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The critically endangered western gorillas (Gorilla gorilla) are divided into two subspecies: the western lowland (G. g. gorilla) and the Cross River (G. g. diehli) gorilla. Given the difficulty in sampling wild great ape populations and the small estimated size of the Cross River gorilla population, only one whole genome of a Cross River gorilla has been sequenced to date, hindering the study of this subspecies at the population level. In this study, we expand the number of whole genomes available for wild western gorillas, generating 41 new genomes (25 belonging to Cross River gorillas) using single shed hairs collected from gorilla nests. By combining these genomes with publicly available wild gorilla genomes, we confirm that Cross River gorillas form three population clusters. We also found little variation in genome-wide heterozygosity among them. Our analyses reveal long runs of homozygosity (>10 Mb), indicating recent inbreeding in Cross River gorillas. This is similar to that seen in mountain gorillas but with a much more recent bottleneck. We also detect past gene flow between two Cross River sites, Afi Mountain Wildlife Sanctuary and the Mbe Mountains. Furthermore, we observe past allele sharing between Cross River gorillas and the northern western lowland gorilla sites, as well as with the eastern gorilla species. This is the first study using single shed hairs from a wild species for whole genome sequencing to date. Taken together, our results highlight the importance of implementing conservation measures to increase connectivity among Cross River gorilla sites. This research was funded by Ministerio de Ciencia, Universidades e Investigación (FPI, PRE2018) (to M.A.-E.), Generalitat de Catalunya (FI_B100131) (to H.P.), European Research Council (ERC) under the European Union’s Horizon 2020 research and innovation programme (grant agreement No. 864203), PID2021-126004NB-100 (MICIIN/FEDER, UE), “Unidad de Excelencia María de Maeztu”, funded by the AEI (CEX2018-000792-M), NIH 1R01HG010898-01A1 and Secretaria d’Universitats i Recerca and CERCA Programme del Departament d’Economia i Coneixement de la Generalitat de Catalunya (GRC 2021 SGR 00177) (to T.M.B.). Illumina Inc. provided support for next-generation library sequencing, Beatriu de Pinós postdoctoral programme of the Government of Catalonia’s Secretariat for Universities and Research of the Ministry of Economy and Knowledge 2017 BP 00265 (to J.O.). The project that gave rise to these results received the support of a fellowship from “la Caixa” Foundation (ID 100010434) and from the European Union’s Horizon 2020 research and innovation programme under the Marie Skłodowska-Curie grant agreement (No 847648. The fellowship code is LCF/BQ/PI20/11760004) (to J.O.).

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.

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.

Wildlife species worldwide are under threat from a range of anthropogenic threats, with declines primarily caused by overexploitation and habitat loss associated with an increasing human population and per capita resource use. Exploitation is driven by numerous factors, but is often the result of illegal activities, such as hunting, logging, and wildlife trade. Protected areas, designed to safeguard threatened species and their habitats, are the foundation of biodiversity conservation, and several analyses have demonstrated that effective protected areas are critical to the maintenance of biodiversity. However, other analyses show that most protected areas suffer from a lack of resources and poor management. Numerous technologies have been developed to address these challenges by facilitating adaptive management via ranger-based data collection, data analysis and visualization, and strategic planning. This chapter reviews the Spatial Monitoring and Reporting Tool (SMART) platform, with a particular emphasis on conservation law enforcement monitoring, and demonstrates both how SMART has been used to improve management of conservation areas, and how complementary systems and emerging technologies can be integrated into a single unified platform for conservation area management. In a relatively short period of time, SMART has grown to become the global standard for conservation area management. More than 800 national parks and other conservation areas are currently using SMART in more than 65 countries worldwide. SMART sites have seen improvements in patrol effectiveness, increases in populations of critically endangered species like tigers, and reductions in threats from poaching and habitat loss.

