Colleen M. Schaffner

Fission-Fusion Dynamics: New Research Frameworks

Renewed interest in fission‐fusion dynamics is due to the recognition that such dynamics may create unique challenges for social interaction and distinctive selective pressures acting on underlying communicative and cognitive abilities. New frameworks for integrating current knowledge on fission‐fusion dynamics emerge from a fundamental rethinking of the term “fission‐fusion” away from its current general use as a label for a particular modal type of social system (i.e., “fission‐fusion societies”). Specifically, because the degree of spatial and temporal cohesion of group members varies both within and across taxa, any social system can be described in terms of the extent to which it expresses fission‐fusion dynamics. This perspective has implications for socioecology, communication, cognitive demands, and human social evolution.

Relationship assessment through emotional mediation

Social relationship is a concept that links the observable social interactions between group members to the inferred group social system. Social relationships allow animals (as well as the human observers) to predict the actions and responses of their partners and therefore guide their own. Social relationships can also be described as investments that benefit the individuals involved in them. Some benefits simply require stable association and some level of mutual tolerance, whereas others depend on the establishment of more fully developed social relationships. The variation in the quality of social relationships leads to a great flexibility in the frequency and quality of interaction with various group members and with the same individual over time. A key issue is therefore to understand the proximate mechanisms underlying such flexibility since individuals need to be able to assess relationship quality in order to maximise the benefits that social relationships provide. Assessment of social relationships should be based on the information contained in the various interactions that the partners exchange. Assessment should therefore require bookkeeping of the various interactions, computation of their relative frequencies, and conversion of their quality and information associated with them into common currencies. We propose emotional mediation as a possible mechanism that fulfils such requirements and provides the individual with a timely assessment to guide its social decision. Emotions are viewed as intervening variables that result from the integration of the information contained in the various interactions between two partners. Before presenting evidence for relationship assessment through emotional mediation, we define the concept of emotion in animals and provide evidence for measuring relevant emotions in non-human primates. Then, we present four examples obtained by combining findings from multiple studies. The examples provide evidence for emotion resulting from information contained in the interactions exchanged by the partners and for emotional differences being at the basis of the variation in social interactions. Thus, animals may appreciate variation in their social relationships through emotional mediation. This is a promising avenue to disclose the proximate mechanisms of relationship assessment and we suggest new lines of research to gather further evidence for the role of emotional mediation.

Intragroup aggression, fission–fusion dynamics and feeding competition in spider monkeys

Scramble competition is related to animals depleting resources before others can use them, whereas contest competition is associated with the monopolization of resources and food-related aggression. One hypothesized major benefit of fission–fusion sociality is the reduction of scramble feeding competition between community members. We studied intragroup aggression in a community of spider monkeys. Despite seasonal variation in food availability, we found no seasonal difference in aggression rates, which, in combination with a basic lack of aggression among adult females, suggests that fission–fusion dynamics may reduce not only scramble, but also the intensity of contest competition. There were, however, numerous attacks from adult females towards subadult females, especially new immigrants. This aggression, although it occurred mainly during feeding, may not reflect contest competition. Instead, it may be used by resident females to limit scramble competition at the community level, which is ultimately affected by the number of community members. The aggressive harassment of subadult females by long-term resident females can be a means to encourage dispersal of natal subadult females and discourage immigration of new subadult females who are both potential future resource competitors.

Aggression and conflict management at fusion in spider monkeys

In social systems characterized by a high degree of fission–fusion dynamics, members of a large community are rarely all together, spending most of their time in smaller subgroups with flexible membership. Although fissioning into smaller subgroups is believed to reduce conflict among community members, fusions may create conflict among individuals from joining subgroups. Here, we present evidence for aggressive escalation at fusion and its mitigation by the use of embraces in wild spider monkeys (Ateles geoffroyi). Our findings provide the first systematic evidence for conflict management at fusion and may have implications for the function of human greetings.

An explicit signature of balancing selection for color vision variation in New World monkeys

Color vision is an important characteristic of primates and, intriguingly, Neotropical monkeys are highly polymorphic for this trait. Recent field studies have challenged the conventional view that trichromatic color vision is more adaptive than dichromatic color vision. No study has investigated the pattern of genetic variation in the long to middle wavelength-sensitive (L-M or red-green) opsin gene as compared with that of other genomic regions (neutral references) in wild populations of New World monkeys to look for the signature of natural selection. Here, we report such a study conducted on spider monkeys and capuchin monkeys inhabiting Santa Rosa National Park, Costa Rica. The nucleotide sequence of the L-M opsin gene was more polymorphic than the sequences of the neutral references, although the opsin-gene sequences were not more divergent between the two species than were the sequences of the neutral references. In a coalescence simulation that took into account the observed nucleotide diversity of the neutral references, the Tajimas D value of the L-M opsin gene deviated significantly in a positive direction from the expected range. These results are the first to statistically demonstrate balancing selection acting on the polymorphic L-M opsin gene of New World monkeys. Taking the results of behavioral and genetic studies together, the balancing selection we detected may indicate that coexistence of different color-vision types in the same population, also characteristic of humans, is adaptive.

Embraces and grooming in captive spider monkeys

Relatively little information is available regarding the role of social grooming and embraces in spider monkeys that live in fission-fusion societies in which individuals are usually split into subgroups. We investigated whether embraces and grooming have similar roles in captive spider monkeys by examining their occurrence in two contexts. One context was fusion, i.e. when the monkeys moved into an area of the enclosure where other monkeys were present, in which individuals from different subgroups were expected to exchange greeting behavior. The other context was the access to young infants, in which females were expected increase their friendly behavior toward mothers. We collected data by observing all individuals within a subgroup and via focal animal sampling. We found that overall embraces occurred more frequently than grooming and that their distributions were not correlated. The frequency of embraces was positively associated with the number of fusion events per observation, whereas the frequency of grooming bouts was not. Furthermore, embraces were more frequent following initial approaches after fusion versus subsequent approaches, and the figure was higher than the corresponding one for grooming. Mothers received more embraces after than before the birth of their infants, whereas there was no such effect for grooming. Embraces, but not grooming, play a role in the regulation of social relationships in spider monkeys. Embraces may serve as signals of benign disposition in contexts that are likely to be associated with tension, such as fusion and access to infants.

