Examining the dual hormone hypothesis in wild male mountain gorillas (Gorilla beringei beringei).

The Challenge Hypothesis is an influential framework for understanding how androgens are involved in the promotion of competitive behavior during mating-related challenges and has been tested extensively in studies across scientific disciplines. Mixed support in psychological research led scholars to develop the Dual Hormone Hypothesis as a potential path forward, which argues that glucocorticoids moderate the relationship between androgens and status-striving. In the current study, we examine the Challenge Hypothesis and the Dual Hormone Hypothesis in wild male mountain gorillas, representing the first time the latter hypothesis has been tested in a non-human primate. In a sample of 30 adult males comprising over 600 days of observation, we find some limited support for the Challenge Hypothesis. Greater daily rates of targeted aggression toward other adult males corresponded to higher fecal androgen metabolites 1-2 days following observations, though this pattern did not fully generalize to dominance rank or other competitive behaviors examined. However, we find no support for the Dual Hormone Hypothesis: neither dominance rank nor any category of competitive behavior was predicted by the interaction between androgens and glucocorticoids. We close by discussing how this initial investigation might be leveraged toward the development of an expanded Dual Hormone Hypothesis that draws on the large evidence base in primate behavioral ecology.

Anoplocephalid tapeworms in mountain gorillas (Gorilla beringei beringei) inhabiting the Volcanoes National Park, Rwanda.

Cestodes of the family Anoplocephalidae parasitize a wide range of usually herbivorous hosts including e.g. rodents, ungulates, primates, elephants and hyraxes. While in some hosts, the epidemiology of the infection is well studied, information is lacking in others. In this study of mountain gorillas in the Virunga Massif, an extensive sample set comprising adult cestodes collected via necropsies, proglottids shed in feces, and finally, fecal samples from both night nests and identified individuals were analysed. Anoplocephala gorillae was the dominant cestode species detected in night nest samples and individually known gorillas, of which only 1 individual hosted a Bertiella sp. It was shown that the 2 species can be distinguished through microscopy based on egg morphology and polymerase chain reaction (PCR) assays for diagnostics of both species were provided. Sequences of mitochondrial (cox 1) and nuclear (ITS1, 18S rDNA, 28S rDNA) markers were used to evaluate the phylogenetic position of the 2 cestodes detected in mountain gorillas. Both types of fecal samples, from night nests and from identified individuals, provided comparable information about the prevalence of anoplocephalid cestodes, although the analysis of samples collected from identified gorilla individuals showed significant intra-individual fluctuation of A. gorillae egg shedding within a short period. Therefore, multiple samples should be examined to obtain reliable data for wildlife health management programmes, especially when application of anthelmintic treatment is considered. However, while A. gorillae is apparently a common symbiont of mountain gorillas, it does not seem to impair the health of its host.

Multiple mechanisms for inbreeding avoidance used simultaneously in a wild ape.

Mating with close kin can have considerable negative fitness consequences, which are expected to result in selective pressure for inbreeding avoidance mechanisms, such as dispersal, mate choice and post-copulatory biases. Captive studies have suggested that inbreeding avoidance through mate choice is far less widespread than expected and may be absent where other mechanisms already limit inbreeding. However, few studies have examined multiple mechanisms of inbreeding avoidance simultaneously, particularly in the wild. We use 13 years of detailed dispersal, copulation and paternity data from mountain gorillas to examine inbreeding avoidance. We find that partial dispersal of both sexes results in high kinship in multimale groups, but that copulations between close kin occur 40% less than expected. We find strong kin discrimination in mate choice, with significant avoidance of maternal kin but more limited avoidance of paternal kin. We find no evidence for post-copulatory inbreeding avoidance. Our analyses support familiarity-based mechanisms of kin identification and age-based avoidance that limits mating between fathers and daughters in their natal group. Our findings demonstrate that multiple complementary mechanisms for inbreeding avoidance can evolve in a single species and suggest that inbreeding avoidance through mate choice may enable more flexible dispersal systems to evolve.

An empirical comparison of several commercial enzyme immunoassays for the non-invasive assessment of adrenocortical and gonadal function in mountain gorillas.