Author(s): Cronin, Drew T.; Benbow, Sophie; Bergl, Richard A.; Bourgault, Liz; Caro, Lina; Dancer, Anthony; Davies, Alasdair; Joachim, Denton; Lewis, Claire; Lynam, Antony J.; Maaz, Julio; Muntifering, Jeff; O'Donnell, Stephanie; Palmer, Jonathan; Slade, James; Spina Avino, Felipe; Stevens, Xia; Wyatt, Alex | Abstract: Protected areas are key to biodiversity conservation and ranger-based monitoring, and law enforcement is the cornerstone upon which effective protected areas are built. Frontline practitioners, however, are often asked to protect large swathes of land or sea with limited resources, support, infrastructure, capacity, and/or training. Technology, when applied effectively and appropriately, has the capacity to empower practitioners, revolutionize ranger operations, improve ranger safety, and enhance wildlife protection and conservation outcomes. To do so, technology must be recognized, from the frontlines through to key decisionmakers, as a force multiplier, but only when it is fit for purpose, accessible, cost-effective, and supportive of rangers’ needs. In this paper we detail the general state of conservation technology and innovation within the ranger context and provide a series of detailed recommendations to help the Universal Ranger Support Alliance (URSA) meet the needs of rangers around the world, including: demystifying technology and clarifying what it can and cannot do, connecting the right technology with the right people and places, focusing technology development and investment on substantive improvements and support, broadening ranger familiarization with technology, building technology capacity in rangers, fostering greater community building and creating opportunities around technologies, engaging the technology sector to innovate and design technology to support rangers, and supporting technology as a complement to traditional knowledge and skills, rather than a replacement. These recommendations constitute an ambitious vision which cannot be delivered by URSA in isolation. Rather, we propose URSA leverages existing efforts to ensure rangers are supported around the world.

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.

Herpesviruses are thought to have evolved in very close association with their hosts. This is notably the case for cytomegaloviruses (CMVs; genus Cytomegalovirus) infecting primates, which exhibit a strong signal of co-divergence with their hosts. Some herpesviruses are however known to have crossed species barriers. Based on a limited sampling of CMV diversity in the hominine (African great ape and human) lineage, we hypothesized that chimpanzees and gorillas might have mutually exchanged CMVs in the past. Here, we performed a comprehensive molecular screening of all 9 African great ape species/subspecies, using 675 fecal samples collected from wild animals. We identified CMVs in eight species/subspecies, notably generating the first CMV sequences from bonobos. We used this extended dataset to test competing hypotheses with various degrees of co-divergence/number of host switches while simultaneously estimating the dates of these events in a Bayesian framework. The model best supported by the data involved the transmission of a gorilla CMV to the panine (chimpanzee and bonobo) lineage and the transmission of a panine CMV to the gorilla lineage prior to the divergence of chimpanzees and bonobos, more than 800,000 years ago. Panine CMVs then co-diverged with their hosts. These results add to a growing body of evidence suggesting that viruses with a double-stranded DNA genome (including other herpesviruses, adenoviruses, and papillomaviruses) often jumped between hominine lineages over the last few million years.

Bushmeat hunting is among the principal threats to larger-bodied vertebrate species, such as primates, in central Africa. However, species vary in their response to differing levels of hunting intensity, making it difficult to not only quantify the effect hunting has on wild populations, but also to plan for effective conservation interventions. In order to investigate the impact of hunting on primates on Bioko Island, Equatorial Guinea, we collected data on primate abundance and hunting intensity across three areas in the Gran Caldera Scientific Reserve (GCSR) via reconnaissance surveys. We assessed how the relative abundance and proportional representation of Bioko's seven diurnal primate species varied among the different sites and in relation to hunting intensity. Overall primate species richness and abundance were highest in areas where gun hunting was minimal, but both measures declined with increasing evidence of gun hunting. Two species of primates, Cercopithecus erythrotis and Cercopithecus nictitans, were relatively tolerant of heavy hunting, but all other species were considered vulnerable. Procolobus pennantii, a species endemic to Bioko, exhibited the highest vulnerability, and its range appears restricted to a very limited area in the southwest of the GCSR. Our analyses indicate that hunting on Bioko is a strong contributor to current patterns of primate abundance and diversity, and suggest that hunting may be driving Bioko's most threatened primates (e.g., P. pennantii) towards extinction. These results highlight the need for rapid intervention on Bioko in order to safeguard the future of its unique primates.

Technologies to identify individual animals, follow their movements, identify and locate animal and plant species, and assess the status of their habitats remotely have become better, faster, and cheaper as threats to the survival of species are increasing. New technologies alone do not save species, and new data create new problems. For example, improving technologies alone cannot prevent poaching: solutions require providing appropriate tools to the right people. Habitat loss is another driver: the challenge here is to connect existing sophisticated remote sensing with species occurrence data to predict where species remain. Other challenges include assembling a wider public to crowdsource data, managing the massive quantities of data generated, and developing solutions to rapidly emerging threats.

