Early field studies of the behaviour of nonhuman primates drew attention to the prominent role of adult males in predation avoidance. DeVore and Washburn (1963; see also Maxim & Buettner-Janusch, 1963) stressed how baboon troops formed a clear travel formation with adult males at the front, flanks and the rear. Males were also reported to interpose themselves in prominent locations between the group and an approaching threat (Hall, 1960).When predators attacked, multiple males were observed engaging in joint counter-attacks (review: Cowlishaw, 1994). The earlier authors interpreted this pattern as an adaptation that maximizes the survival of the group, with expendable males protecting the valuable core of the reproducing females and their offspring. With the rejection of classic group selection (Williams, 1966), this interpretation was no longer tenable, but curiously also meant that interest in the phenomenon waned. Even so, more recent studies also suggest that males appear to be systematically more actively