Virunga Massif Research Articles

Comparative life history patterns of female gorillas.

Several theories have been proposed to explain the impact of ecological conditions on differences in life history variables within and between species. Here we compare female life history parameters of one western lowland gorilla population (Gorilla gorilla gorilla) and two mountain gorilla populations (Gorilla beringei beringei). We compared the age of natal dispersal, age of first birth, interbirth interval, and birth rates using long-term demographic datasets from Mbeli Bai (western gorillas), Bwindi Impenetrable National Park and the Virunga Massif (mountain gorillas). The Mbeli western gorillas had the latest age at first birth, longest interbirth interval, and slowest surviving birth rate compared to the Virunga mountain gorillas. Bwindi mountain gorillas were intermediate in their life history patterns. These patterns are consistent with differences in feeding ecology across sites. However, it is not possible to determine the evolutionary mechanisms responsible for these differences, whether a consequence of genetic adaptation to fluctuating food supplies ("ecological risk aversion hypothesis") or phenotypic plasticity in response to the abundance of food ("energy balance hypothesis"). Our results do not seem consistent with the extrinsic mortality risks at each site, but current conditions for mountain gorillas are unlikely to match their evolutionary history. Not all traits fell along the expected fast-slow continuum, which illustrates that they can vary independently from each other ("modularity model"). Thus, the life history traits of each gorilla population may reflect a complex interplay of multiple ecological influences that are operating through both genetic adaptations and phenotypic plasticity.

Projecting the impact of an ebola virus outbreak on endangered mountain gorillas

Ebola virus is highly lethal for great apes. Estimated mortality rates up to 98% have reduced the global gorilla population by approximately one-third. As mountain gorillas (Gorilla beringei beringei) are endangered, with just over 1000 individuals remaining in the world, an outbreak could decimate the population. Simulation modeling was used to evaluate the potential impact of an Ebola virus outbreak on the mountain gorilla population of the Virunga Massif. Findings indicate that estimated contact rates among gorilla groups are high enough to allow rapid spread of Ebola, with less than 20% of the population projected to survive at 100 days post-infection of just one gorilla. Despite increasing survival with vaccination, no modeled vaccination strategy prevented widespread infection. However, the model projected that survival rates greater than 50% could be achieved by vaccinating at least half the habituated gorillas within 3 weeks of the first infectious individual.

Food availability influences birth seasonality at a small spatial scale in endangered golden monkeys (Cercopithecus mitis kandti).

Studying reproductive seasonality helps us understand changes in the energetically demanding periods of pregnancy and lactation. We investigated how diet variability and key food plant phenology relate to mating and birth seasonality in both remaining populations of endangered golden monkeys in the Virunga massif and the Gishwati forest in Rwanda. Using scan and ad libitum sampling, we recorded feeding, mating, and births in two social groups (K and M) living in Volcanoes National Park (VNP), in the Virunga massif, from 2004 to 2018, and in one group (G) in the Gishwati forest in 2017-2018. We also monitored bamboo shoot and fruit availability in the groups' home ranges in 2017-2018. Mating was observed year-round but peaked four to 6 months prior to each group's respective birth season. Despite the two VNP groups ranging only 16 km apart, they had different birth seasons. Females from group K gave birth during the late bamboo growing season, from September to December, while females from group M gave birth during the early bamboo shooting season, from February to April. This pattern was linked to differences in the availability and consumption of bamboo shoots between low-elevation (group K) and high-elevation (group M) habitat. In group G, births occurred from March to April, coinciding with the period of high fruit availability and consumption in the Gishwati forest. Our findings suggest that food availability shapes golden monkey birth seasons, even at a small spatial scale. Current changes in key food plant regeneration, potentially driven by climate change, need to be closely monitored to inform golden monkey conservation management.