Wildlife researchers seeking to non-invasively examine endocrine function in their study species are presented with a dense and technical 'garden of forking paths' to navigate between collecting a biological sample and obtaining a final measurement. In particular, the choice of which enzyme immunoassay (EIA) to use with collected fecal samples, out of the many options offered by different manufacturers and research laboratories, may be one of the most consequential for final results. However, guidance for making this decision is still emerging. With this gap in mind, we performed a head-to-head comparison of results obtained from four different EIAs for fecal glucocorticoid metabolites (FGCMs), and three different EIAs for fecal androgen metabolites (FAMs), applied to the same set of fecal samples collected from the mountain gorillas (Gorilla beringei beringei) monitored by the Dian Fossey Gorilla Fund in Volcanoes National Park, Rwanda. We provide a) an analytical validation of the different EIAs via tests of parallelism and linearity; b) an estimate of inter-assay correlation between EIA kits designed for the same metabolites; and c) a test of the kits' ecological validity, in which we examine how well each captures endocrine changes following events that theory predicts should result in elevated FGCM and/or FAM concentrations. Our results show that kits differ to some degree in their performance; at the same time, nearly all assays exhibited at least moderate evidence of validity and covariance with others for the same analyte. Our findings, which differ somewhat from similar comparisons performed in other species, demonstrate the need to directly assess assay performance in a species- and context-specific manner as part of efforts to develop the burgeoning discipline of wildlife endocrinology.

Discovery of Novel Herpes Simplexviruses in Wild Gorillas, Bonobos, and Chimpanzees Supports Zoonotic Origin of HSV-2.

Viruses closely related to human pathogens can reveal the origins of human infectious diseases. Human herpes simplexvirus type 1 (HSV-1) and type 2 (HSV-2) are hypothesized to have arisen via host-virus codivergence and cross-species transmission. We report the discovery of novel herpes simplexviruses during a large-scale screening of fecal samples from wild gorillas, bonobos, and chimpanzees. Phylogenetic analysis indicates that, contrary to expectation, simplexviruses from these African apes are all more closely related to HSV-2 than to HSV-1. Molecular clock-based hypothesis testing suggests the divergence between HSV-1 and the African great ape simplexviruses likely represents a codivergence event between humans and gorillas. The simplexviruses infecting African great apes subsequently experienced multiple cross-species transmission events over the past 3 My, the most recent of which occurred between humans and bonobos around 1 Ma. These findings revise our understanding of the origins of human herpes simplexviruses and suggest that HSV-2 is one of the earliest zoonotic pathogens.

Distinguishing mobility and immobility when establishing species-specific activity budgets: A case study with gorillas (Gorilla berengei berengei and Gorilla gorilla gorilla).

Activity budgets characterize the distribution of behavior over a specified time period. In some cases, having comparable data from free-ranging populations can help inform the management of wildlife in zoos and sanctuaries. For example, although variations exist across subspecies, seasons, and study sites, diurnal activity budgets for free-ranging gorillas largely consist of feeding and resting. Unfortunately, most studies do not consistently differentiate between the type of activities gorillas exhibit while locomoting versus stationary. Therefore, it can be difficult to characterize optimal levels of aerobic activity that might enhance body condition or promote gorilla health in zoos and sanctuaries. In this study, we concurrently measured the mobility state and activity of mountain gorillas (Gorilla beringei beringei) in Volcanoes National Park, Rwanda. From June to August 2015, behavioral data were collected using group scan sampling with 15-min intervals in two groups (N = 29 gorillas) monitored by the Dian Fossey Gorilla Fund International. Overall, gorillas spent significantly more time immobile (85.2% of observations) than mobile (14.8%), revealing energy expenditure levels comparable to western lowland gorillas living in zoos. There was no difference in behavioral diversity when gorillas were mobile versus immobile but adult females exhibited substantially less behavioral diversity while immobile than other age-sex classes. There was more diversity in behaviors following the transition from immobile to mobile than vice versa, particularly for adult females. Future studies should concurrently measure mobility state and behavior to improve the precision of activity budget data and serve as a more useful tool for evaluating optimal activity levels for wildlife in human care.

Evidence of demographic buffering in an endangered great ape: Social buffering on immature survival and the role of refined sex-age classes on population growth rate.