Genes encoded by the major histocompatibility complex (MHC) are crucial for the recognition and presentation of antigens to the immune system. In contrast to their closest relatives, chimpanzees and humans, much less is known about variation in gorillas at these loci. This study explored the exon 2 variation of -DPB1, -DQB1, and -DRB genes in 46 gorillas from four populations while simultaneously evaluating the feasibility of using fecal samples for high-throughput MHC genotyping. By applying strict similarity- and frequency-based analysis, we found, despite our modest sample size, a total of 18 alleles that have not been described previously, thereby illustrating the potential for efficient and highly accurate MHC genotyping from non-invasive DNA samples. We emphasize the importance of controlling for multiple potential sources of error when applying this massively parallel short-read sequencing technology to PCR products generated from low concentration DNA extracts. We observed pronounced differences in MHC variation between species, subspecies and populations that are consistent with both the ancient and recent demographic histories experienced by gorillas.

Repetitive movement patterns are commonly observed in zoo elephants. The extent to which these behaviors constitute a welfare concern varies, as their expression ranges from stereotypies to potentially beneficial anticipatory behaviors. Nevertheless, their occurrence in zoo animals is often viewed negatively. To better identify conditions that prompt their performance, observations were conducted on six African elephants (Loxodonta africana) at the North Carolina Zoo. Individuals spent most of their time engaged in feeding, locomotion, resting, and repetitive behavior. Both generalized estimating equation and zero-inflated negative binomial models were used to identify factors associated with increased rates of repetitive behavior. Time of day in conjunction with location on- or off-exhibit best explained patterns of repetitive behavior. Repetitive behaviors occurred at a lower rate in the morning when on-exhibit, as compared to afternoons on-exhibit or at any time of day off-exhibit. Increased repetitive behavior rates observed on-exhibit in the afternoon prior to the evening transfer and feeding were possibly anticipatory responses towards those events. In contrast, consistently elevated frequencies of repetitive behavior off-exhibit at all times of day could be related to differences in exhibit complexity between off-exhibit and on-exhibit areas, as well as a lack of additional foraging opportunities. Our study contributes valuable information on captive elephant behavior and represents a good example of how behavioral research can be employed to improve management of zoo animals. Zoo Biol. 34:201–210, 2015. © 2015 Wiley Periodicals Inc.

Human adenoviruses (HAdV; species HAdV-A to -G) are highly prevalent in the human population, and represent an important cause of morbidity and, to a lesser extent, mortality. Recent studies have identified close relatives of these viruses in African great apes, suggesting that some HAdV may be of zoonotic origin. We analyzed more than 800 fecal samples from wild African great apes and humans to further investigate the evolutionary history and zoonotic potential of hominine HAdV. HAdV-B and -E were frequently detected in wild gorillas (55%) and chimpanzees (25%), respectively. Bayesian ancestral host reconstruction under discrete diffusion models supported a gorilla and chimpanzee origin for these viral species. Host switches were relatively rare along HAdV evolution, with about ten events recorded in 4.5 My. Despite presumably rare direct contact between sympatric populations of the two species, transmission events from gorillas to chimpanzees were observed, suggesting that habitat and dietary overlap may lead to fecal-oral cross-hominine transmission of HAdV. Finally, we determined that two independent HAdV-B transmission events to humans occurred more than 100,000 years ago. We conclude that HAdV-B circulating in humans are of zoonotic origin and have probably affected global human health for most of our species lifetime.

Although captive populations of western gorilla have been maintained in the United States for over a century, little is known about the geographic origins and genetic composition of the current zoo population. Furthermore, although previous mitochondrial analyses have shown that free-range gorilla populations exhibit substantial regional differentiation, nothing is known of the extent to which this variation has been preserved in captive populations. To address these questions, we combined 379 pedigree records with data from 52 mitochondrial sequences to infer individual haplogroup affiliations, geographical origin of wild founders and instances of inter-breeding between haplogroups in the United States captive gorilla population. We show that the current captive population contains all major mitochondrial lineages found within wild western lowland gorillas. Levels of haplotype diversity are also comparable to those found in wild populations. However, the majority of captive gorilla matings have occurred between individuals with different haplogroup affiliations. Although restricting crosses to individuals within the same haplogroup would preserve the phylogeographic structure present in the wild, careful management of captive populations is required to minimize the risk of drift and inbreeding. However, when captive animals are released back into the wild, we recommend that efforts should be made to preserve natural phylogeographic structure.