Distribution and conservation status of the golden monkeyCercopithecus mitis kandtiin Rwanda

AbstractHabitat loss and fragmentation are major threats to primate populations globally. The Endangered golden monkeyCercopithecus mitis kandtiis only found in two small forest fragments: the Virunga massif in Rwanda, Uganda and the Democratic Republic of the Congo, and the Gishwati Forest in Rwanda. Little is known about the abundance and distribution of this subspecies, or threats to its survival. During 2007–2018, we collected data along 893.7 km of line transects and 354.2 km of recce trails in Volcanoes National Park and in Gishwati–Mukura National Park to estimate golden monkey density and examine any threats. In Volcanoes National Park, golden monkeys were found almost exclusively in the bamboo zone, and in Gishwati–Mukura National Park they occurred only in the remnant tropical montane Gishwati Forest. In Volcanoes National Park, density was estimated to be 7.89 (95% CI: 3.85–16.19), 5.41 (2.64–11.08), and 5.47 (3.68–8.14) groups per km2in 2007, 2011 and 2017–2018, respectively. This corresponds to a total of 4,331 individuals (95% CI: 2,723–5,938) in 2007, 4,487 (2,903–6,071) in 2011 and 4,626 (4,165–5,088) in 2017–2018. In Gishwati Forest, group density averaged 1.98 (95% CI: 1.27–3.16) per km2, corresponding to 172 (95% CI: 154–190) individuals in 2017–2018. Survey results from Volcanoes National Park suggest that the golden monkey population has been stable during 2007–2018. Limited habitat, illegal activities such as harvesting of bamboo and firewood, and the presence of feral dogs, threaten the golden monkey in Rwanda and require continued monitoring. The development of a conservation action plan is a priority to protect this species.

Ecological drivers of helminth infection patterns in the Virunga Massif mountain gorilla population.

The Virunga Massif mountain gorilla population has been periodically monitored since the early 1970s, with gradually increasing effort. The population declined drastically in the 1970s, but the numbers stabilized in the 1980s. Since then, the population has been steadily increasing within their limited habitat fragment that is surrounded by a dense human population. We examined fecal samples collected during the Virunga 2015–2016 surveys in monitored and unmonitored gorilla groups and quantified strongylid and tapeworm infections using egg counts per gram to determine environmental and host factors that shape these helminth infections. We showed that higher strongylid infections were present in gorilla groups with smaller size of the 500-m buffered minimum-convex polygon (MCP) of detected nest sites per gorilla group, but in higher gorilla densities and inhabiting vegetation types occurring at higher elevations with higher precipitation and lower temperatures. On the contrary, the impact of monitoring (habituation) was minor, detected in tapeworms and only when in the interaction with environmental variables and MCP area. Our results suggest that the Virunga mountain gorilla population may be partially regulated by strongylid nematodes at higher gorilla densities. New health challenges are probably emerging among mountain gorillas because of the success of conservation efforts, as manifested by significant increases in gorilla numbers in recent decades, but few possibilities for the population expansion due to limited amounts of habitat.

What factors affect species richness and distribution dynamics within two Afromontane protected areas?

Context Biodiversity monitoring programs provide information on the status and trends in wildlife populations. These trends are unknown for most mammals within African montane forests, which harbour many endemic and threatened species. Camera traps are useful for assessing mammal populations, because they allow for the estimation of species richness, occupancy, and activity patterns. Aims We sought to explore the richness and distribution of small- to large-sized mammals by using occupancy models while accounting for imperfect detection in Volcanoes and Mgahinga Gorilla National Parks, in Rwanda and Uganda. Methods We used camera-trap data collected from 2014 to 2017 by the Tropical Ecology Assessment and Monitoring (TEAM) network and multi-season occupancy models with multispecies data to assess the dynamics of species richness and distribution in the Virunga Massif and the influence of site covariates on species detection probability, occupancy, colonisation and extinction. Key results We identified 17 species from 7047 trap-days, with most of them showing an uneven distributional pattern throughout the park. We found that average species richness per site increased from five to seven species in 2017. Average local colonisation was estimated at 0.13 (s.e. 0.014), but the probability of local extinction was 0.17 (s.e. 0.028) and negatively influenced by distance from the park boundary. Detection probability was highest for medium-sized species. For species distribution, we found that black-fronted duiker, Cephalophus nigrifrons, and bushbuck, Tragelaphus scriptus, declined in distribution but remained widespread in our study area, while all other species showed an increase in distribution over the study period. Conclusions Our approach allowed us to draw inferences on rare species, such as African golden cat, Caracal aurata, by estimating detection probability on the basis of shared covariate information with more common, widespread species. As such, we were able to estimate all occupancy parameters across the terrestrial mammal community. Implications The results of this study on the distribution of terrestrial mammal species can be used by park management to inform optimal ranger patrolling efforts for mitigating threats in areas of high species presence. Additionally, the results can highlight locations of potential human–wildlife conflicts on the basis of species commonly found along the park boundary.

Population structure, intergroup interaction, and human contact govern infectious disease impacts in mountain gorilla populations.

