Virunga Gorillas Research Articles

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.

Variability of weaning age in mountain gorillas (Gorilla beringei beringei).

Weaning is a key life history milestone for mammals that represents both the end of nutritional investment from the perspective of mothers and the start of complete nutritional independence for the infants. The age at weaning may vary depending on ecological, social, and demographic factors experienced by the mother and infant. Bwindi mountain gorillas live in different environmental conditions and have longer interbirth intervals than their counterparts in the Virunga Volcanoes, yet other life history characteristics of this population remain less well known. We use long-term data from Bwindi Impenetrable National Park, Uganda to examine factors related to weaning age. We analyzed data on infants born in four mountain gorilla groups in Bwindi to quantify their age of weaning (defined as last nipple contact) and to test if the sex of offspring, parity, and dominance rank of mother influences age of weaning. We also compared the age at weaning and time to conception after resumption of mating in Bwindi and Virunga gorillas. Bwindi gorillas were weaned at an average age of 57.5 months. No difference was found between age of weaning for primiparous and multiparous mothers, nor did maternal dominance rank influence age of weaning, but sons were weaned at a later age than daughters. The majority of Bwindi mothers were still suckling when they resumed mating and mothers generally conceived before they weaned their previous offspring. The age of weaning was significantly later in Bwindi than in Virunga gorillas. After mothers resumed mating, the time to conceiving the next offspring was not significantly longer for Bwindi females than Virungas females (6 vs. 4 months). Later weaning age for sons than daughters is similar to findings of other studies of great apes. Bwindi mountain gorillas are weaned at approximately the same age as western gorillas and chimpanzees, which is more than a year later than Virunga mountain gorillas. The results of this study suggest that variation in ecological conditions of populations living in close geographic proximity can result in variation in life history patterns, which has implications for understanding the evolution of the unique life history patterns of humans.

A Simple and Environmentally Friendly Field Method for Fecal Analysis of Herbivore Diet

ABSTRACTWe developed a simple and environmentally‐friendly protocol for identifying herbivore diets by isolating and identifying microscopic vegetation fragments from fecal materials that can be easily applied in remote areas where laboratories are absent. We used feces from human‐habituated Virunga mountain gorillas (Gorilla beringei beringei), whose diet is well‐known, to develop and validate the protocol. First, fresh materials from parts of the 5 most frequently consumed plants by the Virunga gorillas were collected from June to July, 2016, processed, and their distinctive features photographed using a digital microscope to establish a plant reference collection. Second, we collected fresh fecal samples from 16 known gorillas and successfully identified distinctive vegetation fragments of the 5 key food plants. The method is inexpensive, environmentally friendly, and does not require sophisticated laboratory equipment. It has the potential to be applied to species that cannot be easily studied by direct observation and those ranging in remote regions. The method can also be used in other studies involving plant‐animal interactions, the ontogeny of feeding behavior, and animal ecology. © 2020 The Wildlife Society.

Human Respiratory Syncytial Virus Detected in Mountain Gorilla Respiratory Outbreaks

Respiratory illness (RI) accounts for a large proportion of mortalities in mountain gorillas (Gorilla beringei beringei), and fatal outbreaks, including disease caused by human metapneumovirus (HMPV) infections, have heightened concern about the risk of human pathogen transmission to this endangered species, which is not only critically important to the biodiversity of its ecosystem but also to the economies of the surrounding human communities. Our goal was to conduct a molecular epidemiologic study to detect the presence of HRSV and HMPV in fecal samples from wild human-habituated free-ranging mountain gorillas in Rwanda and to evaluate the role of these viruses in RI outbreaks. Fecal samples were collected from gorillas with clinical signs of RI between June 2012 and February 2013 and tested by real-time and conventional polymerase chain reaction (PCR) assays; comparison fecal samples were obtained from gorillas without clinical signs of RI sampled during the 2010 Virunga gorilla population census. PCR assays detected HMPV and HRSV first in spiked samples; subsequently, HRSV-A, the worldwide-circulating ON1 genotype, was detected in 12 of 20 mountain gorilla fecal samples collected from gorillas with RI during outbreaks, but not in samples from animals without respiratory illness. Our findings confirmed that pathogenic human respiratory viruses are transmitted to gorillas and that they are repeatedly introduced into mountain gorilla populations from people, attesting to the need for stringent biosecurity measures for the protection of gorilla health.