Comparisons of MHC gene content and diversity among closely related species can provide insights into the evolutionary mechanisms shaping immune system variation. After chimpanzees and bonobos, gorillas are humans’ closest living relatives; but in contrast, relatively little is known about the structure and variation of gorilla MHC class I genes (Gogo). Here, we combined long-range amplifications and long-read sequencing technology to analyze full-length MHC class I genes in 35 gorillas. We obtained 50 full-length genomic sequences corresponding to 15 Gogo-A alleles, 4 Gogo-Oko alleles, 21 Gogo-B alleles, and 10 Gogo-C alleles including 19 novel coding region sequences. We identified two previously undetected MHC class I genes related to Gogo-A and Gogo-B, respectively, thereby illustrating the potential of this approach for efficient and highly accurate MHC genotyping. Consistent with their phylogenetic position within the hominid family, individual gorilla MHC haplotypes share characteristics with humans and chimpanzees as well as orangutans suggesting a complex history of the MHC class I genes in humans and the great apes. However, the overall MHC class I diversity appears to be low further supporting the hypothesis that gorillas might have experienced a reduction of their MHC repertoire. Electronic supplementary material The online version of this article (doi:10.1007/s00251-017-0974-x) contains supplementary material, which is available to authorized users.

Mount Cameroon (4095 m), the highest peak and only active volcano in West Africa, is located in the center of the Gulf of Guinea Pleistocene refugium. The associated forests and highlands along the southern Nigerian-Cameroon border and on the island of Bioko, known as the Biafran forests and highlands, are important formations of the Cameroon Volcanic Line owing to their wide elevational range, and on Mount Cameroon, a continuous gradient of unbroken vegetation from sea level to over 4000 m. The montane zones in the region begin 800 m above sea level forming the critically endangered Mount Cameroon and Bioko Montane Forests ecoregion. The broad elevational gradient of the region has resulted in high habitat diversity, leading the region to be a center for species endemism and richness across many taxa. Some of the densest human populations in Africa also occur in this region, putting intense pressure on the forests and highlands mostly due to overexploitation and habitat loss. The governments of ...

In the wild, western lowland gorillas travel long distances while foraging and consume a diet high in fiber and low in caloric density. In contrast, gorillas in zoos typically consume a diet that is low in fiber and calorically dense. Some items commonly used in captive gorilla diets contain high levels of starch and sugars, which are present at low levels in the natural diet of gorillas. Diet items high in simple carbohydrates are associated with obesity and heart disease in humans. Typical captive gorilla diets may also encourage undesirable behaviors. In response to these issues, we tested the behavioral impact of a diet that was biscuit-free, had low caloric density, and which was higher in volume at five institutions. We hypothesized that this diet change would reduce abnormal behaviors such as regurgitation and reingestion (R/R), decrease time spent inactive, and increase time spent feeding. The biscuit-free diet significantly reduced (and in the case of one zoo eliminated) R/R and may have reduced hair-plucking behavior. However, an increase in coprophagy was observed in many individuals following the diet change. The experimental diet caused a general increase in time the gorillas spent feeding, but this increase did not occur across all institutions and varied by individual. Interestingly, the overall time gorillas spent inactive actually increased with this diet change. Future research will examine these behavioral changes in a greater number of individuals to determine if the results remain consistent with these preliminary findings. Additionally, future research will examine the physiological impact of this diet change.

Ebola virus disease (EVD) is a threat to human health and to the survival of African great apes. The disease has led to major population declines in chimpanzees Pan troglodytes and gorillas Gorilla gorilla, and infected great apes play an important role as sources of human EVD outbreaks. The threat posed by EVD raises the question whether vaccination of wild apes is an effective strategy to reduce the occurrence and impact of this disease. We review the current knowledge about EVD in great apes and document the link between outbreaks in apes and in humans, mainly via bushmeat consumption. We discuss the need for control strategies, such as vaccination, and describe aspects of primate behaviour, virus biology, vaccine composition, and vaccination principles that need to be considered when making management decisions about great ape vaccination. Finally, we identify gaps in the understanding of Ebola ecology and highlight surveillance and research that can aid the survival of great apes and reduce human exposure to Ebola virus. The severe impact of EVD indicates the need for efficient monitoring and, ultimately, control of Ebola. However, the unknown reservoir and unpredictable emergence of Ebola, the elusive nature of great apes, and the lack of licensed and suitable vaccines represent major hurdles for such control. Public education about zoonotic diseases and monitoring of great ape health are both important strategies. Experts should also discuss the feasibility of developing safe vaccines that can be delivered efficiently to large populations of elusive wild apes in their natural remote habitats. This review provides a platform for further interdisciplinary discussions, so that management plans can be discussed and adjusted according to possible future changes in the development, availability and cost of vaccines, the status of EVD, knowledge about Ebola ecology, and opinion on wildlife vaccination.