Infectious zoonotic diseases are a threat to wildlife conservation and global health. They are especially a concern for wild apes, which are vulnerable to many human infectious diseases. As ecotourism, deforestation, and great ape field research increase, the threat of human-sourced infections to wild populations becomes more substantial and could result in devastating population declines. The endangered mountain gorillas (Gorilla beringei beringei) of the Virunga Massif in east-central Africa suffer periodic disease outbreaks and are exposed to infections from human-sourced pathogens. It is important to understand the possible risks of disease introduction and spread in this population and how human contact may facilitate disease transmission. Here we present and evaluate an individual-based, stochastic, discrete-time disease transmission model to predict epidemic outcomes and better understand health risks to the Virunga mountain gorilla population. To model disease transmission we have derived estimates for gorilla contact, interaction, and migration rates. The model shows that the social structure of gorilla populations plays a profound role in governing disease impacts with subdivided populations experiencing less than 25% of the outbreak levels of a single homogeneous population. It predicts that gorilla group dispersal and limited group interactions are strong factors in preventing widespread population-level outbreaks of infectious disease after such diseases have been introduced into the population. However, even a moderate amount of human contact increases disease spread and can lead to population-level outbreaks.

High flexibility in diet and ranging patterns in two golden monkey (Cercopithecus mitis kandti) populations in Rwanda.

Many primates exhibit behavioral flexibility which allows them to adapt to environmental change and different habitat types. The golden monkey (Cercopithecus mitis kandti) is a little-studied endangered primate subspecies endemic to the Virunga massif and the Gishwati forest in central Africa. In the Virunga massif, golden monkeys are mainly found in the bamboo forest, while in the Gishwati forest they live in mixed tropical montane forest. Here we describe and compare the diet of golden monkeys in both fragments. Over 24 consecutive months from January 2017 we used scan sampling to record feeding and ranging behavior of two Virunga groups and one Gishwati group totaling ca. 240 individuals. We also examined the phenology of bamboo and fruit trees, key seasonal food plant species for the monkeys. Golden monkeys fed on more than 100 plant species. The Virunga groups were mostly folivorous (between 72.8% and 87.16% of the diet) and fed mostly on young bamboo leaves and bamboo shoots, while 48.69% of the diet of the Gishwati group consisted of fruit from 22 different tree and shrub species. Bamboo shoots and fruit are seasonally available foods and were consumed regularly throughout the period when they were available. Despite being the smallest of the three study groups, the Gishwati group had a larger home range area (150.07 ha) compared to both Virunga groups (25.24 and 91.3 ha), likely driven by the differences in availability and distribution of fruit and bamboo in the habitats. Like other blue monkey subspecies, golden monkeys appear to have a flexible dietary strategy enabling them to adjust diet and ranging behavior to local habitats and available food resources. Additional studies and continuing conservation efforts are needed to better understand how variation in feeding and ranging ecology affects reproduction, population growth, and carrying capacity.

Heterogeneity in patterns of helminth infections across populations of mountain gorillas (Gorilla beringei beringei)

Conservation efforts have led to the recovery of the endangered mountain gorilla populations. Due to their limited potential for spatial expansion, population densities increased, which may alter the epidemiology of infectious diseases. Recently, clinical gastrointestinal illnesses linked to helminth infections have been recorded in both gorilla populations. To understand drivers and patterns of helminth infections we quantified strongylid and tapeworm infections across both Virunga Massif and Bwindi populations using fecal egg counts. We assessed the impact of age, sex, group size, season and spatial differences used as a proxy, which reflects observed variation in the occurrence of gastrointestinal problems, vegetation types, gorilla subpopulation growth and associated social structure on helminth infections. We revealed striking geographic differences in strongylid infections with higher egg counts mostly in areas with high occurrences of gastrointestinal disease. Increased helminth egg counts were also associated with decreasing group size in some areas. Observed spatial differences may reflect mutual effects of variations in subpopulation growth rates, gorilla social structure, and vegetation associated with altitude across mountain gorilla habitat. Helminth infection intensities in Virunga gorillas were lowest in the youngest and the oldest animals. Elucidating parasite infection patterns of endangered species with low genetic diversity is crucial for their conservation management.

Daily defecation outputs of mountain gorillas (Gorilla beringei beringei) in the Volcanoes National Park, Rwanda.