Polymorphic Color Vision in Primates: Evolutionary Considerations

Color provides a reliable cue for object detection and identification during various behaviors such as foraging, mate choice, predator avoidance, and navigation. The total number of colors that a visual system can discriminate is largely dependent on the number of different spectral types of cone opsins present in the retina and the spectral separations among them. Thus, opsins provide an excellent model system to study evolutionary interconnections at genetic, phenotypic, and behavioral levels. Primates have evolved a unique ability for three-dimensional color vision (trichromacy) from the two-dimensional color vision (dichromacy) present in the majority of other mammals. This development was accomplished via allelic differentiation (e.g., most New World monkeys) or gene duplication (e.g., Old World primates) of the middle to long wavelength-sensitive (M/LWS, or red–green) opsin gene. However, questions remain regarding the behavioral adaptations of primate trichromacy. Allelic differentiation of the M/LWS opsins results in extensive color vision variability in New World monkeys, where trichromats and dichromats are found in the same breeding population, enabling us to directly compare visual performances among different color vision phenotypes. Thus, New World monkeys can serve as an excellent model to understand and evaluate the adaptive significance of primate trichromacy in a behavioral context. In this chapter, we summarize recent findings on color vision evolution in primates and other vertebrates and introduce our genetic and behavioral study of vision–behavior interrelationships in free-ranging sympatric capuchin and spider monkey populations in Costa Rica.

Spider Monkeys (Ateles geoffroyi yucatenensis) Cope with the Negative Consequences of Hurricanes Through Changes in Diet, Activity Budget, and Fission–Fusion Dynamics

Hurricanes can bring about dramatic changes to ecosystems and adversely affect animals that live in them. We monitored behavioral responses in wild spider monkeys (Ateles geoffroyi yucatanensis) in the aftermath of two hurricanes (Emily and Wilma) that moved through the Yucatan peninsula in 2005. We predicted that the monkeys would shift their diets depending on relative food availability and experience a change in the distribution of activity patterns. Because spider monkeys’ social organization is characterized by a high degree of fission–fusion dynamics, we predicted they would form smaller subgroups subsequent to the hurricanes to mitigate competition over limited food resources. We compared their responses in the immediate aftermath of Hurricane Emily and across the dry seasons before and after both hurricanes, to control for seasonal changes, by examining their activity budgets, foods consumed, and subgrouping dynamics. In the immediate aftermath of Hurricane Emily spider monkeys spent less time moving and more time feeding on leaves than before the hurricane and they were in smaller subgroups. In the dry season after both hurricanes the monkeys spent more time resting and less time moving than before the hurricanes, and leaves replaced fruit as their primary food resource. They fused into larger subgroups less often and had smaller subgroup sizes in the dry season after than before the hurricanes. Thus, the high degree of fission–fusion dynamics of spider monkeys was instrumental in affording the behavioral flexibility critical to cope with the negative post-hurricane consequences.

Site Fidelity in Space Use by Spider Monkeys (Ateles geoffroyi) in the Yucatan Peninsula, Mexico

Animal home ranges may vary little in their size and location in the short term but nevertheless show more variability in the long term. We evaluated the degree of site fidelity of two groups of spider monkeys (Ateles geoffroyi) over a 10- and 13-year period, respectively, in the northeastern Yucatan peninsula, Mexico. We used the Local Convex Hull method to estimate yearly home ranges and core areas (defined as the 60% probability contour) for the two groups. Home ranges varied from 7.7 to 49.6 ha and core areas varied from 3.1 to 9.2 ha. We evaluated the degree of site fidelity by quantifying the number of years in which different areas were used as either home ranges or core areas. Large tracts were used only as home ranges and only for a few years, whereas small areas were used as either core area or home range for the duration of the study. The sum of the yearly core areas coincided partially with the yearly home ranges, indicating that home ranges contain areas used intermittently. Home ranges, and especially core areas, contained a higher proportion of mature forest than the larger study site as a whole. Across years and only in one group, the size of core areas was positively correlated with the proportion of adult males in the group, while the size of home ranges was positively correlated with both the proportion of males and the number of tree species included in the diet. Our findings suggest that spider monkey home ranges are the result of a combination of long-term site fidelity and year-to-year use variation to enable exploration of new resources.

Variability in core areas of spider monkeys (Ateles geoffroyi) in a tropical dry forest in Costa Rica

Core areas are highly used parts of the home range on which the survival of solitary or group-living animals depends. We investigated the home range and core area size and area fidelity of a spider monkey community in a tropical dry forest over a 4-year period. Home ranges overlapped extensively across years, subgroup sizes, and seasons. In contrast, spider monkeys used core areas that varied in size and location across the study years, subgroup sizes, and seasons. These shifts in core areas suggest that the understanding of core areas, and thus the spatial requirements, of a species in a particular habitat may be limited if based on short-term studies. In this respect, our findings emphasize the importance of long-term studies of the spatial ecology of any species in a particular habitat. Our study also shows that the yearly home range basically includes all the core areas from different years, seasons, and subgroup sizes (i.e., the super-core area). This is conceptually important for territorial species, such as spider monkeys, which defend a stable home range as it contains not only the current, but also the future core areas.