Theoretical and empirical research has shown that increased variability in demographic rates often results in a decline in the population growth rate. In order to reduce the adverse effects of increased variability, life-history theory predicts that demographic rates that contribute disproportionately to population growth should be buffered against environmental variation. To date, evidence of demographic buffering is still equivocal and limited to analyses on a reduced number of age classes (e.g. juveniles and adults), and on single sex models. Here we used Bayesian inference models for age-specific survival and fecundity on a long-term dataset of wild mountain gorillas. We used these estimates to parameterize two-sex, age-specific stochastic population projection models that accounted for the yearly covariation between demographic rates. We estimated the sensitivity of the long-run stochastic population growth rate to reductions in survival and fecundity on ages belonging to nine sex-age classes for survival and three age classes for female fecundity. We found a statistically significant negative linear relationship between the sensitivities and variances of demographic rates, with strong demographic buffering on young adult female survival and low buffering on older female and silverback survival and female fecundity. We found moderate buffering on all immature stages and on prime-age females. Previous research on long-lived slow species has found high buffering of prime-age female survival and low buffering on immature survival and fecundity. Our results suggest that the moderate buffering of the immature stages can be partially due to the mountain gorilla social system and the relative stability of their environment. Our results provide clear support for the demographic buffering hypothesis and its predicted effects on species at the slow end of the slow-fast life-history continuum, but with the surprising outcome of moderate social buffering on the survival of immature stages. We also demonstrate how increasing the number of sex-age classes can greatly improve the detection of demographic buffering in wild populations.

Group structure, but not dominance rank, predicts fecal androgen metabolite concentrations of wild male mountain gorillas (Gorilla beringei beringei).

Androgens are important mediators of male-male competition in many primate species. Male gorillas' morphology is consistent with a reproductive strategy that relies heavily on androgen-dependent traits (e.g., extreme size and muscle mass). Despite possessing characteristics typical of species with an exclusively single-male group structure, multimale groups with strong dominance hierarchies are common in mountain gorillas. Theory predicts that androgens should mediate their dominance hierarchies, and potentially vary with the type of group males live in. We validated the use of a testosterone enzyme immunoassay (T-EIA R156/7, CJ Munro, UC-Davis) for use with mountain gorilla fecal material by (1) examining individual-level androgen responses to competitive events, and (2) isolating assay-specific hormone metabolites via high-performance liquid chromatography. Males had large (2.6- and 6.5-fold), temporary increases in fecal androgen metabolite (FAM) after competitive events, and most captured metabolites were testosterone or 5α-dihydrotestosterone-like androgens. We then examined the relationship between males' dominance ranks, group type, and FAM concentrations. Males in single-male groups had higher FAM concentrations than males in multimale groups, and a small pool of samples from solitary males suggested they may have lower FAM than group-living peers. However, data from two different time periods (n = 1610 samples) indicated there was no clear relationship between rank and FAM concentrations, confirming results from the larger of two prior studies that measured urinary androgens. These findings highlight the need for additional research to clarify the surprising lack of a dominance hierarchy/androgen relationship in mountain gorillas.

A partnership to build scientific capacity of Rwanda's future conservationists: The Memoirs Program.

The future of primates depends on conservationists in primate range countries having the education and opportunities needed to facilitate conservation efforts. However, most primates are found in countries where conservation is underfunded. Rwanda is home to 14 primate species, with three being listed as endangered on the International Union for Conservation of Nature (IUCN) Red List. Their remaining native habitats are scarce, highly fragmented, and surrounded by high human densities. Lacking education opportunities and the loss of qualified individuals following decades of political instability created a need for well-trained conservationists to tackle conservation challenges in Rwanda. The Memoirs Program, a science capacity-building partnership between Dian Fossey Gorilla Fund (DFGF), University of Rwanda (UR), the Rwanda Development Board, and Cleveland Metroparks Zoo, aims to train the next generation of Rwandan conservationists. Annually, the Memoirs Program provides 10-12 top students from the UR Conservation Biology undergraduate program the opportunity to conduct senior research projects (Memoirs) at DFGF addressing conservation issues in and around Rwanda's protected areas. Students receive one-on-one supervision by DFGF scientists and affiliates to supplement UR mentorship, along with targeted teaching modules that reinforce and complement UR curricula. In continuation of the Memoirs Program, participants are offered professional development, employment opportunities, and support to publish their research in peer-reviewed journals. Since inception of the Program in 2003, 123 students have participated with 83% being employed or pursuing higher education degrees in conservation-related fields in Rwanda and the region. Pre- and post-training comparison of oral presentation skills, one of the teaching modules, involving 22 students demonstrated increased post-training performance. The Memoirs Program represents an effective partnership between the government, an international conservation non-governmental organization, and a public higher-learning institution in a developing country to support high-performing students to become tomorrow's conservation leaders. This model could be replicated within Rwanda and in other regions of the world.

