Modern human societies show hierarchical social modularity (HSM) in which lower-order social units like nuclear families are nested inside increasingly larger units. It has been argued that this HSM evolved independently and after the chimpanzee–human split due to greater recognition of, and bonding between, dispersed kin. We used network modularity analysis and hierarchical clustering to quantify community structure within two western lowland gorilla populations. In both communities, we detected two hierarchically nested tiers of social structure which have not been previously quantified. Both tiers map closely to human social tiers. Genetic data from one population suggested that, as in humans, social unit membership was kin structured. The sizes of gorilla social units also showed the kind of consistent scaling ratio between social tiers observed in humans, baboons, toothed whales, and elephants. These results indicate that the hierarchical social organization observed in humans may have evolved far earlier than previously asserted and may not be a product of the social brain evolution unique to the hominin lineage.

Is it possible to slow the rate of ageing, or do biological constraints limit its plasticity? We test the ‘invariant rate of ageing’ hypothesis, which posits that the rate of ageing is relatively fixed within species, with a collection of 39 human and nonhuman primate datasets across seven genera. We first recapitulate, in nonhuman primates, the highly regular relationship between life expectancy and lifespan equality seen in humans. We next demonstrate that variation in the rate of ageing within genera is orders of magnitude smaller than variation in pre-adult and age-independent mortality. Finally, we demonstrate that changes in the rate of ageing, but not other mortality parameters, produce striking, species-atypical changes in mortality patterns. Our results support the invariant rate of ageing hypothesis, implying biological constraints on how much the human rate of ageing can be slowed.

In this paper we report on the context and evaluation of a system for an automatic interpretation of sightings of individual western lowland gorillas (Gorilla gorilla gorilla) as captured in facial fieldVisual data acquisition in the field often captures sufficient information to establish encounters with individual gorillas. However, relevant information is locked within the pixel patterns measured, usually requiring expert knowledge and time-consuming efforts for identification. Computer vision can help to extract gorilla identities by performing automated species detection, followed by individual facial identification. We show that standard deep learning models combined with a traditional SVM classifier can be used for this task. To assist encounter processing, predictions can be presented graphically with known population information as shown.

Evaluating the factors influencing the patterns of female dispersal in mammals is critical to understanding its importance for male and female reproductive strategies and the evolution of social systems. In western lowland gorillas, females emigrate from their natal group (natal dispersal) but also disperse between groups multiple times in their life (secondary dispersal). This strategy is uncommon in primates and may reflect females seeking to reduce feeding competition or female choice for males that can provide protection against infanticide. In this study, we test how inbreeding avoidance, group size, and male competitive ability could influence female dispersal in western gorillas, using 25 natal and 285 secondary dispersal events collected over 20 years at Mbeli Bai. We found that while all females dispersed out of their natal group, presumably to avoid inbreeding, females also left their group before having their first offspring when the dominant male was not their father, pointing towards unconditional dispersal by nulliparous females. Regarding secondary dispersal, we found that females were more likely to stay with males that were at the beginning of their tenure than transfer and more likely to disperse away from (presumably old) males that were near the end of their tenure or closer to their death. Females were also more likely to leave larger groups for smaller ones suggesting a potential effect of within-group feeding competition or avoidance of outsider males or predators. This study highlights the intersexual conflict found in gorillas: while female choice for high-quality males may influence the formation of their relatively rare social system, smaller group size may be more important for females than previously thought, which runs counter to the males' interest of having a high number of females. SignificanceDispersal is a very important life strategy for most group-living mammals. Secondary dispersal by females is a rare behavior but is believed to be a counterstrategy against sexual coercion by males, reduces feeding competition, assists in predator avoidance, and facilitates mate choice. We studied the factors influencing secondary dispersal in western lowland gorillas. We found that the duration of male tenure and the size of the group influences female transfer decisions. This study shows that female reproductive strategies such as secondary dispersal may be very powerful to counter the risk of infanticide through female choice for better protector males but also to reduce the impact of feeding competition or to avoid predators or outsider males.

When mothers continue to support their offspring beyond infancy, they can influence the fitness of those offspring, the strength of social relationships within their groups, and the life-history traits of their species. Using up to 30 years of demographic data from 58 groups of gorillas in two study sites, this study extends such findings by showing that mothers may also contribute to differences in social organization between closely related species. Female mountain gorillas remained with their sons for significantly longer than western gorillas, which may explain why male philopatry and multimale groups are more common among mountain gorillas. The presence of the putative father and other familiar males did not vary significantly between species, and we found only limited support for the socio-ecological theory that the distribution of adult males is influenced by the distribution of females. Within each gorilla species, variations in those distributions may also reflect the different stages in the typical life cycle of a group. Collectively, our results highlight the potentially far-reaching consequences of maternal support that extends beyond infancy, and they illustrate the opportunity to incorporate additional factors into phylogenetic analyses of variations in social organization, including studies of human evolution.

