Spatial cohesion of adult male chimpanzees (Pan troglodytes verus) in Taï National Park, Côte d'Ivoire.

Group living animals can exhibit fission-fusion behavior whereby individuals temporarily separate to reduce the costs of living in large groups. Primates living in groups with fission-fusion dynamics face numerous challenges in maintaining spatial cohesion, especially in environments with limited visibility. Here we investigated the spatial cohesion of adult male chimpanzees (Pan troglodytes verus) living in Taï National Park, Côte d'Ivoire, to better understand the mechanisms by which individuals maintain group cohesion during fission-fusion events. Over a 3-year period, we simultaneously tracked the movements of 2-4 males for 4-12 hr on up to 12 consecutive days using handheld GPS devices that recorded locations at one-minute intervals. Analyses of the male's inter-individual distance (IID) showed that the maximum, median, and mean IID values across all observations were 7.2 km, 73 m, and 483 m, respectively. These males (a) had maximum daily IID values below the limits of auditory communication (<1 km) for 63% of the observation time, (b) remained out of visual range (≥100 m) for 46% of observation time, and (c) remained within auditory range for 70% of the time when they were in different parties. We compared the observed distribution of IIDs with a random distribution obtained from permutations of the individuals' travel paths using Kolmogorov-Smirnov tests. Observation IID values were significantly smaller than those generated by the permutation procedure. We conclude that these male chimpanzees actively maintain cohesion when out of sight, and that auditory communication is one likely mechanism by which they do so. We discuss mechanisms by which chimpanzees may maintain the level of cohesion observed. This study provides a first analysis of spatial group cohesion over large distances in forest chimpanzees using high-resolution tracking, and illustrates the utility of such data for quantifying socio-ecological processes in primate ecology.