Dominance rank but not body size influences female reproductive success in mountain gorillas

According to life history theory, natural selection has shaped trade-offs for allocating energy among growth, reproduction and maintenance to maximize individual fitness. In social mammals body size and dominance rank are two key variables believed to influence female reproductive success. However, few studies have examined these variables together, particularly in long-lived species. Previous studies found that female dominance rank correlates with reproductive success in mountain gorillas (Gorilla beringei beringei), which is surprising given they have weak dominance relationships and experience seemingly low levels of feeding competition. It is not currently known whether this relationship is primarily driven by a positive correlation between rank and body size. We used the non-invasive parallel laser method to measure two body size variables (back breadth and body length) of 34 wild adult female mountain gorillas, together with long-term dominance and demography data to investigate the interrelationships among body size, dominance rank and two measures of female reproductive success (inter-birth interval N = 29 and infant mortality N = 64). Using linear mixed models, we found no support for body size to be significantly correlated with dominance rank or female reproductive success. Higher-ranking females had significantly shorter inter-birth intervals than lower-ranking ones, but dominance rank was not significantly correlated with infant mortality. Our results suggest that female dominance rank is primarily determined by factors other than linear body dimensions and that high rank provides benefits even in species with weak dominance relationships and abundant year-round food resources. Future studies should focus on the mechanisms behind heterogeneity in female body size in relation to trade-offs in allocating energy to growth, maintenance and lifetime reproductive success

3D enamel profilometry reveals faster growth but similar stress severity in Neanderthal versus Homo sapiens teeth

Early life stress disrupts growth and creates horizontal grooves on the tooth surface in humans and other mammals, yet there is no consensus for their quantitative analysis. Linear defects are considered to be nonspecific stress indicators, but evidence suggests that intermittent, severe stressors create deeper defects than chronic, low-level stressors. However, species-specific growth patterns also influence defect morphology, with faster-growing teeth having shallower defects at the population level. Here we describe a method to measure the depth of linear enamel defects and normal growth increments (i.e., perikymata) from high-resolution 3D topographies using confocal profilometry and apply it to a diverse sample of Homo neanderthalensis and H. sapiens anterior teeth. Debate surrounds whether Neanderthals exhibited modern human-like growth patterns in their teeth and other systems, with some researchers suggesting that they experienced more severe childhood stress. Our results suggest that Neanderthals have shallower features than H. sapiens from the Upper Paleolithic, Neolithic, and medieval eras, mirroring the faster growth rates in Neanderthal anterior teeth. However, when defect depth is scaled by perikymata depth to assess their severity, Neolithic humans have less severe defects, while Neanderthals and the other H. sapiens groups show evidence of more severe early life growth disruptions

Facial asymmetry tracks genetic diversity among Gorilla subspecies

Mountain gorillas are particularly inbred compared to other gorillas and even the most inbred human populations. As mountain gorilla skeletal material accumulated during the 1970s, researchers noted their pronounced facial asymmetry and hypothesized that it reflects a population-wide chewing side preference. However, asymmetry has also been linked to environmental and genetic stress in experimental models. Here, we examine facial asymmetry in 114 crania from three Gorilla subspecies using 3D geometric morphometrics. We measure fluctuating asymmetry (FA), defined as random deviations from perfect symmetry, and population-specific patterns of directional asymmetry (DA). Mountain gorillas, with a current population size of about 1000 individuals, have the highest degree of facial FA (explaining 17% of total facial shape variation), followed by Grauer gorillas (9%) and western lowland gorillas (6%), despite the latter experiencing the greatest ecological and dietary variability. DA, while significant in all three taxa, explains relatively less shape variation than FA does. Facial asymmetry correlates neither with tooth wear asymmetry nor increases with age in a mountain gorilla subsample, undermining the hypothesis that facial asymmetry is driven by chewing side preference. An examination of temporal trends shows that stress-induced developmental instability has increased over the last 100 years in these endangered apes

Skeletal ageing in Virunga mountain gorillas.

Bone loss and heightened fracture risk are common conditions associated with ageing in modern human populations and have been attributed to both hormonal and other metabolic and behavioural changes. To what extent these age-related trends are specific to modern humans or generally characteristic of natural populations of other taxa is not clear. In this study, we use computed tomography to examine age changes in long bone and vertebral structural properties of 34 wild-adult Virunga mountain gorillas (Gorilla beringei beringei) whose skeletons were recovered from natural accumulations. Chronological ages were known or estimated from sample-specific dental wear formulae and ranged between 11 and 43 years. Gorillas show some of the same characteristics of skeletal ageing as modern humans, including endosteal and some periosteal expansion. However, unlike in humans, there is no decline in cortical or trabecular bone density, or in combined geometric-density measures of strength, nor do females show accelerated bone loss later in life. We attribute these differences to the lack of an extended post-reproductive period in gorillas, which provides protection against bone resorption. Increases in age-related fractures (osteoporosis) in modern humans may be a combined effect of an extended lifespan and lower activity levels earlier in life. This article is part of the theme issue 'Evolution of the primate ageing process'

