Long-term home range stability provides foraging benefits in western gorillas.

OBJECTIVES: Long-term home range stability presumably emerges because familiarity with an area improves fitness through increased foraging efficiency, reduced predation risk, or reduced costs of intergroup aggression. While the use of spatial memory by primates has been widely demonstrated, few studies have examined whether long-term space use creates opportunities for interannual reuse of spatial knowledge. Here we examine the ranging behavior of western gorillas (Gorilla gorilla) to assess the degree of long-term site fidelity and the foraging consequences of reuse of space. METHODS: We measured interannual home range overlap over a 10-year period for a single group of gorillas at the Mondika Research Center, using both grid-based and kernel density estimation. By plotting the total area used over time, we identified periods of home-range stability and expansion. We compared foraging and ranging behavior in familiar versus unfamiliar areas, considering fruit trees visited, dietary diversity, and daily path length, to determine whether the lack of spatial knowledge in unfamiliar areas was associated with foraging costs. RESULTS: Average interannual home range overlap by the group remained high throughout the study. During periods of home range expansion, daily path lengths increased but not the number of fruit trees visited, suggesting that reduced familiarity with the area led to decreased foraging efficiency because individuals lacked prior knowledge of where to find resources. DISCUSSION: Western gorillas at Mondika exhibit long-term home range stability, presumably reflecting a strategy that relies on the use of spatial memory to increase foraging efficiency that is favored by their reliance on ephemeral fruit resources.

Spatial cognition in western gorillas (Gorilla gorilla): an analysis of distance, linearity, and speed of travel routes.

Spatial memory allows animals to retain information regarding the location, distribution, and quality of feeding sites to optimize foraging decisions. Western gorillas inhabit a complex environment with spatiotemporal fluctuations of resource availability, prefer fruits when available, and travel long distances to reach them. Here, we examined movement patterns-such as linearity, distance, and speed of traveling-to assess whether gorillas optimize travel when reaching out-of-sight valued resources. Our results show that gorillas travel patterns are affected by the activity they perform next, the type of food they feed on, and their preference level to specific fruits, suggesting they are able to optimize foraging based on spatial knowledge of their resources. Additionally, gorillas left in the direction of the next resource as soon as they started traveling and decelerated before approaching food resources, as evidence that they have a representation of their exact locations. Moreover, home range familiarity did not influence gorillas' movement patterns, as travel linearity in the core and periphery did not differ, suggesting that they may not depend wholly on a network of paths to navigate their habitat. These results show some overlap with chimpanzees' spatial abilities. Differences between the two ape species exist, however, potentially reflecting more their differences in diet (degree of frugivory) rather than their cognitive abilities. Further studies should focus on determining whether gorillas are able to use shortcuts and/or approach the same goal from multiple directions to better identify the spatial abilities used by this species.

Against all odds: Numerical assessment by tufted capuchin monkeys.

When competitors are able to assess the asymmetry in the resource holding potential before interacting, individuals or groups should avoid interacting with stronger opponents, thereby avoiding the energy costs and risk of injury associated with aggressive intergroup encounters. Thus, escalated aggression is expected only between closely matched competitors. Among Argentine tufted capuchin monkeys (Sapajus nigritus), intergroup dominance is decided by the asymmetry in male group size. Using playback experiments, I simulated intergroup encounters with neighboring groups, manipulating both the apparent numerical asymmetry and the resource context. During experimental trials, I recorded the approach behavior of the focal individual, as well as changes in neighbor density and individual travel speed following the presentation of the playback stimulus, to assess whether individual willingness to participate in resource defense was affected by the probability of winning the encounter. In spite of the competitive disadvantage, neither males nor females showed a decreased probability of approach when the numerical odds strongly favored the opposing group. Instead decisions regarding whether to participate appear to be driven primarily by the resource context. Nevertheless, changes in individual behavior during approaches suggest that tufted capuchin monkeys are sensitive to the relative odds. Individuals accelerated less when approaching a larger group, although no changes in neighbor density were apparent. The absence of an effect of the numerical asymmetry on willingness to approach the playback speaker suggests that subordinate groups benefit from engaging in intergroup aggression with larger neighbors, despite the high probability of losing. These encounters may serve to assess the current subjective resource valuation of the neighboring group or limit territorial expansion by large groups by decreasing the marginal value of home range exclusivity. Because these encounters are riskier, however, individuals appear to alter their approaches, becoming more tentative as the numerical odds increasingly favor the opposing group.

Group benefit associated with polymorphic trichromacy in a Malagasy primate (Propithecus verreauxi).

In some primate lineages, polymorphisms in the X-linked M/LWS opsin gene have produced intraspecific variation in color vision. In these species, heterozygous females exhibit trichromacy, while males and homozygous females exhibit dichromacy. The evolutionary persistence of these polymorphisms suggests that balancing selection maintains color vision variation, possibly through a 'trichromat advantage' in detecting yellow/orange/red foods against foliage. We identified genetic evidence of polymorphic trichromacy in a population of Verreaux's sifaka (Propithecus verreauxi) at Kirindy Mitea National Park in Madagascar, and explored effects of color vision on reproductive success and feeding behavior using nine years of morphological, demographic, and feeding data. We found that trichromats and dichromats residing in social groups with trichromats exhibit higher body mass indices than individuals in dichromat-only groups. Additionally, individuals in a trichromat social group devoted significantly more time to fruit feeding and had longer fruit feeding bouts than individuals in dichromat-only groups. We hypothesize that, due to small, cohesive sifaka social groups, a trichromat advantage in detecting productive fruit patches during the energetically stressful dry season also benefits dichromats in a trichromat's group. Our results offer the first support for the 'mutual benefit of association' hypothesis regarding the maintenance of polymorphic trichromacy in primates.