Prado-Martinez, Javier et al., Most great ape genetic variation remains uncharacterized1,2; however, its study is critical for understanding population history3–6, recombination7, selection8 and susceptibility to disease9,10.Herewe sequence to high coverage a total of 79 wild- and captive-born individuals representing all six great ape species and seven subspecies and report 88.8 million single nucleotide polymorphisms. Our analysis provides support for genetically distinct populations within each species, signals of gene flow, and the split of common chimpanzees into two distinct groups: Nigeria–Cameroon/western and central/ eastern populations.Wefind extensive inbreeding in almost all wild populations, with eastern gorillas being the most extreme. Inferred effective population sizes have varied radically over timein different lineages and this appears to have a profound effect on the genetic diversity at, or close to, genes in almost all species. We discover and assign 1,982 loss-of-function variants throughout the human and great ape lineages, determining that the rate of gene loss has not been different in the human branch compared to other internal branches in the great ape phylogeny. This comprehensive catalogue of great ape genomediversity provides a framework for understanding evolution and a resource for more effective management of wild and captive great ape populations., We thank the following funding agencies: ERC Starting Grant (260372) to T.M.-B.; NIH grants HG002385 to E.E.E., R01_HG005226 to K.R.V., A.E.W., M.F.H., L.S. and J.D.W., GM100233 and NSF HOMINID grant 1032255 to D.R. and He.Li.;MICINN(Spain)BFU2011-28549to T.M.-B.,BFU2010-19443to Ja.Be.,Spanish Government and FEDER for grants BFU2009-13409-C02-02 and BFU2012-38236 to A.N. and J.P.-M., Direccio´ General de Recerca, Generalitat de Catalunya (Grup de Recerca Consolidat 2009 SGR 1101) to Ja.Be., D.C., A.N. and T.M.-B.; ERC Advanced Grant (233297) and Max Planck Society to S. Paabo; Danish Council for Independent Research Natural Sciences to H.S.; Spanish Grant (CGL-2010-20170) and Zoo de Barcelona (Beca PRIC) to A.R.-H.; EUPRIM-Net to BPRC; DP1ES022577-04 NIH grant to S.A.T.; NSF Grant 0755823 to M.K.G.; P.G. is supported by the G. Harold and Leila Y. Mathers Foundation. A.N. and T.M.-B. are ICREA Research Investigators (Institut Catala d’Estudis i RecercaAvancats delaGeneralitat deCatalunya). J.P.-M. is supported by the Zoo de Barcelona and l’Ajuntament de Barcelona.

It has long been hypothesized that polyomaviruses (PyV; family Polyomaviridae ) codiverged with their animal hosts. In contrast, recent analyses suggested that codivergence may only marginally influence the evolution of PyV. We reassess this question by focusing on a single lineage of PyV infecting hominine hosts, the Merkel cell polyomavirus (MCPyV) lineage. By characterizing the genetic diversity of these viruses in seven African great ape taxa, we show that they exhibit very strong host specificity. Reconciliation analyses identify more codivergence than noncodivergence events. In addition, we find that a number of host and PyV divergence events are synchronous. Collectively, our results support codivergence as the dominant process at play during the evolution of the MCPyV lineage. More generally, our results add to the growing body of evidence suggesting an ancient and stable association of PyV and their animal hosts. IMPORTANCE The processes involved in viral evolution and the interaction of viruses with their hosts are of great scientific interest and public health relevance. It has long been thought that the genetic diversity of double-stranded DNA viruses was generated over long periods of time, similar to typical host evolutionary timescales. This was also hypothesized for polyomaviruses (family Polyomaviridae ), a group comprising several human pathogens, but this remains a point of controversy. Here, we investigate this question by focusing on a single lineage of polyomaviruses that infect both humans and their closest relatives, the African great apes. We show that these viruses exhibit considerable host specificity and that their evolution largely mirrors that of their hosts, suggesting that codivergence with their hosts played a major role in their diversification. Our results provide statistical evidence in favor of an association of polyomaviruses and their hosts over millions of years.

