Comparative life history patterns of female gorillas.

OBJECTIVES: 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). MATERIALS AND METHODS: 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). RESULTS: 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. DISCUSSION: 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.

Population dynamics of western gorillas at Mbeli Bai.

Long-term studies of population dynamics can provide insights into life history theory, population ecology, socioecology, conservation biology and wildlife management. Here we examine 25 years of population dynamics of western gorillas at Mbeli Bai, a swampy forest clearing in Nouabalé-Ndoki National Park, the Republic of Congo. The Mbeli population more than doubled from 101 to 226 gorillas during the study. After adjusting for a net influx of gorillas into the study population, the increase represents an inherent growth rate of 0.7% per year, with 95% confidence limits between -0.7% and 2.6%. The influx of gorillas mainly involved immigration of individuals into existing study groups (social dispersal), but it also included the appearance of a few previously unknown groups (locational dispersal). The average group size did not change significantly during the study, which is consistent with the possibility that western gorillas face socioecological constraints on group size, even when the population is increasing. We found no significant evidence of density dependence on female reproductive success or male mating competition. The distribution of gorillas among age/sex categories also did not change significantly, which suggests that the population had a stable age structure. Our results provide evidence of population stability or growth for some western gorillas (albeit within a small area). The results highlight the value of law enforcement, long-term monitoring, and protected areas; but they do not diminish the importance of improving conservation for this critically endangered species.

The long lives of primates and the 'invariant rate of ageing' hypothesis.

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.

Infant survival in western lowland gorillas after voluntary dispersal by pregnant females.

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.

Hierarchical social modularity in gorillas.

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.

Mothers may shape the variations in social organization among gorillas.

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.