Geology Reference
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with 24, 17, and 21 observed associaions with
groups of . mitis, respecively. In Kibale, . b.
tephrosceles (Struhsaker, 1975) and in Mwanihana,
. b. gordonorum (Wasser, Chapter 13) also asso-
ciated with groups of . mitis. On no occasion in
the Zanzibar study was a solitary red colobus
observed to associate with . mitis. On the other
hand, solitary red colobus in Kibale (Struhsaker,
1975), Gambia (Starin, 1981) and Mwanihana
(Wasser, Chapter 13) associated with other
primate species. In Kibale and Mwanihana more
than two primate species were found together
simultaneously on some occasions.
In both Kibale and Zanzibar red colobus
associations were mostly initiated and disbanded
by the other primates. However, in Gambia, red
colobus approached vervet monkeys (Cercopithecus
aethiops) in most cases (Starin, 1981). The dura-
ion of the associaions varied but it was very short
among the Zanzibar red colobus and . mitis, 1-
15 min, compared with 130.4 and 121.8 min
between the Kibale red colobus and . ascanius
and . mitis, respecively (Struhsaker, 1975).
Associations involving Gambia red colobus
generally lasted for several days up to a year
(Starin,
1981).
In the lringa red colobus some
associations are of long duraion; S. K. Wasser
personal communicaions) reports repeated
observaions of the same polyspecific grouping
over a period of several days.
Resting was the most common acivity recorded
during the associaion ofthe Zanzibar red colobus
and . mitis, comprising 56.6% of the records.
Feeding accounted for 21.9%, allogrooming
5.5%, and playing 3.8% of the records. In the
Kibale red colobus the most common social
acivity was grooming followed by playing (Struh-
saker, 1975). In both Zanzibar and Kibale red
colobus, play mostly involved young animals.
lnterspecific aggression between the red colobus
and the other primate species was not observed in
Zanzibar, while there was occasional aggression in
Kibale (Struhsaker, 1975) and Gambia (Starin,
1981).
Discussion
The Zanzibar red colobus food habits, group
composiion, intraspecific relaions and primate
polyspecific associaions show great similariies to
those of other red colobus that have been studied.
There are very few behavioural aspects in which
any paricular subspecies of red colobus stands
unique. In the majority of cases a behaviour is
common in two or more subspecies irrespecive of
their geographical proimity. This secion discus-
ses the similarities and differences among the red
colobus in the above respects, and attempts to
show why the differences and similariies eist.
The red colobus populaions so far studied are
mainly folivores. This is likely to be related to
feeding adaptaions they have for uilising leaves,
namely, by possession of a large chambered
stomach analogous to that of ruminants (Moir,
1965; Bauchop & Martucci, 1968; Kay, Hoppe &
Maloiy, 1976). They prefer young leaves, with
high nurient and low antifeedant content. (An
antifeedant is defined as a plant chemical com-
pound potentially harmful to herbivores.) Young
leaves commonly contain more protein and less
fibre,
(Dougall & Drysdale, 1964;
Struhsaker,
1975;
Hladik,
1977;
Waterman & Choo,
1981;
Mturi, 1991), and also have less toin than mature
leaves (Waterman & Choo, 1981; Mturi, 1991).
In ruminants, foods containing a high proporion
of fibre take longer to ferment Ganis, 1976). Thus
it may be important for red colobus to utilise
young leaves, which are more digesible, especi-
ally as colobines are near the bottom of the size
range for which fore-stomach fermentation is
viable (Parra, 1978).
All the red colobus monkeys studied are very
selective in their fe eding. This may be aimed at
maimising the intake of nutrients while minimis-
ing the intake of deleterious compounds (Free-
land & Janzen, 1974; Westoby, 1974; Pulliam,
1975; Glander, 1982; Waterman, 1984). Selec-
tivity for paricular plant parts is obseved in red
colobus; for example, mature leaf peioles, which
contain less fibre, are preferred to mature leaf
blades (Waterman & Choo, 1981; Baranga, 1982;
Mturi, 1991). The peioles also have higher
digesibility and ermentable carbohydrates, and
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