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Poaching is one of the greatest threats to wildlife conservation world-wide. However, the spatial and temporal patterns of poaching activities within protected areas, and the effectiveness of ranger patrols and ranger posts in mitigating these threats, are relatively unknown. We used 10 years (2006–2015) of ranger-based monitoring data and dynamic multi-season occupancy models to quantify poaching-related threats, to examine factors influencing the spatio-temporal dynamics of these threats and to test the efficiency of management actions to combat poaching in Nyungwe National Park (NNP), Rwanda. The probability of occurrence of poaching-related threats was highest at lower elevations (1,801–2,200 m), especially in areas that were close to roads and tourist trails; conversely, occurrence probability was lowest at high elevation sites (2,601–3,000 m), and near the park boundary and ranger posts. The number of ranger patrols substantially increased the probability that poaching-related threats disappear at a site if threats were originally present (i.e. probability of extinction of threats). Without ranger visits, the annual probability of extinction of poaching-related threats was an estimated 7%; this probability would increase to 20% and 57% with 20 and 50 ranger visits per year, respectively. Our results suggest that poaching-related threats can be effectively reduced in NNP by adding ranger posts in areas where they do not currently exist, and by increasing the number of patrols to sites where the probability of poaching activities is high. Synthesis and applications. Our application of dynamic occupancy models to predict the probability of presence of poaching-related threats is novel, and explicitly considers imperfect detection of illegal activities. Based on the modelled relationships, we identify areas that are most vulnerable to poaching, and offer insights regarding how ranger patrols can be optimally deployed to reduce poaching-related threats and other illegal activites, while taking into account potential sampling biases. We show that poaching can be effectively reduced by increasing ranger patrols to areas under high risk of poaching activities, and by adding ranger patrols near these sites. These findings are broadly applicable to national parks and protected areas experiencing a high degree of poaching and other illegal activities.

Bushmeat hunting is widely cited as cause for declines of wildlife populations throughout Africa. Forest duikers (Bovidae, Cephalophinae) are among the most exploited species. Whether current harvest rates imperil duikers is debated because of the difficulty of accurately assessing population trends. To assess population trends, we first reviewed literature for historical duiker population estimates. Second, we used systematic camera-trap monitoring to assess population trends for 15 populations of nine duiker species in six national parks in Central and East Africa. We analysed annual monitoring data using Royle-Nichols heterogeneity-induced occupancy models to estimate abundance/sample point and derive occupancy estimates. Published density estimates indicate that duiker populations declined significantly throughout Africa between 1973 and 2013. There was a wide range of densities depending on species ((Formula presented.) range: 0.26–20.6 km−1) and whether populations were hunted ((Formula presented.) =6.3 km−1) or unhunted ((Formula presented.) = 16.3 km−1). More recent analysis of camera-trap monitoring produced different results. Estimated mean point abundance over time was between 0 and 0.99 individuals/point for four populations, between 1.0 and 1.99 for six populations, and greater than 2.0 for five populations. We observed five populations of duikers with negative trends in point abundances, although only one trend was significant and point abundance estimates for three populations were above 2.0 in the final survey year. Six populations showed positive trends in point abundance (three significant), and the remaining populations displayed no trends. Average occupancy was high (Ψ > 0.60) except for three populations. While literature indicates that historical population declines have occurred, most duiker populations appear relatively healthy in monitored parks. Our results indicate that these parks are effective in protecting most duikers despite hunting pressure. We recommend that systematic, standardized camera-trap monitoring be initiated in other African parks in combination with point-abundance models to objectively assess forest ungulate population trends.

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

We present the first cross continental comparison of the flowering and fruiting phenology of tropical forests across Africa. Flowering events of 5,446 trees from 196 species across 12 sites, and fruiting events of 4,595 trees from 191 species, across 11 sites were monitored over periods of 6 to 29 years, and analysed to describe phenology at the continental level. To study phenology we used Fourier analysis to identify the dominant cycles of flowering and fruiting for each individual tree and we identified the time of year African trees bloom and bear fruit and their relationship to local seasonality. Reproductive strategies were diverse and no single regular cycle was found in >50% of individuals across all 12 sites. Additionally, we found annual flowering and fruiting cycles to be the most common. Sub-annual cycles were the next most common for flowering whereas supra-annual patterns were the next most common for fruiting. We also identify variation in different subsets of species, with species exhibiting mainly annual cycles most common in West and West-Central African tropical forests, while more species at sites in East-Central and Eastern African forests showed cycles ranging from sub-annual to supra-annual. Despite many trees showing strong seasonality, at most sites some flowering and fruiting occurred all year round. Environmental factors with annual cycles are likely to be important drivers of seasonal periodicity in trees across Africa, but proximate triggers are unlikely to be constant across the continent

