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populations. An example of the effect of interference on carnivore population
dynamics is the American marten, a mustelid carnivore in the forests of the USA and
Canada (Fryxell et al. 1999).

Many wildlife species are territorial, meaning that they defend an area of (more
or less) exclusive access from usage by other members of the population. Males,
females, or both sexes may be territorial, depending on the ecological circumstances
that apply. Territories may be defended solely during the breeding season, as in many
birds, or throughout the year, as in many vertebrate carnivores; and they may be
defended by individuals, such as in tigers, or by a pack of individuals, such as in
gray wolves. The multitude of territorial forms mean that many different factors con-
tribute to the adaptive significance of territoriality. Conversely, the consequences of
territoriality at the population and community levels can also vary considerably.
Central to most arguments about the ecological basis of territoriality is the notion
of economic defendability(Brown 1964; Dill 1978; Kodric-Brown and Brown 1978;
Schoener 1983; Stephens and Dunbar 1993; Fryxell and Lundberg 1997). It would
make little sense to try to defend any territory offering trivial benefits or whose costs
are astronomical. Let us assume that the purpose of a territory is to gain access to

THE ECOLOGY OF BEHAVIOR 75

1

0.1
0.1 1 10 100
Predator density

Food intake

1.0

0.8

0.6

0.4

0.2

(^0123)
Log 10 oystercatcher density
Log
10
feeding rate
Site 1
Site 2
Fig. 5.11Changes in
food intake as a
function of predator
(forager) density due to
interference between
predators.
Fig. 5.12Food intake
by oystercatchers at two
sites in the Netherlands
declines as population
density increases. (After
Sutherland 1996.)
5.6.3Territoriality
WECC05 18/08/2005 14:42 Page 75

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