W A G A C: Orld Tlas of Reat Pes and Their Onservation

WORLD ATLAS OF GREAT APES AND THEIR CONSERVATION
intact.115 This pattern is in marked contrast to that instead form their own groups, sometimes taking
seen in western gorillas, among which multimale females with them from the natal group.201
groups are extremely rare.120 Both natal dispersal (leaving the natal group
Group size is variable among eastern gorillas; and transferring to a new group), and secondary
groups ranging from two to 53 individuals have been dispersal (subsequent transfer to yet another
observed.80, 189, 196 In general, median group size is group), occur among female eastern gorillas.
similar for both eastern and western gorillas, Females have also been known to remain and re-
across various habitat types and the different diets produce within their natal group.61, 134, 169, 183 Female
associated with them.103, 120, 200, 203 In the Virungas, mountain gorillas of the Virungas usually transfer
median and mean group size are eight and 11 from their natal group alone, while female eastern
individuals respectively (see Table 8.2).80 At Bwindi, lowland gorillas sometimes transfer with another
a mean group size of about 10 has been reported.92 female and their offspring.201 If a female is pregnant
In the area surrounding Tshivanga in Kahuzi-Biega, or has an infant when she transfers to a new group,
the mean group size of eastern lowland gorillas there is a risk that the new silverback will kill the
(excluding solitary males) is almost 10.69 Mean infant.173 Infanticide has been observed occasionally
group size in the highland sector of Kahuzi-Biega in eastern gorillas,173, 202 although not in every in-
decreased from about 16 per group in 1978, to 11 in stance of transfer with an infant133 (see Box 8.2).
1990, to 10 in 1996.201 Other studies indicate a mean Female transfer could offer a number of
group size of seven animals in Kahuzi-Biega but possible advantages, such as the opportunity of
only three in the adjacent Kasese region.50 higher social rank,35 especially following migration
The sex ratio at birth in both the Virungas and into a small or new group; avoidance of inbreeding;
Kahuzi-Biega is approximately 1:1.174, 201 Upon increased choice of mates; improved reproductive
reaching maturity, most males and females leave success; reduced feeding competition; or improved
the group in which they were born (their natal protection against infanticide.134
group). Males that emigrate usually remain solitary Females may have preferences with regard
until they can attract females and establish their to mates, and this choice may be influenced by
own groups; occasionally, males form all-male male behavior.134 From the male point of view, good
groups. After emigration from the natal group, relationships with females are important to mating
some males spend a large proportion of their time access and breeding success, as a female is free
alone, although in the home range of their natal to leave the group. Although gorilla groups are
group.20, 53 It is very unusual for fully adult males to essentially controlled harems, males cannot there-
migrate into other groups.53, 115, 118, 195 Young males fore afford to make them unduly oppressive ones.
may also stay within the natal group and eventually Both aggressive and affiliative interactions
inherit its leadership.58, 115, 118 Most multimale between males and females have been observed.
groups, but not all, may be the result of males Males have been seen to direct aggressive displays
maturing and remaining in their natal groups,115 and toward females, and females to appease those
are therefore believed to contain several related males (see Box 8.3), although the reasons for these
adult males. Genetic studies confirm that this is displays and their impact on female mate choice
often, but not always, the case.100 remain unclear.134 Males may also vocalize and en-
Whether a young male remains in his natal gage in nonaggressive behaviors toward females,
group or emigrates could be determined by a range possibly to maintain proximity with females.134
of factors including changes in social relationships Females may sometimes intervene in an attempt to
and demographic structure, such as the availa- end aggressive interactions between adult males.132
bility of mating opportunities within the group, the Silverback males in mixed-sex groups do not
death of a parent, or disintegration of the natal interact much with each other but, when they do,
group.28, 115, 118 Males that develop strong affiliative the behavior tends to be more competitive and
(friendly) relationships with the dominant silverback aggressive than affiliative, presumably as a result
while they are infants are more likely to be close of competition over access to mates.118 Affiliative
to the leading male during adolescence, and are interactions are rarely seen,116 but occasional co-
therefore more likely to remain in their natal operation by males within the same group has
group.58 Male eastern lowland gorillas in Kahuzi- been observed, apparently to prevent females from
Biega rarely stay with their putative fathers but leaving the group.131 Relations between silverbacks
136

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