Increasing population density can increase infectious disease risk and thus reduce population growth and size. Host-parasite interactions of threatened animals that remain in small protected forest fragments therefore need to be monitored carefully. Due to extreme conservation efforts, the mountain gorilla (Gorilla beringei beringei) population in the 450-km2 Virunga Massif has more than doubled since 1973, reaching 604 individuals in 2016. To better understand changes in the transmission risks of soil-borne and other enteric pathogens for mountain gorillas, we determined defecation outputs of different age-sex classes and the diurnal variation in feces deposition. We weighed 399 wet fecal samples deposited at nest sites and on trails between nest sites by gorillas of varying age and sex, determined by lobe diameter, from five social groups (n = 58 gorillas) that range in the Volcanoes National Park, Rwanda. We found increasing daily average defecation outputs with increasing age-sex class (infants, 435g; juveniles, 1346g; medium-sized gorillas, 2446g; silverbacks, 3609g). Gorillas deposited two- to threefold the amount of feces at nest sites compared to on trails, suggesting that nest sites may function as hotspots for enteric pathogen infections through direct contact or when gorillas ingest foods contaminated with infectious larvae during site revisits in intervals matching the maturation period of environmentally transmitted gastrointestinal parasites. In combination with ranging and demographic data, these findings will enable the modeling of spatiotemporal variation in soil contamination and infectious disease risk for Virunga gorillas as their population density continues to increase.

ANESTHESIA WITH MEDETOMIDINE-KETAMINE AND DEXMEDETOMIDINE-KETAMINE IN MOUNTAIN GORILLAS (GORILLA BERINGEI BERINGEI).

Between December 2002 and September 2017, 125 anesthetic procedures involving free-living and orphaned captive mountain gorillas (Gorilla beringei beringei) were performed in the Virunga Massif and Bwindi Impenetrable Forest in East-Central Africa. Of these 125 immobilizations, 114 records were complete enough for inclusion into this study. Anesthetic and physiologic data from these 114 cases were analyzed, of which 57 used medetomidine-ketamine and 57 used dexmedetomidine-ketamine administered intramuscularly. With the use of estimated weights, the mean induction dosage (mg/kg ± SD) for medetomidine was 0.033 ± 0.003 (n = 42), for dexmedetomidine 0.018 ± 0.005 (n = 53), and for ketamine 3.66 ± 0.95 (n = 95). Mean time from injection of induction dose to recumbency was 6.8 ± 3.1 min (n = 74). Atipamezole was administered intramuscularly to reverse anesthesia. First signs of recovery occurred at 5.0 ± 4.0 min, and full recovery was 19.0 ± 17.0 min after administration of the reversal agent. No significant differences in physiologic parameters or anesthetic time variables were noted between healthy and unhealthy individuals. Mean heart rate was 72.0 ± 17.6 beats/min (n = 83) and mean oxygen saturation was 96.5% ± 4.2 (n = 62). Mean respiratory rate was 27 ± 9 breaths/min (n = 84) and mean body temperature 36.6°C ± 1.2 (n = 61). The current protocol has several advantages for field use in this species given its quick induction, few observed side effects, and ability to reverse so that the animal can return more quickly to its social group.

Integrating Remote Sensing and GIS to Model Forest Fire Rik in Virunga Massif, Central - Eastern Africa

This study aimed at developing a forest fire risk model using a combination of GIS and Remote sensing techniques, which helped to identify the level of forest fire vulnerability in Virunga Massif, located at the edge of central and eastern Africa. The Analytic Hierarchical Process (AHP) approach was employed to rank and weigh the key variables and combine them into different fire risk input factors which were later integrated into the main forest fire risk model. The main input datasets, which were linked with a potential source of a forest fire, include the land cover (specifically vegetation type data generated through the Landsat 8 image classification); topographic variables such as slope, elevation and aspect retrieved from the existing Digital Elevation Model (DEM) of Rwanda; the concentration of illegal activities and proximity to beehives sites; as well as visibility from the road and human settlements. Input factor maps were generated, assigned weights and combined in a GIS environment to produce a Virunga massif fire risk model map, which was validated using the existing burnt areas map, and ground truth points recorded using GPS. The study found that the ignition factors are the most forest fire triggering factors in Virunga massif, followed by topographic factors which play a major role in the fire spreading across the ecosystem. The high forest fire risk areas were found in steep slope location around the peaks of the volcanoes, whereas areas with the lowest risk of forest fire were found inside the forest at gentle slopes. The model was validated at 75% accuracy using ground truth data. The study proposes measure to halt the ignition factors through prevention of illegal activities in the Virunga massif for the successful prevention of the forest fire risk in the ecosystem, with much effort invested during the dry season, along with the relocation of beehives to a farther distance from the ecosystem’s edge.
 Keywords: Forest Fire Risk Modelling, Biodiversity, Illegal Activities, Ignition Factors, Topographic Factors, Analytic Hierarchy Process