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.

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

Social and ecological factors alter stress physiology of Virunga mountain gorillas (Gorilla beringei beringei).

Living in a rapidly changing environment can alter stress physiology at the population level, with negative impacts on health, reproductive rates, and mortality that may ultimately result in species decline. Small, isolated animal populations where genetic diversity is low are at particular risks, such as endangered Virunga mountain gorillas (Gorilla beringei beringei). Along with climate change‐associated environmental shifts that are affecting the entire population, subpopulations of the Virunga gorillas have recently experienced extreme changes in their social environment. As the growing population moves closer to the forest's carrying capacity, the gorillas are coping with rising population density, increased frequencies of interactions between social units, and changing habitat use (e.g., more overlapping home ranges and routine ranging at higher elevations). Using noninvasive monitoring of fecal glucocorticoid metabolites (FGM) on 115 habituated Virunga gorillas, we investigated how social and ecological variation are related to baseline FGM levels, to better understand the adaptive capacity of mountain gorillas and monitor potential physiological indicators of population decline risks. Generalized linear mixed models revealed elevated mean monthly baseline FGM levels in months with higher rainfall and higher mean maximum and minimum temperature, suggesting that Virunga gorillas might be sensitive to predicted warming and rainfall trends involving longer, warmer dry seasons and more concentrated and extreme rainfall occurrences. Exclusive use of smaller home range areas was linked to elevated baseline FGM levels, which may reflect reduced feeding efficiency and increased travel efforts to actively avoid neighboring groups. The potential for additive effects of stress‐inducing factors could have short‐ and long‐term impacts on the reproduction, health, and ultimately survival of the Virunga gorilla population. The ongoing effects of environmental changes and population dynamics must be closely monitored and used to develop effective long‐term conservation strategies that can help address these risk factors.

Are the gorillas in Bwindi Impenetrable National Park “true” mountain gorillas?

The gorillas that inhabit Bwindi Impenetrable National Park in Uganda are the least known of the eastern gorillas. Because they are an allopatric population living a minimum of 25 km from the well-studied population of mountain gorillas (Gorilla beringei beringei) in Rwanda and have certain morphological and ecological differences from these gorillas, their taxonomic status has been in question in recent years. This study presents new craniodental metrics from Bwindi individuals and compares them to Virunga individuals as well as to eastern lowland gorillas, G. gorilla graueri. Multivariate statistics, including MANCOVA, least-squares, regression, and principal components analyses, were used to evaluate how closely the Bwindi crania resemble the Virunga crania and how both relate to G. g. graueri. Results indicate that the Bwindi gorillas have generally smaller crania than the Virunga gorillas, but when metrics are log-transformed, the only variable that distinguishes the Bwindi individuals is a longer face. When both populations are compared to G. g. graueri, they cluster together separately from the eastern lowland gorillas, sharing such features as higher rami, wider bigonia, longer mandibles, and wider and shorter mandibular symphyses in relation to G. g. graueri. Functional morphological explanations for these differences are discussed, but lacking measurements of the physical properties of G. g. graueri, they cannot fully be explained. Results clearly indicate that at least pertaining to the cranium, upon which most gorilla taxonomy is based, the Bwindi gorillas are proper mountain gorillas (G. b. beringei).

