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80<br />

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

species can have significant negative ecological and environmental effects<br />

while having positive economic effects (e.g., cows). Third, it is important to<br />

have a baseline against which changes associated with invasion can be judged.<br />

We argue that comparing the ecology of an introduced species to that of populations<br />

in its native range will set the most relevant baseline by which to<br />

measure changes in ecology resulting from translocation. If we first understand<br />

the causes of demographic variation among populations within the<br />

native range, then we can compare introduced populations to this gradient<br />

(Torchin et al. 2001).<br />

For some species, population demographics may be similar between the<br />

native and novel ranges, and mechanisms of their success may be no different.<br />

Such species may be quite benign when introduced, or if they cause economic<br />

problems in their native range, then often they will in their introduced range<br />

as well (e.g., insect pests such as Western corn root worm, Diabrotica virgifera,<br />

which is a native pest in North America, and an invasive pest in Europe;<br />

Chap. 2). If a species is more abundant, dense, or widespread in the novel<br />

range than in the native range, then – by definition – something has fundamentally<br />

changed in its ecology or perhaps evolution. Because invasiveness is<br />

a combined function of the invaded community and the invader, the changes<br />

leading to greater success in the new range can be extrinsic changes in the<br />

environment that favor the invading species, or they can be intrinsic to the<br />

invading species. We refer to species that experience significant positive<br />

demographic changes that contribute to invasion success as strong invaders,<br />

and those with no change or significant negative changes as weak invaders.As<br />

Hierro et al. (2005) point out, there are remarkably few comparative data<br />

between the native and novel ranges of species documenting whether the<br />

demography of invaders changes (but see Torchin et al. 2001).<br />

Introduced species that are weak invaders can be important economically<br />

and environmentally, and also give rise to interesting and urgent ecological<br />

questions. However, weak invaders are less useful in helping us address questions<br />

regarding fundamental ecological and evolutionary changes that appear<br />

to underlie the most damaging invasions (e.g., tamarisk and zebra mussel in<br />

North America). To further an understanding of the causes of biological invasion,<br />

it is vital to know whether or not most species considered to be “invasive”<br />

are strong invaders that have experienced dramatic demographic<br />

changes relative to their native range.<br />

Herein, we propose a metric to quantify the continuum from weak<br />

invaders to strong invaders. Response ratios are used to compare the means<br />

of experimental treatments (X E ) and controls (X C ), where R=X E /X C (Hedges<br />

et al. 1999). If R>1, then the experimental treatment is larger than the control,<br />

and if R

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