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88<br />

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

(1) differential parasitism in populations of a single species, both in its native<br />

and introduced region (within-species, cross-regional comparison), (2) native<br />

versus introduced species in the same region (between-species comparison of<br />

ecologically analogous competitors), (3) introduced species that are not invasive<br />

with introduced species that are invasive (between introduced species),<br />

and (4) population growth rates and enemy abundance among populations<br />

within a range (within species, within region) will help fully evaluate enemy<br />

release and the role of enemies in biological invasions. When used in combination,<br />

such comparisons will clarify the extent to which natural enemies<br />

keep their host populations in check, and the consequences of release from<br />

these natural enemies on population growth. To our knowledge, this joint<br />

approach has not yet been employed.<br />

6.3.2 Refining the Evolution of Increased Competitive Ability Hypothesis<br />

Evidence for adaptive evolution in invaders supporting the EICA hypothesis,<br />

particularly local adaptation to the new environment, is growing (e.g., Leger<br />

and Rice 2003; Lee et al. 2003; Blair and Wolfe 2004; Maron et al. 2004), but is<br />

not found in all cases (e.g., Willis et al. 1999; van Kleunen and Schmid 2003).<br />

Similarly to cases dealing with enemy release, much of the conflict in the data<br />

may be due to the specific comparisons that are made. For example, most of<br />

the data come from common garden experiments with plants, largely comprised<br />

of specimens that simply happened to be available for study, without<br />

particular knowledge of the origin of an invasion (but see Maron et al. 2004).<br />

Often, these are not comprehensive samples taken from across the native and<br />

introduced ranges (but see Blair and Wolfe 2004). If common gardens include<br />

inappropriate comparisons, then the data may not reflect the role of evolution<br />

in invasions.<br />

6.4 Recent Syntheses and Synergies Between Hypotheses<br />

Efforts have been made to synthesize across hypotheses. Shea and Chesson<br />

(2002) view biological invasions from the perspective of community ecology<br />

theory. They discuss invasions in terms of niche opportunities that are determined<br />

largely by resources, natural enemies, and the physical environment.<br />

Top-down population regulation is exerted by both specialist and generalist<br />

enemies, and for invaders, specialists may be relatively rare in the new range.<br />

The availability of resources that are present in a habitat is governed by other<br />

members of the community using those resources. Thus, Shea and Chesson<br />

(2002) bring together aspects of the enemy release, biotic resistance, and<br />

empty niche hypotheses.

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