Habitat loss and fragmentation are among the major threats to wildlife populations in tropical forests. Loss of habitat reduces the carrying capacity of the landscape and fragmentation disrupts biological processes and exposes wildlife populations to the effects of small population size, such as reduction of genetic diversity and increased impact of demographic stochasticity. The Critically Endangered Cross River gorillaGorilla gorilla diehliis threatened in particular by habitat disturbance because its population is small and it lives in an area where high human population density results in intense exploitation of natural resources. We used remotely-sensed data to assess the extent and distribution of gorilla habitat in the Cross River region and delineated potential dispersal corridors. Our analysis revealed > 8,000 km2of tropical forest in the study region, 2,500 km2of which is in or adjacent to areas occupied by gorillas. We surveyed 12 areas of forest identified as potential gorilla habitat, 10 of which yielded new records of gorillas. The new records expand the known range of the Cross River gorilla by > 50%, and support genetic analyses that suggest greater connectivity of the population than previously assumed. These findings demonstrate that considerable connected forest habitat remains and that the area could potentially support a much larger gorilla population if anthropogenic pressures such as hunting could be reduced.

In small and fragmented populations, genetic diversity may be reduced owing to increased levels of drift and inbreeding. This reduced diversity is often associated with decreased fitness and a higher threat of extinction. However, it is difficult to determine when a population has low diversity except in a comparative context. We assessed genetic variability in the critically endangered Cross River gorilla (Gorilla gorilla diehli), a small and fragmented population, using 11 autosomal microsatellite loci. We show that levels of diversity in the Cross River population are not evenly distributed across the three genetically identified subpopulations, and that one centrally located subpopulation has higher levels of variability than the others. All measures of genetic variability in the Cross River population were comparable to those of the similarly small mountain gorilla (G. beringei beringei) populations (Bwindi and Virunga). However, for some measures both the Cross River and mountain gorilla populations show lower levels of diversity than a sample from a large, continuous western gorilla population (Mondika, G. gorilla gorilla). Finally, we tested for the genetic signature of a bottleneck in each of the four populations. Only Cross River showed strong evidence of a reduction in population size, suggesting that the reduction in size of this population was more recent or abrupt than in the two mountain gorilla populations. These results emphasize the need for maintaining connectivity in fragmented populations and highlight the importance of allowing small populations to expand. Am. J. Primatol. 70:848–859, 2008. © 2008 Wiley-Liss, Inc.

The forests and highlands along the southern portion of the Nigeria-Cameroon border and on the island of Bioko have long been recognized as being biologically diverse. This region (referred to as the Biafran forests and highlands) is a center of endemism for a wide variety of taxa including, but not limited to, primates, anuran amphibians, birds, freshwater fish, butterflies, dragonflies, and trees. Though these groups have diverse distributions, conservation efforts have to date largely been focused on lowland areas. We conducted a GIS-based analysis of point locality records for three groups characterized by high endemism (primates, anuran amphibians, birds) in order to examine both their spatial and altitudinal distribution throughout the study area. We also evaluated the distribution of existing and potential protected areas relative to highland areas and the distribution of endemics. Our analysis suggests that the existing protected area system provides poor coverage of montane habitats and their associated endemic taxa. Complementarity analysis suggests that, if the protected area network were expanded to include a small number of highland sites, coverage of endemic taxa could be significantly improved. Many of these important highland sites are currently under intense pressure from habitat loss and hunting. If the full range of biodiversity present in the Biafran forests and highlands is to be preserved, new protected areas should be gazetted that take the varied distributions of the regions endemic taxa into account.