Abundance and spatial distribution of the main food species for mountain gorillas in the Virunga Massif, Rwanda

The conservation of endangered species can benefit from a clear understanding of the quantity and distribution of their main foods. The population of mountain gorillas (Gorilla beringei beringei) living in the Virunga Massif of Rwanda, Uganda, and Democratic Republic of Congo has doubled in size since the 1980s, due to success in conservation efforts in and around their habitat. However, this increase in population size along with pressures on gorilla habitat raises concerns about spatial-temporal changes in the gorillas’ food plants. This study modelled the abundance and distribution of gorilla food species in the Virunga Massif. A total of 1050 vegetation recordings were collected on five plant species that are known to be frequently consumed by gorillas in one region of the Virungas, the Karisoke area. Two types of datasets collected along vegetation zones were combined: one with plant abundance expressed with Braun-Blanquet scores; and the other with abundance expressed as biomass. Moreover, ecological characteristics of locations where these species occur were extracted from satellite imagery. Analysis of variance and linear regression models were used to examine relationships between food species abundances and predictor variables. Subsequently, maps for the food species were created using boosted regression trees (BRTs). The abundance of species differed across vegetation zones, and the differences were statistically significant among vegetation zones with enough species observations. The accuracy of the BRTs indicated greater than random predictions (AUC > 0.65). This study shows the suitable areas for these gorilla food species and relevant ecological variables determining their distribution. The results provide insights into habitat occupancy by mountain gorillas, and help to design a baseline for monitoring changes in the abundance of gorilla food species under changing climate and anthropogenic pressure.

Effect of mountain gorilla (<i>Gorilla beringei beringei</i>) population growth to their key food plant biomass in Volcanoes National Park, Rwanda

High densities of large herbivores can have detrimental effects on plant biomass. Understanding the relationship between animal densities and plant distribution and abundance is essential for the conservation of endangered species and ecosystems. Mountain gorilla censuses conducted for different periods in the last three decades have revealed a steady increase of gorilla population in Virunga Massif whereby the recent number of gorillas has doubled compared to their number in the 1980s. It is unclear whether the continuous population growth of the herbivorous Virunga gorilla within an isolated forest ‘island’ has been affecting gorilla food plant biomass. This study investigated the effect of varying mountain gorilla densities on the biomass of the five key food plant species (Galium spp., Carduus nyassanus, Peucedanum linderi, Rubus spp., Laportea alatipes) that make up >70% of the mountain gorilla diet. We used plant biomass data collected in a central part of the Virunga massif, commonly known as Karisoke sector from 2009 to 2011, and GPS records of gorilla groups ranging in the same area nine months prior biomass assessment. Gorilla densities were estimated using the Kernel Utilization Distribution (KDE) analysis (functions: ‘kernelUD’ and ‘getvolumeUD’) from the Adehabitat package in R software, which provides the probability density of gorilla occurrence at each coordinate (x, y) of the study area. Analyses using GLMs suggest that gorilla densities (a proxy of previous gorilla utilization intensity) did neither affect the total biomass of key food plant species nor the biomass of each key food plant species (p>0.05). These results may indicate that current revisit rates of feeding sites by gorillas allow for complete plant regeneration, and no signs of overharvesting. Alternatively, feeding sites characterized by very high biomass may be preferred by gorillas and remain sites with the highest biomass even after being frequently used by gorillas. Findings also suggest that carrying capacity of the gorilla population in the study areas may not yet be reached if food is the driving constraint. However, monitoring of the relationship between gorilla densities and food plant biomass must continue while the Virunga population continues growing. Future studies also need to incorporate other sympatric large herbivores in the Virungas who share food plants with mountain gorillas.Keywords: habitat use, gorilla density, plants biomass

Status, diversity and trends of the bird communities in Volcanoes National Park and surrounds, Rwanda

