changes: they were higher during food availability and males marked twice as often as females at this time (x2 =5.1, p <0.03, df =1, Figure 4).

Agonism Within-Group

The only form of within-group aggression recorded was cuffing and was rarely observed (n = 20, rate 0.01/hr DJO, n = 19, rate 0.007/hr SRT). Only two of these exchanges occurred during feeding in February when food was scarce. The majority of cuffs (n = 20) were observed between parents and unweaned offspring. In each of these cases, the adult female carrying the infant approached the adult male, cuffed him, and turned to rub the infant off her back or belly and onto the male's back. The male would then either carry the infant or rub the infant off his back onto a branch so that the infant was forced to travel on its own. The remaining incidents (n = 17) occurred between an adult and yearling offspring between August (just prior to the birth season) and October (when infants were approximately 1 month old).

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