Agriculture Reference
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of the same host species, unfalteringly supported the ERH. However, evidence
from community studies, which compared the levels of attack between NIS
and native competitors, was much more equivocal. A number of alternate
explanations were proposed to help explain this apparent discrepancy: (1) the
'enemy of my enemy hypothesis' (EEH) occurs when a co-introduced enemy
does more damage to native competitors than to their co-evolved hosts because
of native host naiveté to introduced enemies (see also IFH, below); (2) the
'enemy inversion hypothesis' (EIH) describes a documented situation where a
co-introduced enemy may have actually increased the dispersal ability of its
host by complex interactions between the host, its natural enemy, and a native
species [83-85]; and (3) the 'increased susceptibility hypothesis' (ISH), which
predicts that genetic bottlenecks during the invasion process may render a host
species more susceptible to the effects of enemies, even though this may be
inflicted by a smaller number of species. Colautti et al. [82] also note that bio-
geographical studies may be biased by a failure to consider the confounding
effects of propagule pressure. In other words, many introduced species were
likely founded by a small subset of individuals from a restricted part of their
native range, and therefore a number of natural enemies had no chance of
introduction. Indeed, this seems to be the case for the European starling
Sturnus vulgaris , as the 'apparent' reduction in the number of enemy species,
calculated as a reduction in enemies between its native and introduced ranges,
was a gross overestimate of the 'realized' reduction, calculated after account-
ing for sub-sampling of the source region [85]. It is worth noting the difference
between the ERH, which predicts that invaders succeed because they escape
natural enemies, and the BRH, which predicts that invaders often fail to estab-
lish in the invaded range because of negative effects of enemies native to the
introduced range [55]. Indeed, all invaders likely lose enemies during the inva-
sion process, yet only a small proportion of these have appreciable ecological
or economic impacts.
Invasional facilitation hypothesis (IFH)
Contrary to the BRH and the ERH, the IFH predicts an increasing rate of inva-
sions through time [86]. The term 'invasional meltdown' was introduced by
Dan Simberloff and Betsy Von Holle at the University of Tennessee to describe
a process whereby facilitative interactions among NIS may increase invasion
rates [86]. Facilitative interactions among invaders have been demonstrated in
many systems [63, 87-89]. For example, invasion by the zebra mussel
Dreissena polymorpha in the Laurentian Great Lakes has drastically altered
the ecology of the lakes, including an increase in the diversity of macroinver-
tebrates, which use zebra mussel shells as refuge from predators [90]. The
invasion and subsequent proliferation of D. polymorpha in 1988 was closely
followed by invasions of three co-evolved species, which are also found in
high abundance: in 1994 by Echinogammarus ischnus , a macroinvertebrate
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