Comparing measures of social complexity: larger mountain gorilla groups do not have a greater diversity of relationships.

Social complexity reflects the intricate patterns of social interactions in societies. Understanding social complexity is fundamental for studying the evolution of diverse social systems and the cognitive innovations used to cope with the demands of social life. Social complexity has been predominantly quantified by social unit size, but newer measures of social complexity reflect the diversity of relationships. However, the association between these two sets of measures remains unclear. We used 12 years of data on 13 gorilla groups to investigate how measures of social complexity relate to each other. We found that group size was a poor proxy for relationship diversity and that the social complexity individuals experienced within the same group varied greatly. Our findings demonstrate two fundamental takeaways: first, that the number of relationships and the diversity of those relationships represent separate components of social complexity, both of which should be accounted for; and second, that social complexity measured at the group level may not represent the social complexity experienced by individuals in those groups. These findings suggest that comprehensive studies of social complexity, particularly those relating to the social demands faced by individuals, may require fine-scale social data to allow accurate comparisons across populations and species.

Inter-group relationships influence territorial defence in mountain gorillas.

Many species show territoriality, in which territory owners have exclusive or priority use of a region. In humans, tolerance of others within our space also depends greatly on our social relationships with them. This has been hypothesized as one potential driver of the evolution of long-term, inter-group relationships, through enabling shared access of resources and easing disputes over space. However, extremely little is known about the importance of social relationships between neighbouring groups in non-humans for how space is used and shared. Using 16 years of data on the simultaneous movement and interaction patterns of 17 mountain gorilla groups, we investigated how the occurrence of aggressive and affiliative behaviour during inter-group encounters was influenced by both their social and spatial context. We found evidence of territorial defence, with rates of aggression increasing towards the centre of home ranges. Groups which had previously split from each other showed higher levels of affiliation during encounters with each other and experienced lower levels of aggression when within the other's peripheral home range. However, encounters within core areas of the home range consistently elicited higher aggression, regardless of the groups' history. Our findings indicate that not only are the social relationships between groups retained after they split from one another but also that these relationships enable groups to access certain areas with a reduced risk of aggression. This suggests that reduced aggression when accessing areas within neighbours' home ranges may be an advantage for the maintenance of inter-group relationships and a potential driver in the evolution of long-term, post-dispersal relationships and complex multi-level societies.

Urticaceae leaves with stinging trichomes were already present in latest early Eocene Okanogan Highlands, British Columbia, Canada.

PREMISE: Paleontologists use tooth form to assess diets of fossil mammals. Plants would also be expected to adapt their morphology to respond to herbivory. Fossil nettle leaves with definitive stinging trichomes (tribe Urticeae, family Urticaceae) are described from the early Eocene upland lacustrine floras of the Okanogan Highlands, British Columbia, Canada. This is the first report of stinging trichomes in the fossil record. Their occurrence in western North America at a time of major large herbivorous mammal radiation suggests they acted, as they do today, as a deterrent for mammal herbivory. METHODS: Fossil leaf compressions and extant leaves were photographed with standard methods. Focus-shift stacking was used to layer photos of the fossil leaves. RESULTS: Urticaceous fossil leaves from the Okanogan Highlands greatly resemble their modern relatives in leaf morphology and particularly in both stinging and nonstinging trichomes. Nettles are common components of the flora of the Volcanoes National Park in Rwanda. This region is used as a modern analogue for the Okanogan Highlands, based on comparable elevation, equable conditions that host both similar floras and large folivores. CONCLUSIONS: Nettles in tribe Urticeae (Urticaceae) producing leaves with stinging and nonstinging trichomes were already present in the early Eocene of western North America at a pivotal time during the early radiation of modern mammalian herbivore groups. They offer tantalizing evidence of a selective response that plants may have developed to protect themselves from the evolving mammalian herbivores of that time.

