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90<br />

R.A. Hufbauer and M.E. Torchin<br />

introduced ranges, examining how their population dynamics are influenced<br />

by several types of interspecific interactions and the abiotic environment. The<br />

biotic interactions they examined comprise predation, parasitism, mutualism,<br />

and competition, making this the first theoretical integration of these key<br />

groups.<br />

Furthermore, Mitchell et al. (2006) highlight that phylogenetic relationships<br />

between an invader and the members of the invaded community may<br />

play a critical role in the outcome of an introduction. Introduced species<br />

invading communities with close relatives should be more susceptible to enemies<br />

in that community, and thus accumulate a broader suite of enemies compared<br />

to invasions of communities in which relatives are absent (Strong et al.<br />

1984; Mack 1996). However, they may also be more likely to gain mutualists<br />

from their relatives. The comparative importance of enemies and mutualists,<br />

and their relative differences in host specificity will determine whether invading<br />

a community containing close relatives is a disadvantage for invaders. The<br />

relatedness of members of the community may also be an indicator of suitability<br />

of the abiotic environment, when related species have similar environmental<br />

requirements.<br />

Additional connections and synergies between hypotheses are possible,<br />

and for the field to advance, those links should be clarified and formalized in<br />

a predictive framework. We offer an initial attempt at such a framework<br />

(Table 6.2), focusing on interactions among enemy release, biotic resistance,<br />

and evolutionary change (including both evolution of increased competitive<br />

ability, and hybridization). By formalizing the connections among hypotheses<br />

to explain invasion, we can generate specific, testable predictions. These predictions<br />

can guide research efforts, and resultant data can feed back into the<br />

predictive framework.<br />

For example, we suggest that genetic variation and changes associated with<br />

introductions may interact directly with enemy release and biotic resistance<br />

in at least two key ways. First, with a severe bottleneck in population size upon<br />

introduction, a species will lose genetic variation, but is also more likely to<br />

lose natural enemies (Torchin et al. 2002). Thus, adaptive evolution may be<br />

most limited by lack of genetic variation when enemy release is likely to be<br />

greatest, setting up a useful comparison between two potentially opposing<br />

pathways for invasion success (Drake 2003; Table 6.2, hypothesis 4a). Additionally,<br />

reductions in variation will define the context within which new<br />

interactions with enemies develop, and may affect the ability of the invader to<br />

defend against enemies.<br />

A second way that genetic variation may interact with enemy release and<br />

biotic resistance is through changes associated with hybridization. Often, it is<br />

only in novel environments that hybrid and backcross offspring are fitter than<br />

their parents. One key novel aspect of a new range may be the lack of enemies.<br />

Thus, ways in which hybridization alters interactions with natural enemies<br />

could be particularly important. One mechanism may be simply that

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