SummaryThe Volcanoes National Park (VNP) in Rwanda, part of the Virunga massif in the Albertine Rift region, boasts some of Rwanda’s best remaining natural vegetation and is home to many endangered and endemic species. The park has suffered from high levels of degradation and destruction, reducing in size by 50% during the 1960s and 1970s, and remains under threat from illegal activities, human population pressure and climate change. This study is the first to investigate the status and trends of bird communities in the VNP, using a multi-year dataset. We use a five-year dataset, totalling over 3,200 point-counts, both within and around the VNP, to assess the conservation value of the VNP for birds in comparison with other national parks and non-protected areas. We assess bird communities and population trends within and around the parks and identify important habitat factors for birds within the VNP. We found that the VNP hosts a unique bird community compared to other localities, with several Albertine Rift endemics and threatened species occurring in the VNP.Hagenia/ Hypericumwoodland, herbaceous vegetation, brush ridge and mixed forest host the highest levels of bird diversity in the VNP, whilst the park’s waterbodies provide key habitat for the endangered Albertine Rift endemic Grauer’s Swamp-warbler (Bradypterus graueri). Elevation had a negative effect on bird diversity in the VNP, whilst the basal area of dead trees,Hageniaand vines had a positive effect. Both inside and outside the VNP, there was a significant decline in abundance and species richness over the sampling years; however, we advocate for further monitoring to confirm these trends. Based on our findings, we recommend effective, targeted management of key habitats for birds within the park, including those identified in this study, in order to mitigate bird declines and conserve the unique bird communities in the VNP.

Morbidity and mortality in infant mountain gorillas (Gorilla beringei beringei): A 46‐year retrospective review

Long-term studies of morbidity and mortality in free-ranging primates are scarce, but may have important implications for the conservation of extant populations. Infants comprise a particularly important age group, as variation in survival rates may have a strong influence on population dynamics. Since 1968, the Mountain Gorilla Veterinary Project (MGVP, Inc.) and government partners have conducted a comprehensive health monitoring and disease investigation program on mountain gorillas (Gorilla beringei beringei) in Rwanda, Uganda, and the Democratic Republic of the Congo. In an effort to better understand diseases in this species, we reviewed reliable field reports (n = 37), gross post-mortem (n = 66), and histopathology (n = 53) reports for 103 infants (less than 3.5 years) mountain gorillas in the Virunga Massif. Our aim was to conduct the first comprehensive analysis of causes of infant mortality and to correlate histological evidence with antemortem morbidity in infant mountain gorillas. Causes of morbidity and mortality were described, and compared by age, sex, and over time. Trauma was the most common cause of death in infants (56%), followed by respiratory infections and aspiration (13%). Gastrointestinal parasitism (33%), atypical lymphoid hyperplasia (suggestive of infectious disease) (31%), and hepatic capillariasis (25%) were the most significant causes of antemortem morbidity identified post-mortem. Identifying the causes of mortality and morbidity in infants of this critically endangered species will help to inform policy aimed at their protection and guide ante- and post-mortem health monitoring and clinical decision-making in the future.

Causes, mechanisms, and consequences of contest competition among female mountain gorillas in Rwanda

Socioecological models predict that contest competition will arise when high quality foods can be usurped or monopolized, leading to more favorable energy balances and higher reproductive success for high-ranking females. Gorillas are interesting species for studying such predictions due to the variety of ecological conditions that they experience in different locations. Using data from 23 female mountain gorillas in 3 social groups in the Virunga Massif, we examined food characteristics that may influence contest competition (food site residence times [FSRT]), proximate mechanisms of such competition (aggression and avoidance), and potential consequences of competition (rank-related differences in energy intake rates, travel expenditures, and activity budgets). The average FSRT of each female was significantly correlated with dominance rank, which suggests that high-ranking females may have greater access to foods that are easier to usurp (as predicted with contest competition). High-ranking females were significantly more aggressive than low-ranking females, and both aggression and avoidance were significantly higher while feeding than during other activities. Contrary to predictions for contest competition, however, rank was not significantly correlated with energy intake rates nor with the proportion of time spent traveling versus feeding. Thus, we did not find any energetic benefits to explain why high-ranking females had significantly higher reproductive success in earlier studies. We propose several alternative explanations and discuss the potential complications of assessing contest competition in species with weak dominance relationships.

Energetic responses to variation in food availability in the two mountain gorilla populations (Gorilla beringei beringei).

