A PRELIMINARY-STUDY OF VOCAL COMMUNICATION IN WILD LONG-TAILED MACAQUES (MACACA-FASCICULARIS) .2. POTENTIAL OF CALLS TO REGULATE INTRAGROUP SPACING

This study is a preliminary assessment of the potential of long-tailed macaque (Macaca fascicularis) calls to operate in systems of within-group spacing. Covariation in the rate of occurrence of calls with party spread, size, and activity among wild individuals of one group suggested that four classes of calls may function in intragroup spacing. Two of them are "clear" calls of long duration and pronounced frequency modulation. Calling rate increased with party spread for low- and high-frequency variants of these calls during resting and feeding, respectively, suggesting possible utility in maintenance of spatial relations over moderately long distances. A third "harsh" call was negatively correlated with party spread during foraging and may thus function to increase dispersion among foraging individuals. Another harsh call with a tonal onset was unique among all calls in the vocal repertoire in being more frequently performed by lone, isolated individuals than by macaques accompanied by others, suggesting a possible function in reestablishing contact that has been severed. The functional significance of these calls with respect to their acoustic structures is discussed. Macaques that use calls to regulate intragroup spacing can control communication distance and direction by their choice of acoustically different vocalizations. This choice may be affected not only by varying environmental constraints on sound transmission, but also by social and ecological factors such as intragroup competition.