Populations of the endangered eastern chimpanzee (Pan troglodytes schweinfurthii) are declining throughout their range. Although Nyungwe National Park (NNP) harbors the largest remaining eastern chimpanzee population in Rwanda, we know little about their space use and dietary patterns. We studied home range, movement, and diet of two communities of chimpanzees in NNP using daily tracking data (6:00 am to 6:00 pm) collected from 2000 to 2015. One community, Mayebe, resided in the forest center, and the other community, Cyamudongo, inhabited a forest fragment located about 10 km from the main forest. Home range estimated with the 95% kernel density estimation (KDE) method was 21 km2 for the Mayebe community and 4 km2 for the Cyamudongo community. Chimpanzee home range sizes were smaller during the dry versus wet season and varied monthly throughout the year. The Mayebe community had an average hourly step length of 75 ± SE 5 m with a daily movement range of 987 ± SE 71 m, while the Cyamudongo community had a shorter hourly step length of 52 ± SE 3 m with a daily movement range of 651 ± SE 71 m. Both chimpanzee communities fed primarily on Ficus spp. Other important dietary items included fruits of Symphonia globulifera, Syzygium guineense, and Chrysophyllum gorungosanum for the Mayebe community and Trilepisium madagascariense for the Cyamudongo community. Food choice varied monthly and seasonally for each chimpanzee community. Our study provides the first estimates of home range size and movement parameters for chimpanzees in Rwanda and documents their food habits and seasonal variations therein. We also identified the 50% core home range for each chimpanzee community and suggest this area as the focus of management actions. These results could help park management reduce threats to chimpanzees and other sympatric species by improving the efficiency of ranger patrols.

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.

The elusive, Vulnerable owl-faced monkey Cercopithecus hamlyni is a rare and little studied species and one of the least known of the African Cercopithecidae. This study describes the distribution and relative abundance of the only known population in East Africa, in Nyungwe National Park, Rwanda. This species is restricted to a small (32 km2) area of bamboo and bamboo–forest mix in the southern sector of the Park, close to the international border with Burundi. We present the first empirical data of its abundance in the bamboo forests of Nyungwe. A total length of 185 km of transect were surveyed to estimate relative abundance of diurnal primates. Encounter rates with the owl-faced monkey were 0.081 groups km-1 (n = 15). Mean group size was 3.6 individuals. Eight independent photographs of C. hamlyni were obtained from five camera traps during 182 camera-days. Four other species of primates occur in the bamboo forest: eastern chimpanzee Pan troglodytes schweinfurthii, Angola colobus Colobus angolensis, L’Hoest’s monkey Cercopithecus l’hoesti and blue monkey Cercopithecus mitis. The main threats to the bamboo forest are from the illegal harvesting of bamboo, trapping and tree-cutting. These threats originate from both Rwanda and Burundi. There is an urgent need for conservation action to halt the destruction and degradation of the bamboo forest and to ensure the long-term survival of the owl-faced monkey in Rwanda.

We examined frugivore visitation and seed dispersal of five large-seeded (≥ 5 mm) tree species in tropical montane forest based on their occurrence in frugivorous primate diets: Ekebergia capensis, Olea capensis, Parinari excelsa, Prunus africana, and Syzygium guineense. A total of 21 frugivores in five assemblages (i.e., chimpanzees, cercopithecines, large-bodied birds, small-bodied birds, and squirrels) were observed over the study period (August 2006 and October–April 2007). We observed seed dispersal in four of five tree species studied; no dispersal was observed for P. excelsa. Frugivore assemblages did not visit tree species equally. Primates spent the most time in trees and had the largest group size. Large-bodied birds (LB) and chimpanzees dispersed the highest number of seeds per minute. LB and cercopithecines potentially dispersed the greatest number of seeds for E. capensis, and chimpanzees for S. guineense. Our analyses indicated that the mean fruiting duration of the focal tree, time in the tree, and number of species present are important predictor variables for seed dispersal by small- and large-bodied birds, and cercopithecines. The number of fruiting trees in the immediate vicinity of the focal tree further predicted seed dispersal for small-bodied birds (SB). Large-bodied birdseed dispersal also was predicted by time in tree by SB, and the number of individuals for SB and cercopithecines. Cercopithecines (CS) were further explained by the time in tree and number of species (SB & LB), and number of individuals for CS. Our study highlights the complexity of describing the relative importance of a frugivore assemblage to the dispersal of a tree species seeds. RESUME Nous avons examine comment se passent les visites par les frugivores et la dissemination des grandes graines (> 0.5 cm) de cinq especes d’arbres importants dans l’alimentation des primates dans la foret tropicale de montagne : Ekebergia capensis, Olea capensis, Parinari excelsa, Prunus africana, et Syzygium guineense. Au total 21 especes de frugivores classees en cinq groups (a savoir les chimpanzes, cercopithecines, grand-corps les oiseaux, les petits corps des oiseaux et des ecureuils) ont ete observees au cours de la periode d’etude (aout 2006 et octobre – avril 2007). Nous avons observe la dispersion des graines dans quatre des cinq especes d’arbres etudiees, aucune dispersion n’a ete observee pour P. excelsa. Different groupes de frugivores ont visite les especes d’arbres de differente facon. Les primates ont passe plus de temps dans les arbres et en plus grands groups. Les grands oiseaux (LB) et les chimpanzes ont disperse le plus grand nombre de graines par minute. LB et cercopithecines ont potentiellement disperse le plus grand nombre de graines de E. capensis, tandis que les chimpanzes celles de S. guineense. Nos analyses ont indique que la duree moyenne de fructification de l’arbre observe, le temps passe en arbre et le nombre d’especes presentes est d’importants variables predictifs de la dispersion des graines par les petits et les grands oiseaux ainsi que les cercopithecines. La fructification des arbres entourant l’arbre focal a en outre predit la dissemination des graines par les petits oiseaux (SB). La dissemination des graines par les LB a eteegalement predite par le temps que les SB ont passe dans un arbre et le nombre d’individus de ces SB et les cercopithecines. La dispersion des graines par les cercopithecines a en outre ete expliquee par le temps passe dans un arbre et le nombre d’especes (SB & LB), et la taille du group de cercopithecines. Notre etude met en evidence la complexite de decrire l’importance relative de differents frugivores dans la dispersion des graines des especes d’arbres.

