HAdV-B viruses, among them a virus nearly identical to the novel AdV isolated from captive gorillas. This shows that the discovered virus is a member of a group of HAdV-B viruses that naturally infect gorillas. The mixed phylogenetic clusters of gorilla, chimpanzee, bonobo and human AdVs within the HAdV-B species indicate that host switches may have been a component of the evolution of human and non-human primate HAdV-B viruses.