Population dynamics of the Bwindi mountain gorillas

Mountain gorillas are critically endangered, with just a few hundred animals remaining in each of two populations: in Bwindi Impenetrable National Park in south-western Uganda and the nearby Virunga Volcanoes on the borders of Uganda, Rwanda and the Democratic Republic of Congo. While the life-history and population dynamics of the Virunga gorillas have been studied extensively, comparable information from Bwindi has not been reported. Such studies are difficult to conduct because gorillas are long-lived, have delayed reproduction, and monitoring known individuals requires habituation of social groups. Bwindi has experienced lower levels of human disturbance than the Virungas, yet its gorilla population has shown little or no growth over the past 20 years, while a subpopulation of study groups in the Virungas have grown by 3–4% per year. Here we show that the lower growth rate at Bwindi may arise mainly from lower fertility than the Virunga study groups, rather than higher mortality. This difference may indicate that the more frugivorous Bwindi gorillas have a slower life-history, or that they are closer to the carrying capacity of their habitat. The study groups at Bwindi had a higher growth rate than the broader population, possibly because they receive veterinary care and better protection from poachers, but further analysis is necessary to understand the complex interactions among human disturbance, ecology, and the gorillas’ population dynamics. Meanwhile, efforts to increase the Bwindi population should place emphasis on reducing human disturbances, improving our understanding of the impact of habitat quality, and ensuring that the gorillas can expand their home ranges into under-utilized areas of the park.

Nutritional composition of the diet of the gorilla (Gorilla beringei): a comparison between two montane habitats

The mountain gorilla (Gorilla beringei beringei) lives in two geographically separated populations, Bwindi Impenetrable National Park, Uganda and in three national parks spanning the Virunga mountain region in Rwanda, Democratic Republic of Congo, and Uganda. The altitude, climate and plant composition of these habitats differ. Our goal was to compare the diets of gorillas living in each of these habitats. The nutrients in staple foods and in the diets of individuals in a group of gorillas in Bwindi (N = 12 individuals) and a group in the Virungas (N = 7 individuals) were compared to determine if differences in dietary composition affected concentrations of nutrients in their diets. At both sites gorilla diets consisted primarily of herbaceous leaves, but the diet of Bwindi gorillas contained more tree leaves, fruit, pith and dry wood, and fewer stems. Despite differences in habitat and dietary composition, the nutrient concentrations in both gorilla diets were remarkably similar. On a dry matter basis, the diets and staple foods of Bwindi and Virunga gorillas contained similar concentrations of crude protein (CP), fibre (NDF) and non-structural carbohydrates (TNC). Bwindi gorillas ate diets containing 18% CP, 43% NDF and 19% TNC on a dry-matter basis, while the diets of the Virunga gorillas contained 17% CP, 41% NDF and 18% TNC. Our results demonstrate that gorillas consume diets that differ by plant species and part, but contain similar concentrations of nutrients. This suggests that classifying animals by broad dietary strategy (e.g. frugivory and folivory) does not provide a reliable indicator of the nutritional quality of their diet, and that our previous assumptions about these categories should be re-evaluated.

Gorillas of Bwindi-Impenetrable Forest and the Virunga Volcanoes: Taxonomic implications of morphological and ecological differences.

Based on their geographic proximity to the Virunga Volcanoes (≈ 25 km), the Bwindi-Impenetrable Forest gorillas have been referred to the subspecies Gorilla gorilla beringei. Differences in anatomy, habitat, ecology, and behavior, however, suggest Bwindi gorillas are distinct from those in the Virungas. Relative to Virunga gorillas, Bwindi gorillas live at lower elevations, in warmer temperatures, are much more arboreal, have longer day ranges and larger home ranges, and eat much more fruit and pith, and less bamboo and leaves. Morphological differences reflect the differences in ecology, habitat, and behavior. Bwindi gorillas measured have smaller bodies, relatively longer limbs, hands, and feet, shorter trunks, thumbs, big toes, and tooth row lengths, and narrower trunks and orbital breadths than Virunga gorillas. These differences indicate Bwindi gorillas do not belong to G.g. beringei and should not be referred to as "mountain gorillas." How unique the distinguishing features of Bwindi gorillas are, and whether or not they should be assigned to a new taxon, depends on the expression of these features in eastern lowland gorillas (G.g. graueri). © 1996 Wiley-Liss, Inc.

