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nificantly in their average monthly rate of presence, whereas age classes did;
juvenile cats had the highest average monthly rate of presence, and adult males
the lowest. Furthermore, each sex-age class displayed a favorite nearest-neigh-
bor distance in terms of its tendency to maintain a given level of proximity to
other cats (figure 10.6).
At the large colony, when kittens were within 1 m of an adult female, this
was their mother on 71.1 percent and 58.3 percent of occasions for male and
female kittens, respectively, whereas adult males tended to be within 1 m of
male juveniles most frequently and of male kittens least frequently. Despite
such generalizations, there were also significant differences in the pattern of
proximity between the colonies. For example, at the large colony, adult males
were significantly more frequently within 1 m of adult females, but at the
medium-sized colony no such tendency emerged. A further corollary of spatial
structure was that certain females occupied spatially peripheral positions,
whereas others lived centrally, and the latter had significantly higher reproduc-
tive success (figure 10.7).
Taking into account indices of association, it is possible to explore the fre-
quency, rate, and quality at which cats interact. Take three cats, cat A spending
twice as much time in the company of cat B as of cat C (figure 10.8). If A rubs
on B and C 40 and 8 times respectively, and attacks them 10 and 2 times
respectively, then A rubs on and attacks B at only 2.5 times the rate at which it
rubs on or attacks C, although A rubs and attacks B five times as often as C.
The quality of A's relationship with B and C is the same in that 80 percent of
its interactions with both are rubbing and 20 percent are attacking.
In reality, at Kerby and Macdonald's (1988) large colony, individual central
males averaged 602 (±254, SE) initiations relative to 502 (±142) for peripheral
males, but the latter were present so seldom that their hourly rate of interac-
tion when present averaged 6.9 (±1.5), in comparison to the central males at
2.3 (±0.7). The situation was opposite for females: Peripheral females inter-
acted at low frequency, and when their presence was taken into account, they
interacted at an even lower rate. Furthermore, in the context of grooming,
Macdonald et al. (1987) showed that whereas the flow of licking differed sig-
nificantly between pairs of interacting cats, it was invariably symmetric. In
marked contrast, the flow of rubbing, though also differing between dyads,
correlated with aggression but not grooming and was generally highly asym-
metric. They concluded that in the society of female cats, rubbing as a measure
of status flowed centripetally toward central dominant individuals; grooming,
which was exchanged reciprocally, was a measure of social (and often genetic)
alliances, but unrelated to status (figure 10.9).