Genes encoded by the major histocompatibility complex (MHC) are crucial for the recognition and presentation of antigens to the immune system. In contrast to their closest relatives, chimpanzees and humans, much less is known about variation in gorillas at these loci. This study explored the exon 2 variation of -DPB1, -DQB1, and -DRB genes in 46 gorillas from four populations while simultaneously evaluating the feasibility of using fecal samples for high-throughput MHC genotyping. By applying strict similarity- and frequency-based analysis, we found, despite our modest sample size, a total of 18 alleles that have not been described previously, thereby illustrating the potential for efficient and highly accurate MHC genotyping from non-invasive DNA samples. We emphasize the importance of controlling for multiple potential sources of error when applying this massively parallel short-read sequencing technology to PCR products generated from low concentration DNA extracts. We observed pronounced differences in MHC variation between species, subspecies and populations that are consistent with both the ancient and recent demographic histories experienced by gorillas.

Population estimates using genetic capture–recapture methods from non-invasively collected wildlife samples are more accurate and precise than those obtained from traditional methods when detection and resampling rates are high. Recently, detection dogs have been increasingly used to find elusive species and their by-products. Here we compared the effectiveness of dog- and human-directed searches for Cross River gorilla (Gorilla gorilla diehli) faeces at two sites. The critically endangered Cross River gorilla inhabits a region of high biodiversity and endemism on the border between Nigeria and Cameroon. The rugged highland terrain and their cryptic behaviour make them difficult to study and a precise population size for the subspecies is still lacking. Dog-directed surveys located more fresh faeces with less bias than human-directed survey teams. This produced a more reliable population estimate, although of modest precision given the small scale of this pilot study. Unfortunately, the considerable costs associated with use of the United States-based detection dog teams make the use of these teams financially unfeasible for a larger, more comprehensive survey. To realize the full potential of dog-directed surveys and increase cost-effectiveness, we recommend basing dog-detection teams in the countries where they will operate and expanding the targets the dogs are trained to detect.

Recently developed methods of individual-based analysis of genetic data allow an unprecedented opportunity to understand the relationships among fragmented populations. By defining population structure and identifying migrant individuals, such analyses can provide a framework to aid in evaluating the threats posed by inbreeding and reduced genetic variability as a consequence of limited gene flow among fragments. Here we investigate population structure in the critically endangered Cross River gorilla ( Gorilla gorilla diehli ) by applying a suite of individual-based analyses to data obtained from between onequarter and one-third of the estimated total population through the use of noninvasively collected DNA samples. The population structure inferred using data from 11 autosomal microsatellite loci was broadly consistent with geography and habitat fragmentation, but showed no simple isolation-by-distance effects. In contrast to previous field surveys, which suggested that all gorilla localities were isolated from one another, we infer low levels of gene flow and identify migrants between habitat fragments as well as individuals of admixed ancestry, suggesting persistent recent reproductive contact between many of the localities. These results are encouraging for the conservation of the Cross River gorilla population. Conservation efforts should strive to maintain connectivity between subpopulations that are still in migratory contact and attempt to restore connectivity where it has been lost.

Films, as part of a larger environmental education program, have the potential to influence the knowledge and attitudes of viewers. However, to date, no evaluations have been published reporting the effectiveness of films, when used within primate range countries as part of a conservation themed program. The Great Ape Education Project was a year-long environmental education program implemented in Uganda for primary school students living adjacent to Kibale National Park (KNP) and Bwindi Impenetrable National Park (BINP). Students viewed a trilogy of conservation films about great apes, produced specifically for this audience, and participated in complementary extra-curricular activities. The knowledge and attitudes of students participating in the program from KNP, but not BINP were assessed using questionnaires prior to (N = 1271) and following (N = 872) the completion of the program. Following the program, students demonstrated a significant increase in their knowledge of threats to great apes and an increase in their knowledge of ways that villagers and students can help conserve great apes. Additionally, student attitudes toward great apes improved following the program. For example, students showed an increase in agreement with liking great apes and viewing them as important to the environment. These data provide evidence that conservation films made specifically to address regional threats and using local actors and settings can positively influence knowledge of and attitudes toward great apes among students living in a primate range country.

