subject to large-scale phenomena such as drought
and severe winters may not be protected from such
die-offs by population subdivision. But, where the
crashes are produced by disease epidemics, subdi-
vision may be beneficial, and the creation of linking
corridors, and translocation efforts, may be harmful.
Young noted in his review that most herbivore die-
offs were due to starvation, but carnivore die-offs
were more often attributed to disease. An epidemic
need not eliminate all the individuals in a popula-
tion for it to have a crucial role in causing the even-
tual extinction of a population from a reserve.
Combining these considerations with meta-
population modelling, Hess (1996) argues that
there should theoretically be an optimal degree of
movement within a metapopulation to enable the
movement of propagules of the target species but
not of disease. But much will depend on the nature
of the intervening landscape matrix and the extent
to which it filters target and ‘pest’ species (Ricketts
2001; Ewers and Didham 2005).
The spread of exotic competitors and predators
and of fire into reserves can also be problematic
(Spellerberg and Gaywood 1993; Lockwood and
Moulton 1994). It has been suggested that alien
plant invasion can be reduced by isolating reserves
and surrounding them with simplified cropland
oceans (Brothers and Spingarn 1992). Short and
Turner (1994) found that the decline and extinction
of certain species of native Australian mammals
280 ISLAND THEORY AND CONSERVATION
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Wilderness
Diamond
Figure 10.11A possible spatial and size-frequency distribution of old-growth habitat islands arranged along riparian strips at progressively
greater distances from a current wilderness area in the Willamette National Forest, USA, designed to provide the optimal degree of connectivity
between patches in a system structured by the dendritic pattern of drainage. In this hypothetical system, the most distant islands are generally
larger in order to counter presumed lower dispersal. (Redrawn from Harris 1984, Fig. 10.2.)