Here, we compare food availability and relate this to differences in energy intake rates, time spent feeding, and daily travel distance of gorillas in the two populations. Comparative intraspecific studies investigating spatiotemporal variation in food availability can help us understand the complex relationships between ecology, behavior, and life history in primates and are relevant to understanding hominin evolution. Differences in several variables have been documented between the two mountain gorilla populations in the Virunga Massif and Bwindi Impenetrable National Park, but few direct comparisons that link ecological conditions to feeding behavior have been made. Using similar data collection protocols we conducted vegetation sampling and nutritional analysis on important foods to estimate food availability. Detailed observations of feeding behavior were used to compute energy intake rates and daily travel distance was estimated through GPS readings. Food availability was overall lower and had greater temporal variability in Bwindi than in the Virungas. Energy intake rates and time spent feeding were similar in both populations, but energy intake rates were significantly higher in Bwindi during the period of high fruit consumption. Daily travel distances were significantly shorter in the Virungas. Overall, despite the differences in food availability, we did not find large differences in the energetics of gorillas in the two populations, although further work is needed to more precisely quantify energy expenditure and energy balance. These results emphasize that even species with high food availability can exhibit behavioral and energetic responses to variable ecological conditions, which are likely to affect growth, reproduction, and survival.

Challenges in the use of genetic mark-recapture to estimate the population size of Bwindi mountain gorillas (Gorilla beringei beringei)

Monitoring the population dynamics of endangered species is a critical component of conservation management strategies, but attaining accurate and precise estimates of population sizes using cost and time effective methods can be challenging. Routine censuses of the two populations of critically endangered mountain gorillas (Gorilla beringei beringei) have been conducted over the last decades to monitor populations and evaluate the effectiveness of conservation strategies. A census in 2006 of the mountain gorillas in Bwindi Impenetrable National Park, Uganda, showed the value of genetic analysis of fecal samples collected at nest sites by revealing discrepancies between the numbers of nests and uniquely identified gorillas. In this study, we censused the Bwindi gorilla population using a ‘mark-recapture’ method which involved genetic analysis of fecal samples collected in 2011 during two ‘sweep’ surveys of the entire park. We found that a notable proportion of gorillas were missed in either of the two sweeps (minimum 35% and 31%, respectively). Based on the number of genotyped gorillas and correction factors, we estimated the population to contain a minimum of 400 individuals. Using the mark-recapture approach, we infer possibly as many as 430 gorillas (95% confidence interval: 398–487). As the 2010 census of the Virunga Massif population found 480 gorillas, the total number of mountain gorillas worldwide is at least 880 individuals. Simulations using different mark-recapture models suggest that a future census of Bwindi mountain gorillas would benefit by increasing the number of sweeps in order to achieve accurate and precise results. Finally, based on our results, we recommend a sequential approach incorporating a pilot study and simulations for optimizing time and resources in large mammal genetic census studies.

Fine-scale genetic structure analyses suggest further male than female dispersal in mountain gorillas.

BackgroundMolecular studies in social mammals rarely compare the inferences gained from genetic analyses with field information, especially in the context of dispersal. In this study, we used genetic data to elucidate sex-specific dispersal dynamics in the Virunga Massif mountain gorilla population (Gorilla beringei beringei), a primate species characterized by routine male and female dispersal from stable mixed-sex social groups. Specifically, we conducted spatial genetic structure analyses for each sex and linked our genetically-based observations with some key demographic and behavioural data from this population.ResultsTo investigate the spatial genetic structure of mountain gorillas, we analysed the genotypes of 193 mature individuals at 11 microsatellite loci by means of isolation-by-distance and spatial autocorrelation analyses. Although not all males and females disperse, female gorillas displayed an isolation-by-distance pattern among groups and a signal of dispersal at short distances from their natal group based on spatial autocorrelation analyses. In contrast, male genotypes were not correlated with spatial distance, thus suggesting a larger mean dispersal distance for males as compared to females. Both within sex and mixed-sex pairs were on average genetically more related within groups than among groups.ConclusionsOur study provides evidence for an intersexual difference in dispersal distance in the mountain gorilla. Overall, it stresses the importance of investigating spatial genetic structure patterns on a sex-specific basis to better understand the dispersal dynamics of the species under investigation. It is currently poorly understood why some male and female gorillas disperse while others remain in the natal group. Our results on average relatedness within and across groups confirm that groups often contain close relatives. While inbreeding avoidance may play a role in driving female dispersal, we note that more detailed dyadic genetic analyses are needed to shed light on the role of inbreeding avoidance as an ultimate cause of female dispersal in mountain gorillas.