Assessing Host-Virus Codivergence for Close Relatives of Merkel Cell Polyomavirus Infecting African Great Apes.

UNLABELLED: 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.

Body growth and life history in wild mountain gorillas (Gorilla beringei beringei) from Volcanoes National Park, Rwanda.

OBJECTIVES: Great apes show considerable diversity in socioecology and life history, but knowledge of their physical growth in natural settings is scarce. We characterized linear body size growth in wild mountain gorillas from Volcanoes National Park, Rwanda, a population distinguished by its extreme folivory and accelerated life histories. METHODS: In 131 individuals (0.09-35.26 years), we used non-invasive parallel laser photogrammetry to measure body length, back width, arm length and two head dimensions. Nonparametric LOESS regression was used to characterize cross-sectional distance and velocity growth curves for males and females, and consider links with key life history milestones. RESULTS: Sex differences became evident between 8.5 and 10.0 years of age. Thereafter, female growth velocities declined, while males showed increased growth velocities until 10.0-14.5 years across dimensions. Body dimensions varied in growth; females and males reached 98% of maximum body length at 11.7 and 13.1 years, respectively. Females attained 95.3% of maximum body length by mean age at first birth. Neonates were 31% of maternal size, and doubled in size by mean weaning age. Males reached maximum body and arm length and back width before emigration, but experienced continued growth in head dimensions. CONCLUSIONS: While comparable data are scarce, our findings provide preliminary support for the prediction that mountain gorillas reach maximum body size at earlier ages compared to more frugivorous western gorillas. Data from other wild populations are needed to better understand comparative great ape development, and investigate links between trajectories of physical, behavioral, and reproductive maturation.

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

OBJECTIVES: 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. MATERIAL AND METHODS: 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. RESULTS: 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. DISCUSSION: 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.

Cumulative early-life adversity does not predict reduced adult longevity in wild gorillas.

Extensive research across fields has repeatedly confirmed that early-life adversity (ELA) is a major selective force for many taxa, in part via its ties to adult health and longevity.1,2,3 Negative effects of ELA on adult outcomes have been documented in a wide range of species, from fish to birds to humans.4 We used 55 years of long-term data collected on 253 wild mountain gorillas to examine the effects of six putative sources of ELA on survival, both individually and cumulatively. Although cumulative ELA was associated with high mortality in early life, we found no evidence that it had detrimental consequences for survival later in life. Experiencing three or more forms of ELA was associated with greater longevity, with a 70% reduction in the risk of death across adulthood, driven specifically by greater longevity in males. Although this higher survival in later life is likely a consequence of sex-specific viability selection5 during early life due to the immediate mortality consequences of adverse experiences, patterns in our data also suggest that gorillas have significant resilience to ELA. Our findings demonstrate that the detrimental consequences of ELA on later life survival are not universal, and indeed largely absent in one of humans' closest living relatives. This raises important questions about the biological roots of sensitivity to early experiences and the protective mechanisms that contribute to resiliency in gorillas, which could be critical for understanding how best to encourage similar resiliency to early-life shocks in humans.

African Queens find mates when males are rare.

In butterflies and moths, male-killing endosymbionts are transmitted from infected females via their eggs, and the male progeny then perish. This means that successful transmission of the parasite relies on the successful mating of the host. Paradoxically, at the population level, parasite transmission also reduces the number of adult males present in the final population for infected females to mate with. Here we investigate if successful female mating when males are rare is indeed a likely rate-limiting step in the transmission of male-killing Spiroplasma in the African Monarch, Danaus chrysippus. In Lepidoptera, successful pairings are hallmarked by the transfer of a sperm-containing spermatophore from the male to the female during copulation. Conveniently, this spermatophore remains detectable within the female upon dissection, and thus, spermatophore counts can be used to assess the frequency of successful mating in the field. We used such spermatophore counts to examine if altered sex ratios in the D. chrysippus do indeed affect female mating success. We examined two different field sites in East Africa where males were often rare. Surprisingly, mated females carried an average of 1.5 spermatophores each, regardless of male frequency, and importantly, only 10-20% remained unmated. This suggests that infected females will still be able to mate in the face of either Spiroplasma-mediated male killing and/or fluctuations in adult sex ratio over the wet-dry season cycle. These observations may begin to explain how the male-killing mollicute can still be successfully transmitted in a population where males are rare.