Phylogenetic and environmental effects on limb bone structure in gorillas

Objectives The effects of phylogeny and locomotor behavior on long bone structural proportions are assessed through comparisons between adult and ontogenetic samples of extant gorillas. Materials and Methods A total of 281 wild-collected individuals were included in the study, divided into four groups that vary taxonomically and ecologically: western lowland gorillas (G. g. gorilla), lowland and highland grauer gorillas (G. b. graueri), and Virunga mountain gorillas (G. b. beringei). Lengths and articular breadths of the major long bones (except the fibula) were measured, and diaphyseal cross-sectional geometric properties determined using computed tomography. Ages of immature specimens (n = 145) were known or estimated from dental development. Differences between groups in hind limb to forelimb proportions were assessed in both adults and during development. Results Diaphyseal strength proportions among adults vary in parallel with behavioral/ecological differences, and not phylogeny. The more arboreal western lowland and lowland grauer gorillas have relatively stronger forelimbs than the more terrestrial Virunga mountain gorillas, while the behaviorally intermediate highland grauer gorillas have intermediate proportions. Diaphyseal strength proportions are similar in young infants but diverge after 2 years of age in western lowland and mountain gorillas, at the same time that changes in locomotor behavior occur. There are no differences between groups in length or articular proportions among either adults or immature individuals. Conclusion Long bone diaphyseal strength proportions in gorillas are developmentally plastic, reflecting behavior, while length and articular proportions are much more genetically canalized. These findings have implications for interpreting morphological variation among fossil taxa

Tooth wear and feeding ecology in mountain gorillas from Volcanoes National Park, Rwanda

Objectives: Ecological factors have a dramatic effect on tooth wear in primates, although it remains unclear how individual age contributes to functional crown morphology. The aim of this study is to determine how age and individual diet are related to tooth wear in wild mountain gorillas (Gorilla beringei beringei) from Volcanoes National Park, Rwanda. Material and Methods: We calculated the percent of dentine exposure (PDE) for all permanent molars (M1–M3) of known-age mountain gorillas (N = 23), to test whether PDE varied with age using regression analysis. For each molar position, we also performed stepwise multiple linear regression to test the effects of age and percentage of time spent feeding on different food categories on PDE, for individuals subject to long-term observational studies by the Dian Fossey Gorilla Fund International's Karisoke Research Center. Results: PDE increased significantly with age for both sexes in all molars. Moreover, a significant effect of gritty plant root consumption on PDE was found among individuals. Our results support prior reports indicating reduced tooth wear in mountain gorillas compared to western gorillas, and compared to other known-aged samples of primate taxa from forest and savanna habitats. Discussion: Our findings corroborate that mountain gorillas present very low molar wear, and support the hypothesis that age and the consumption of particular food types, namely roots, are significant determinants of tooth wear variation in mountain gorillas. Future research should characterize the mineral composition of the soil in the Virunga habitat, to test the hypothesis that the physical and abrasive properties of gritty foods such as roots influence intra- and interspecific patterns of tooth wear.Grant sponsor: National Science Foundation; Grant numbers: BCS 0852866, 0964944; Grant sponsors: The Leakey Foundation; National Geographic Society’s Committee for Exploration and Research; the Center and Institute Facilitating Fund of The George Washington University; the 2010 Max Planck Research Award endowed by the German Federal Ministry of Education and Research to the Max Planck Society; Alexander von Humboldt Foundation

Facial asymmetry tracks genetic diversity among Gorilla subspecies

Mountain gorillas are particularly inbred compared to other gorillas and even the most inbred human populations. As mountain gorilla skeletal material accumulated during the 1970s, researchers noted their pronounced facial asymmetry and hypothesized that it reflects a population-wide chewing side preference. However, asymmetry has also been linked to environmental and genetic stress in experimental models. Here, we examine facial asymmetry in 114 crania from three Gorilla subspecies using 3D geometric morphometrics. We measure fluctuating asymmetry (FA), defined as random deviations from perfect symmetry, and population-specific patterns of directional asymmetry (DA). Mountain gorillas, with a current population size of about 1000 individuals, have the highest degree of facial FA (explaining 17% of total facial shape variation), followed by Grauer gorillas (9%) and western lowland gorillas (6%), despite the latter experiencing the greatest ecological and dietary variability. DA, while significant in all three taxa, explains relatively less shape variation than FA does. Facial asymmetry correlates neither with tooth wear asymmetry nor increases with age in a mountain gorilla subsample, undermining the hypothesis that facial asymmetry is driven by chewing side preference. An examination of temporal trends shows that stress-induced developmental instability has increased over the last 100 years in these endangered apes