Dissecting the two mechanisms of scramble competition among the Virunga mountain gorillas.

ABSTRACT: Two mechanisms have been proposed to explain why scramble competition can increase the travel requirements of individuals within larger groups. Firstly, individuals in larger groups may be more likely to encounter food sites where other group members have already eaten, leading to greater asynchronous "individual" travel to find fresh sites. Secondly, when food sites are aggregated into patches, larger groups may need to visit more patches to obtain the same amount of food per capita, leading to greater synchronous "group" travel between patches. If the first mechanism can be mitigated by increasing group spread, then we expect the second mechanism to be more sensitive to group size. Here, we examine the individual travel and group travel of the Virunga mountain gorillas, along with potential implications for the two mechanisms of scramble competition. Asynchronous individual travel accounted for 67% of the total travel time, and the remainder arose from group travel. Group spread increased significantly for larger groups, but not enough to prevent an increase in individual travel. Contrary to expectations, group travel decreased with size among most groups, and we found only limited evidence of patch depletion that would cause the second mechanism of scramble competition. Collectively, our results illustrate how the influence of group size can differ for individual travel versus group travel, just as it differs among species for overall travel. Studies that distinguish between the two mechanisms of scramble competition may enhance our understanding of ecological constraints upon group size, including potential differences between frugivores and folivores. SIGNIFICANCE STATEMENT: Feeding competition provides insight into how group size can influence the foraging patterns of social animals, but two key mechanisms are not typically compared. Firstly, larger groups may visit more patches to access the same amount of food per capita (group travel). Secondly, their individuals may also need to move past more spots where another member has already eaten (individual travel). Contrary to expectations, we found that group travel decreased with size for most groups of mountain gorillas, which may reflect extra travel by smaller groups to avoid larger groups. Individual travel increased with size in most groups, even though gorillas in larger groups compensated by spreading out over a broader area. The two mechanisms revealed patterns that were not apparent in our previous study of overall travel. Our approach may help to explain potential differences between folivores and frugivores.

Quadratic relationships between group size and foraging efficiency in a herbivorous primate.

The effect of feeding competition on foraging efficiency is an important link between ecological factors and the social organization of gregarious species. We examined the effects of group size on daily travel distances, activity budgets, and energy intake of mountain gorillas in Rwanda. We measured daily travel distances of five groups, activity budgets of 79 gorillas in nine groups, and energy intake data for 23 adult females in three groups over a 16-month period. Travel distances and the proportion of time spent traveling increased with size for most groups, which would be expected if their foraging efficiency is limited by intragroup feeding competition. However, travel distances and times decreased for the largest group, which also had higher energy intake rates than intermediate sized groups. The improved foraging efficiency of the largest group may be explained by advantages in intergroup contest competition. The largest group had much lower home range overlap than the other study groups which may be due to groups avoiding one another as a result of male mating competition. Collectively, our results indicate that intermediate sized groups had the lowest foraging efficiency and provide a new twist on the growing evidence of non-linear relationships between group size and foraging efficiency in primates.

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.

Energetic responses to variation in food availability in the two mountain gorilla populations (Gorilla beringei beringei).

OBJECTIVE: Here, we compare food availability and relate this to differences in energy intake rates, time spent feeding, and daily travel distance of gorillas in the two populations. Comparative intraspecific studies investigating spatiotemporal variation in food availability can help us understand the complex relationships between ecology, behavior, and life history in primates and are relevant to understanding hominin evolution. Differences in several variables have been documented between the two mountain gorilla populations in the Virunga Massif and Bwindi Impenetrable National Park, but few direct comparisons that link ecological conditions to feeding behavior have been made. MATERIALS AND METHODS: Using similar data collection protocols we conducted vegetation sampling and nutritional analysis on important foods to estimate food availability. Detailed observations of feeding behavior were used to compute energy intake rates and daily travel distance was estimated through GPS readings. RESULTS: Food availability was overall lower and had greater temporal variability in Bwindi than in the Virungas. Energy intake rates and time spent feeding were similar in both populations, but energy intake rates were significantly higher in Bwindi during the period of high fruit consumption. Daily travel distances were significantly shorter in the Virungas. CONCLUSIONS: Overall, despite the differences in food availability, we did not find large differences in the energetics of gorillas in the two populations, although further work is needed to more precisely quantify energy expenditure and energy balance. These results emphasize that even species with high food availability can exhibit behavioral and energetic responses to variable ecological conditions, which are likely to affect growth, reproduction, and survival.

Long-term temporal and spatial dynamics of food availability for endangered mountain gorillas in Volcanoes National Park, Rwanda.

Monitoring temporal and spatial changes in the resource availability of endangered species contributes to their conservation. The number of critically endangered mountain gorillas (Gorilla beringei beringei) in the Virunga Volcano population has doubled over the past three decades, but no studies have examined how food availability has changed during that period. First, we assessed if the plant species consumed by the gorillas have changed in abundance and distribution during the past two decades. In 2009-2010, we replicated a study conducted in 1988-1989 by measuring the frequency, density, and biomass of plant species consumed by the gorillas in 496 plots (ca. 6 km(2)) in the Karisoke study area in Volcanoes National Park, Rwanda. We expected to observe a decreased presence of major gorilla food plants as a likely result of density-dependent overharvesting by gorillas. Among the five most frequently consumed species (composing approximately 70% of the gorilla's diet, excluding bamboo), two have decreased in availability and abundance, while three have increased. Some species have undergone shifts in their altitudinal distribution, possibly due to regional climatic changes. Second, we made baseline measurements of food availability in a larger area currently utilized by the gorillas. In the extended sampling (n = 473 plots) area (ca. 25 km(2) ), of the five most frequently consumed species, two were not significantly different in frequency from the re-sampled area, while two occurred significantly less frequently, and one occurred significantly more frequently. We discuss the potential impact of gorilla-induced herbivory on changes of vegetation abundance. The changes in the species most commonly consumed by the gorillas could affect their nutrient intake and stresses the importance of monitoring the interrelation among plant population dynamics, species density, and resource use.