In group-living mammals, individual efforts to maximize reproductive success result in conflicts and compromises between the sexes. Females utilize counterstrategies to minimize the costs of sexual coercion by males, but few studies have examined the effect of such behaviors on female reproductive success. Secondary dispersal by females is rare among group-living mammals, but in western gorillas, it is believed to be a mate choice strategy to minimize infanticide risk and infant mortality. Previous research suggested that females choose males that are good protectors. However, how much female reproductive success varies depending on male competitive ability and whether female secondary dispersal leads to reproductive costs or benefits has not been examined. We used data on 100 females and 229 infants in 36 breeding groups from a 20-year long-term study of wild western lowland gorillas to investigate whether male tenure duration and female transfer rate had an effect on interbirth interval, female birth rates, and offspring mortality. We found that offspring mortality was higher near the end of males' tenures, even after excluding potential infanticide when those males died, suggesting that females suffer a reproductive cost by being with males nearing the end of their tenures. Females experience a delay in breeding when they dispersed, having a notable effect on birth rates of surviving offspring per female if females transfer multiple times in their lives. This study exemplifies that female counterstrategies to mitigate the effects of male-male competition and sexual coercion may not be sufficient to overcome the negative consequences of male behavior. Significance Individual reproductive success is maximized through conflicts and compromises between the sexes in social mammals. In species with high dimorphism, females can experience sexual coercion by males and develop counterstrategies to reduce these costs. We studied this link in western lowland gorillas, which exhibit female secondary dispersal, a strategy that is believed to reflect female choice for the protective abilities of males. We found that females are faced with the dilemma of staying with a silverback at the end of his tenure and risk higher infant mortality versus dispersing and suffering reproductive delays and lower Communicated by M. A. van Noordwijk Electronic supplementary material The online version of this article (

Zoos and aquariums undoubtably have an important role to play in supporting species conservation through a diverse range of activities, including public awareness and financial support of field‐conservation projects. Charismatic megafauna, such as gorillas Gorilla spp, play a crucial role as flagship species in their range countries, acting as ambassadors for conservation. Here, we present examples from the Mbeli Bai Study (MBS) in northern Congo, a long‐term Western lowland gorilla Gorilla gorilla gorilla field‐conservation project. We show how collaborations between zoos and field‐conservation projects can support each other in numerous ways. Over the last two decades the MBS has received substantial support from zoos. A large portion of the operational budget has been provided by zoos and this has ensured a degree of financial independence over the long term. Intensive exchange of conservation‐education techniques in terms of material and staff expertise has been helpful for MBS's education project ‘Club Ebobo’. This has not only provided training for field educators but has also given zoo personnel and visitors better insight into the realities of conservation in range countries. Such exchanges can further help to promote awareness, raising the role of zoos in supporting field‐conservation projects. Finally, the knowledge gained through scientific research has proven to be beneficial for animals both in the field (e.g. improved conservation measures) and in captivity (e.g. better‐adapted group structure, feeding strategies and environment). Here, we discuss our experience and describe some of the benefits and challenges of conservation collaborations between zoos and field‐conservation projects.

In many social species, after the alpha male has been replaced or the group disintegrates, a female’s infant is at risk of infanticide by a male. Female gorillas have developed the rare strategy of secondary dispersal in which they transfer between reproductive groups during the limited time period between weaning an infant and conceiving the next one (voluntary dispersal). By doing so they leave a weaker silverback near the end of his tenure and join a stronger silverback at an earlier stage of his own tenure, thereby mitigating the risk of infanticide if the former dies. If females are pregnant or have unweaned offspring when the only male in the group dies, their offspring are vulnerable to infanticide by the new silverback that they join (via involuntary dispersal). In the few known cases of female gorillas transferring when pregnant (mainly after group disintegration), their offspring were killed. We report here on three adult females that transferred voluntarily while pregnant multiple times between two groups yet their offspring were not killed by the new group’s silverback. The gorillas were observed from 1995 to 2015 at the Mbeli Bai research site in northern Republic of the Congo. The females gave birth 5–6 months (gestation period 8.5 months) after their last transfer. To our knowledge, these observations are the first to show that wild female western lowland gorillas can transfer voluntarily while pregnant without incurring infanticide by a new silverback. These observations highlight the behavioural plasticity shown by female gorillas in response to sexual coercion by males.