Repetitive movement patterns are commonly observed in zoo elephants. The extent to which these behaviors constitute a welfare concern varies, as their expression ranges from stereotypies to potentially beneficial anticipatory behaviors. Nevertheless, their occurrence in zoo animals is often viewed negatively. To better identify conditions that prompt their performance, observations were conducted on six African elephants (Loxodonta africana) at the North Carolina Zoo. Individuals spent most of their time engaged in feeding, locomotion, resting, and repetitive behavior. Both generalized estimating equation and zero-inflated negative binomial models were used to identify factors associated with increased rates of repetitive behavior. Time of day in conjunction with location on- or off-exhibit best explained patterns of repetitive behavior. Repetitive behaviors occurred at a lower rate in the morning when on-exhibit, as compared to afternoons on-exhibit or at any time of day off-exhibit. Increased repetitive behavior rates observed on-exhibit in the afternoon prior to the evening transfer and feeding were possibly anticipatory responses towards those events. In contrast, consistently elevated frequencies of repetitive behavior off-exhibit at all times of day could be related to differences in exhibit complexity between off-exhibit and on-exhibit areas, as well as a lack of additional foraging opportunities. Our study contributes valuable information on captive elephant behavior and represents a good example of how behavioral research can be employed to improve management of zoo animals.

We conducted a review of the wildlife conservation literature to identify broad trends in the publishing record and focal areas of research over the past 20 years. A total of 5,853 papers were reviewed with an emphasis on decadal changes between 1993, 2002, and 2012. For each paper we identified the journal and common keywords, and also determined the research scope, conservation issues and applications, and geographic focus. We found that both the number of journals publishing in the field, as well as the number of published articles, has increased significantly over time. The proportional contribution of the most prominent journals in the field has decreased over time, but not the importance of the articles within those journals. Previously reported biases in the literature towards mammals and birds, persisted in our study, leaving large proportions of globally threatened taxa (e.g. amphibians) underrepresented. There was also a disparity in the number of publications from particular geographic regions, however, the proportional contribution of under-represented geographic regions (e.g., Central & South America) increased over time. Finally, using the prevalence of keywords, we identified wildlife/adaptive management, hunting/bushmeat, and human wildlife conflict as contemporary (1998-2012) research priorities. The persistence of biases towards charismatic taxa can hinder conservation efforts, and we suggest that researchers refocus their efforts towards vulnerable regions and taxa in order to better address conservation priorities. The aim of wildlife conservation is to support the preservation of biological diversity in the natural

Recently developed methods of individual-based analysis of genetic data allow an unprecedented opportunity to understand the relationships among fragmented populations. By defining population structure and identifying migrant individuals, such analyses can provide a framework to aid in evaluating the threats posed by inbreeding and reduced genetic variability as a consequence of limited gene flow among fragments. Here we investigate population structure in the critically endangered Cross River gorilla (Gorilla gorilla diehli) by applying a suite of individual-based analyses to data obtained from between one-quarter and one-third of the estimated total population through the use of noninvasively collected DNA samples. The population structure inferred using data from 11 autosomal microsatellite loci was broadly consistent with geography and habitat fragmentation, but showed no simple isolation-by-distance effects. In contrast to previous field surveys, which suggested that all gorilla localities were isolated from one another, we infer low levels of gene flow and identify migrants between habitat fragments as well as individuals of admixed ancestry, suggesting persistent recent reproductive contact between many of the localities. These results are encouraging for the conservation of the Cross River gorilla population. Conservation efforts should strive to maintain connectivity between subpopulations that are still in migratory contact and attempt to restore connectivity where it has been lost.

In this chapter we summarize information on the morphology, genetics, and natural history of the West African gorilla population inhabiting the forests on the Nigeria–Cameroon border at the northern headwaters of the Cross River, a region at the western and northern limits of the species' range. A recent morphological analysis of skeletal specimens from this population has shown that they are sufficiently distinct from other western gorillas to justify being classified as the subspecies Gorilla gorilla diehli , a taxonomic name originally applied to them in the early twentieth century (Sarmiento and Oates, 2000). Just as the distinctiveness of the Cross River gorillas is being appreciated, their continued survival is in jeopardy. Recent surveys suggest that approximately 250 probably remain, concentrated in nine or more isolated hill areas. Because these gorillas are still hunted for their meat and parts of their habitat are under threat, they are one of Africa's most endangered primate taxa. After reviewing data on the status of the Cross River gorillas, our chapter ends by discussing some options for improving their prospects for survival. In addition to the literature, the information we summarize derives from our own research: Field surveys in Nigeria (by JFO and KLM) and Cameroon (by JLG); an ecological study of a subpopulation inhabiting Afi Mountain Wildlife Sanctuary, Nigeria (by KLM); and the sequencing of mtDNA extracted from hairs of Nigerian gorillas shed into sleeping nests (by RAB and JML in the laboratory of TRD). All our results should be regarded as preliminary.