Population Viability Estimates: Theory and Practice for a Wild Gorilla Population

The logic of demographic modeling, the apparent simplicity of its quantifiably substantiated answers, and the ready availability of software correlate with increasing use of demographic modeling as the means of applying biology to the conservation of potentially endangered populations. I investigated that use by considering a small population (about 300 individuals) of a large, forest‐dwelling mammal of the tropics, the Virunga gorilla (Gorilla gorilla) of Zaire, Uganda, and Rwanda. Because censuses of forest populations are so inaccurate and data on variance of some parameters takes so long to collect, models might not be broadly applicable. Therefore, simple demographic indices of potential extinction should replace sophisticated models. The current best index could be problematic, however, because it is based on detecting adult mortality, perhaps the most difficult demographic parameter to measure. Models of the Virunga gorilla population that incorporate aspects of demographic heterogeneity valuably indicate genetic and demographic persistence for several hundred years. Deterministic change in habitat is a greater threat than stochastic demographic variation, and yet our ecological ignorance is such that we could not begin to model the consequences of removal of even the main food plant. We must add to our ability to model outcomes of demographic perturbation a far greater understanding of the processes by which the perturbations occur. Demography allows us to model demographic response to demographic change, but we usually need ecology to tell us how the threat produced the demographic change in the first place. In a time of change, accurate prediction requires ecological understanding of process as well as demographic understanding of outcome.

Intra- and inter-group interactions of an all-male group of virunga mountain gorillas (Gorilla gorilla beringei)

Six unrelated male gorillas formed an all-male group within the Virunga mountain gorilla population. Frequent homosexual interactions characterized the high cohesiveness of this group. Such homosexual behavior reduced the inter-individual distances and increased the social tension between the two silverbacks in the group. The silverbacks retained “ownership” of the homosexual partners, but competed and fought with each other violently when the partners avoided or ignored their courtship. Neither submissive nor reassurance behavior was noted between the silverbacks. Thus, their relationships may not be explained in terms of dominance and subordinancy. However, the loser-support and mediating behavior observed in the group prevented them from engaging in severe fights. Aggression was always directed from the elder and dominant males to the younger and subordinate males, while supporting interactions occurred in the opposite direction. The blackbacks frequently supported the subadult aggressees by attacking the silverbacks, and the younger males displayed mediating behavior in violent fights between the silverbacks. The group's ranging was influenced by encounters with neighboring social units. The members avoided contact with other units and shifted their range after several encounters. On the other hand, when and after a subadult male had immigrated into their group, they frequently encountered other units and did not move away from the encounter site. An all-male group may not be a favorable unit for females to transfer to, but may be profitable for maturing males to associate with. Its formation is probably related to recent social change in the Virunga gorilla population.

A census of the Virunga gorillas

Mountain gorillas exist in the wild today only in the Virunga volcanoes, which span the borders of Zaire, Rwanda and Uganda, and in the Bwindi Forest Reserve of Uganda, and number probably fewer than 400. The authors describe the 1981 survey of the entire Virunga gorilla population, the first to be done since 1973. The results clearly show how the gorillas in the Rwanda. section of the volcanoes have benefited from the protection afforded by the activities of the Karisoke Research Centre and the Mountain Gorilla Project in Rwanda. There is also optimism that the Zaire population, which has retained the potential for increase, could recover if given better protection from poaching and disturbance.

Population dynamics of the virunga gorillas: 1959–1978

The Virunga gorilla population declined from 400–500 individuals in 1959–1960 to 260–290 in 1971–1973. Most of this decline is attributed to heavy hunting pressure in the Zairian sector. Large-scale habitat destruction and cattle disturbance in the Rwandan and Ugandan sectors are held responsible for further gorilla losses. Changes are documented in group size and composition, as well as in total numbers. Between 1971–1973 and 1976–1978, the population stabilised in the range of 252–285 gorillas. A tendency toward larger group sizes was observed in certain sectors, and the overall percentage of infants increased. Comparison of the 1976–1978 group compositions with a simulated stable age structure model indicates a more healthy population than in 1971–1973. Potential for population growth is shown to be greater than previously predicted. Failure to increase in the inter-census period is attributed to continued poaching pressure. Current conservation efforts are described.