Individual participation in intergroup contests is mediated by numerical assessment strategies in black howler and tufted capuchin monkeys.

Asymmetries in resource-holding potential between opposing groups frequently determine outcomes of intergroup contests. Since both numerical superiority and high intergroup dominance rank may confer competitive advantages, group members should benefit from assessing the relative strength of rivals prior to engaging in defensive displays. However, differences in individual assessment may emerge when cost-benefit trade-offs differ among group members. We examine the influence of numerical superiority and intergroup dominance relationships on individual participation in intergroup encounters in black howler monkeys (Alouatta pigra) and tufted capuchin monkeys (Sapajus nigritus). Black howlers responded with longer vocal displays during encounters with neighbours with an equal number of resident males, while tufted capuchins increased their participation with increasing relative male group size. Within each species, males and females responded similarly to varying numerical odds, suggesting that despite pay-off asymmetries between males and females, both sexes were similarly influenced by numerical asymmetries in deciding to participate in collective group defence. Whereas the outcome of contests among tufted capuchins was determined by relative male group size, reflected in a pronounced intergroup dominance hierarchy, the absence of dominance relationships among black howler groups may have provoked prolonged vocal displays in order to assess rival groups with matching competitive abilities.

Rates of agonism among female primates: a cross-taxon perspective.

Agonism is common in group-living animals, shaping dominance relationships and ultimately impacting individual fitness. Rates of agonism vary considerably among taxa, however, and explaining this variation has been central in ecological models of female social relationships in primates. Early iterations of these models posited a link to diet, with more frequent agonism predicted in frugivorous species due to the presumed greater contestability of fruits relative to other food types. Although some more recent studies have suggested that dietary categories may be poor predictors of contest competition among primates, to date there have been no broad, cross-taxa comparisons of rates of female-female agonism in relation to diet. This study tests whether dietary variables do indeed predict rates of female agonism and further investigates the role of group size (i.e., number of competitors) and substrate use (i.e., degree of arboreality) on the frequency of agonism. Data from 44 wild, unprovisioned groups, including 3 strepsirhine species, 3 platyrrhines, 5 colobines, 10 cercopithecines, and 2 hominoids were analyzed using phylogenetically controlled and uncontrolled methods. Results indicate that diet does not predict agonistic rates, with trends actually being in the opposite direction than predicted for all taxa except cercopithecines. In contrast, agonistic rates are positively associated with group size and possibly degree of terrestriality. Competitor density and perhaps the risk of fighting, thus, appear more important than general diet in predicting agonism among female primates. We discuss the implications of these results for socio-ecological hypotheses.

Variation in grouping patterns, mating systems and social structure: what socio-ecological models attempt to explain.

Socio-ecological models aim to predict the variation in social systems based on a limited number of ecological parameters. Since the 1960s, the original model has taken two paths: one relating to grouping patterns and mating systems and one relating to grouping patterns and female social structure. Here, we review the basic ideas specifically with regard to non-human primates, present new results and point to open questions. While most primates live in permanent groups and exhibit female defence polygyny, recent studies indicate more flexibility with cooperative male resource defence occurring repeatedly in all radiations. In contrast to other animals, the potential link between ecology and these mating systems remains, however, largely unexplored. The model of the ecology of female social structure has often been deemed successful, but has recently been criticized. We show that the predicted association of agonistic rates and despotism (directional consistency of relationships) was not supported in a comparative test. The overall variation in despotism is probably due to phylogenetic grade shifts. At the same time, it varies within clades more or less in the direction predicted by the model. This suggests that the model's utility may lie in predicting social variation within but not across clades.

Consequences of lethal intragroup aggression and alpha male replacement on intergroup relations and home range use in tufted capuchin monkeys (Cebus apella nigritus).

In conflicts between primate groups, the resource-holding potential (RHP) of competitors is frequently related to group size or male group size, which can remain relatively constant for long periods of time, promoting stable intergroup dominance relationships. Demographic changes in neighboring groups, however, could introduce uncertainty into existing relationships. Among tufted capuchin monkeys (Cebus apella nigritus), dominant male replacement is a relatively infrequent demographic event that can have a profound effect on both the composition and size of the social group. Here, we report such a case and the consequences for home range use and intergroup aggression. Between June 2008 and August 2010, we periodically followed two neighboring groups (Macuco and Rita) in Iguazú National Park, recording daily paths (N = 143) and encounters between the groups (N = 28). We describe the events leading to a change in the male dominance hierarchy in the larger group (Macuco), which resulted in the death or dispersal of all adult males, followed by the succession of a young adult male to the dominant position. This takeover event reduced the numerical advantage in number of males between the two groups, although the ratio of total group sizes remained nearly constant. Following this shift in numerical asymmetry, the degree of escalation of intergroup aggression increased, and we observed reversals in the former intergroup dominance relationship. These changes in behavior during intergroup encounters were associated with changes in the use of overlapping areas. In the 6 months following the takeover, the area of home range overlap doubled, and the formerly dominant group's area of exclusive access was reduced by half. These results suggest that RHPin tufted capuchin monkeys is related to male group size. Furthermore, they highlight the importance of considering rare demographic events in attempts to understand the dynamics of aggression between primate groups.