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the juveniles leave voluntarily, their behavior being innately determined by their
genes (e.g. in Belding’s ground squirrels (Spermophilus beldingi); Holekamp 1986);
or that adults forcibly exclude juveniles. Saturationdispersal is seen in many large
mammals (Sinclair 1992). In this case dispersal occurs when a population reaches a
threshold density determined by food limitation. Dispersal then is density dependent
(see Chapter 8 for an explanation of this mechanism) so that population density remains
the same on the initial area. Examples of this have been described for Himalayan
tahr (Hemitragus jemlahicus) as they spread through the Southern Alps of New
Zealand (Parkes and Tustin 1985), and for wood bison (Bison bison) as their popu-
lation increased through their former range in the boreal forest of Canada (Larter
et al. 2000) (see Section 7.6 for modeling range expansion).
The likelihood of dispersal differs markedly between individuals of a population.
Figure 7.1 shows a sample of distances dispersed by juvenile kangaroo rats
(Dipodomys spectabilis) ( Jones 1987), a solitary, nocturnal, grain-eating desert
rodent. Females averaged 29 m and males 66 m, but the majority of individuals did
not disperse at all. Jones (1987) reported that adults of this species do not disperse
much: 70% of adult males and 61% of adult females remained in one mound for the
rest of their lives. Juvenile females of red deer (Cervus elaphus) seldom disperse but
adopt home ranges that overlap those of their mothers. In contrast, males leave the
natal home range between the ages of 2 and 3 years, mostly joining stag groups in
the vicinity (Clutton-Brock et al. 1982).
Patterns of dispersal are related to the type of mating system (Greenwood 1980,
1983; Dobson 1982; Greenwood and Harvey 1982). Thus, in mammals, females are
concerned with obtaining resources while males compete for mates. In general, males
disperse in promiscuous and polygynous species because they are more likely to find
new mates by doing so, while females are philopatric (i.e. remain at their birth site)
because they are more likely to find food in areas they know well. Both sexes dis-
perse in monogamous species. Amongst higher vertebrates, one sex is more prone
to dispersal than the other. Thus, in mammals males are the dispersers whereas in
birds it is the females which disperse, although there are exceptions for both groups.
For example, in mammals females are the dispersers in wild dogs (Lycaon pictus) and
zebra (Equus burchelli). In fishers (Martes pennanti) and wolves, both sexes disperse
equally (Arthur et al. 1993; Boyd and Pletscher 1999).
The causes of dispersal fall into three broad categories: competition for mates, avoid-
ance of inbreeding, and competition for resources ( Johnson and Gaines 1990). In

DISPERSAL, DISPERSION, AND DISTRIBUTION 91

50

40

30

20

10

0
0 1 50 100 150 200 250 300
Dispersal distance (m)

Number of individuals

Female (n = 70) Male (n = 72)
50

40

30

20

10

0
0 1 50 100 150 200 250 300

Fig. 7.1The frequency
distributions of
distances dispersed by
juvenile rat kangaroos.
(Data from Jones 1987.)

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