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Nitrogen Enrichment and Plant Invasions 173<br />

Fig. 10.2 Correlation<br />

between total N wet<br />

deposition as nitrate<br />

and ammonium and the<br />

proportion of nonnative<br />

plant species in forest<br />

ecoregions of the USA.<br />

Data taken from the US<br />

National Atmospheric<br />

Deposition Program<br />

(NADP 2006) and from<br />

the World Resources<br />

Institute (WRI 2000).<br />

Wet deposition represents<br />

means±s.e. of<br />

annual averages of 223<br />

monitoring sites located<br />

in forest systems<br />

non-native plant species (%)<br />

> 30<br />

(n=14)<br />

21-30<br />

(n=49)<br />

11-20<br />

(n=80)<br />

5-10<br />

(n=77)<br />

< 5<br />

(n=3)<br />

0 1 2 3 4 5<br />

N deposition wet<br />

(kg N ha -1 yr -1 )<br />

10.3.3.2 Observational Studies<br />

More reliable data on the relationship between N deposition and invader success<br />

are available only on a local or plot scale,i.e.,from a few square kilometers<br />

to several square meters. Theory predicts that a plant community should<br />

become more invasible if there is an increase in the amount of unused<br />

resources (Davis et al. 2000), i.e., if the use of resources by resident species<br />

declines,or if resource supply increases at a rate faster than the resident species<br />

can make use of. The latter may occur in the case of eutrophication, e.g., by N<br />

deposition.There are indeed several reports of promoted invasion on more fertile<br />

soils. For instance, in New Zealand’s Nothofagus forests, areas invaded by<br />

Hieracium species had higher N availabilities as well as higher P, Ca, and Mg<br />

concentrations than those recorded in uninvaded areas (Wiser et al. 1998).<br />

Likewise, in southern Utah, exotic plant invasions were highly correlated to<br />

soils high in C, N, and P (Bashkin et al. 2003).Although variations in soil fertility<br />

were of natural origin in these studies, the authors concluded that because<br />

habitats containing fertile soils appeared more vulnerable to exotic invasions<br />

than those with less-fertile soils, shifts in soil conditions induced by N deposition<br />

could shift the balance for native and exotic species locally. Similarly, the<br />

clonal-growing species Arundo donax,which is known to be very responsive to<br />

N enrichment, invades riparian areas in the USA preferentially at sites of high<br />

nitrogen availability (Decruyenaere and Holt 2005). Its higher success under<br />

such conditions is attributable to year-round activity, greater overall ramet<br />

flux, and higher foraging. Although eutrophication of riparian ecosystems is<br />

related more to nutrient enrichment of surface waters than to atmospheric N

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