Exploring the potential effect of COVID-19 on an endangered great ape.

The current COVID-19 pandemic has created unmeasurable damages to society at a global level, from the irreplaceable loss of life, to the massive economic losses. In addition, the disease threatens further biodiversity loss. Due to their shared physiology with humans, primates, and particularly great apes, are susceptible to the disease. However, it is still uncertain how their populations would respond in case of infection. Here, we combine stochastic population and epidemiological models to simulate the range of potential effects of COVID-19 on the probability of extinction of mountain gorillas. We find that extinction is sharply driven by increases in the basic reproductive number and that the probability of extinction is greatly exacerbated if the immunity lasts less than 6 months. These results stress the need to limit exposure of the mountain gorilla population, the park personnel and visitors, as well as the potential of vaccination campaigns to extend the immunity duration.

Rapid transmission of respiratory infections within but not between mountain gorilla groups.

Minimizing disease transmission between humans and wild apes and controlling outbreaks in ape populations is vital to both ape conservation and human health, but information on the transmission of real infections in wild populations is rare. We analyzed respiratory outbreaks in a subpopulation of wild mountain gorillas (Gorilla beringei beringei) between 2004 and 2020. We investigated transmission within groups during 7 outbreaks using social networks based on contact and proximity, and transmission between groups during 15 outbreaks using inter-group encounters, transfers and home range overlap. Patterns of contact and proximity within groups were highly predictable based on gorillas' age and sex. Disease transmission within groups was rapid with a median estimated basic reproductive number (R0) of 4.18 (min = 1.74, max = 9.42), and transmission was not predicted by the social network. Between groups, encounters and transfers did not appear to have enabled disease transmission and the overlap of groups' ranges did not predict concurrent outbreaks. Our findings suggest that gorilla social structure, with many strong connections within groups and weak ties between groups, may enable rapid transmission within a group once an infection is present, but limit the transmission of infections between groups.

Social groups buffer maternal loss in mountain gorillas.

Mothers are crucial for mammals' survival before nutritional independence, but many social mammals reside with their mothers long after. In these species the social adversity caused by maternal loss later in life can dramatically reduce fitness. However, in some human populations these negative consequences can be overcome by care from other group members. We investigated the consequences of maternal loss in mountain gorillas and found no discernible fitness costs to maternal loss through survival, age at first birth, or survival of first offspring through infancy. Social network analysis revealed that relationships with other group members, particularly dominant males and those close in age, strengthened following maternal loss. In contrast to most social mammals, where maternal loss causes considerable social adversity, in mountain gorillas, as in certain human populations, this may be buffered by relationships within cohesive social groups, breaking the link between maternal loss, increased social adversity, and decreased fitness.

Most mammals depend entirely upon their mothers when they are born. In these species, losing a mother at a young age has dramatic consequences for survival. In cases where orphaned individuals do reach adulthood, they often suffer negative effects, like reduced reproductive success or lower social status. But this is not the case for humans. If a child loses their mother; relatives, friends and the wider community can take over. This does not tend to happen in nature. Even our closest relatives, chimpanzees, are much less likely to survive if their mothers die before they reach adolescence. Although orphan survival is not the norm for mammals, humans may not be entirely unique. Mountain gorillas also live in stable family groups, usually with a dominant male and one or more females who care for their offspring for between 8 and 15 years. It is possible that gorillas may also be able to provide community support to orphans, which could buffer the costs of losing a mother, just as it does in humans. To answer this question, Morrison et al. examined 53 years of data collected by the Dian Fossey Gorilla Fund to assess the effects of maternal loss in mountain gorillas. The analysis examined survival, reproduction and changes in social relationships. This revealed that, like humans, young gorillas that lose their mothers are not at a greater risk of dying. There is also no clear long-term effect on their ability to reproduce. In fact, gorillas who lost their mothers ended up with stronger social relationships, especially with the dominant male of the group and young gorillas around the same age. It seems that gorilla social groups, like human families, provide support to young group members that lose their mothers. These findings suggest that the human ability to care for others in times of need may not be unique. It is possible that the tendency to care for orphaned young has its origins in our evolutionary past. Understanding this in more depth could provide clues into the social mechanisms that help to overcome early life adversity, and have a positive impact on future health and survival.