With group sizes sometimes >300 individuals, the Angolan black-and-white colobus (Colobus angolensis ruwenzorii) population in Nyungwe Forest, Rwanda is an intriguing exception to the tendency for folivores to live in smaller groups than expected relative to body size. Researchers have hypothesized that the unusually high quality of foliage at Nyungwe allows colobus there to avoid intragroup feeding competition, releasing constraints on the formation of large groups (Fimbel et al., 2001). We collected data on the activity and ranging patterns of a >300-member Nyungwe colobus group and compared our results to those from smaller groups in other black-and-white colobus (Colobus spp.) populations. Colobus at Nyungwe spent far more time feeding and moving (62%) and far less time resting (32%) than black-and-white colobus at any other site. The annual home range of the Nyungwe colobus was also many times larger (95% minimum convex polygon: 20.7 km 2 ; 95% fixed kernel: 24.4 km 2 ) than those for other populations. We terminated our research after the group engaged in an unprecedented migration among black-and-white colobus by moving 13 km south of their former range. Our results suggest that intragroup scramble competition may be more intense than originally believed within the large colobus groups at Nyungwe and that long periods of resource renewal may be necessary after a large colobus group passes through an area, thereby potentially helping to explain their wide ranging patterns. We discuss the socioecological convergence between the Nyungwe colobus and Chinese snub-nosed monkeys (Rhinopithecus spp.) and suggest directions for future research on the unique black-and-white colobus population at Nyungwe.

Angolan black-and-white colobus monkeys (Colobus angolensis, Sclater 1980) in the Nyungwe Forest Reserve, Rwanda, are anomalous compared with conspecifics and congeners elsewhere, because they form exceptionally large groups, in excess of 300 individuals, and because of their unusually high consumption of mature leaves (∼40% of annual diet). Site factors at the Nyungwe Forest in southern Rwanda are believed to contribute to the unusual socio-ecology of the C. angolensis residing there. Foremost, the forest provides an abundance of mature leaves that are higher in nutritional quality (protein : fibre) compared with mature leaves analysed at other African sites. The combination of a cool high elevation locale, and a relatively open canopy, may contribute to the production of high quality mature leaves at ground level and in the canopy at the Nyungwe Forest. We believe that the almost unrestricted availability of acceptable foods for Nyungwe's folivorous colobines minimizes the potential for exploitation competition for food, and is a key factor in allowing them to form exceptionally large groups. Resume Les colobes noir et blanc d'Angola (Colobus angolensis, Sclater 1980) vivant dans la Reserve Forestiere de Nyungwe, au Rwanda, presentent des anomalies si on les compare aux autres membres de l'espece et leurs congeneres d'ailleurs, parce qu'ils forment des groupes exceptionnellement grands, depassant 300 individus, et a cause de leur consommation inhabituellement elevee de feuilles matures (≅40% du regime annuel). On pense que les caracteristiques du site de la Foret de Nyungwe, au sud du Rwanda, contribuent a la socio-ecologie inhabituelle des C. angolensis qui y vivent. Avant tout, cette la foret procure une abondance de feuilles matures de qualite nutritionnelle superieure (proteines et fibres) a celle relevee dans les feuilles matures analysees dans d'autres sites africains. La combinaison d'une altitude locale elevee avec temperature frai^che et d'une canopee relativement ouverte peut contribuer a la production de feuilles matures de haute qualite tant au niveau du sol que de la canopee de la Foret de Nyungwe. Nous pensons que la disponibilite quasi illimitee de nourriture acceptable pour les colobes mangeurs de feuilles de Nyungwe minimise la competition pour l'exploitation de la nourriture et qu'elle est un facteur cle dans la formation de ces groupes exceptionnellement nombreux.

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