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Nitrogen Enrichment and Plant Invasions 169<br />

10.2.5 Nitrogen Fixation Suppressed by Invasion<br />

Nitrogen fixation could also be negatively affected by plant invasions. For the<br />

invasion of African grasses in Hawaii, Ley and D’Antonio (1998) demonstrated<br />

that these species eliminate the fire-sensitive native tree Metrosideros<br />

polymorpha by promoting fire events. Non-symbiotic N 2 -fixing bacteria are<br />

active on the litter of M. polymorpha, but not on that of the invasive grasses.<br />

Hence, elimination of these native species led to reduced nitrogen contents in<br />

invaded soils. In addition, invasive species that do not themselves fix N 2 may<br />

also indirectly affect the rate of fixation in co-occurring N 2 -fixing species,<br />

possibly through an allelopathic mechanism (Ehrenfeld 2003). For example,<br />

in glasshouse and pasture studies in New Zealand, Wardle et al. (1994) found<br />

evidence that decomposing leaves of the invasive thistle Carduus nutans<br />

inhibit nitrogen fixation by the introduced Trifolium repens. C. nutans has<br />

invaded many areas of western North America. Therefore, it might be possible<br />

that this thistle also adversely affects growth and nitrogen fixation of native<br />

legumes.<br />

10.3 Nitrogen Deposition and Exotic Invasions<br />

10.3.1 N Deposition and Eutrophication in Natural Ecosystems<br />

The input of nitrogen compounds into terrestrial ecosystems is a major component<br />

of global environmental change, and can have substantial effects on<br />

ecological processes and the biogeochemistry of these systems (Vitousek et<br />

al. 1997). Current estimates of airborne, mineral nitrogen deposition show<br />

that large portions of North America, Europe, and Asia have rates<br />

>75 kg N ha –1 year –1 , and some regions receive >100 kg N ha –1 year –1 (Galloway<br />

et al. 2004), exceeding the natural background rate by two orders of magnitude.<br />

Anthropogenic emissions of nitrogen compounds (NH x and NO x ) from<br />

fossil fuel combustion, fertilizer use, and animal husbandry are the underlying<br />

cause for this elevated deposition of biologically active N. Beside the relatively<br />

well-documented effects on biogeochemical cycles, N deposition can<br />

also have severe economic impacts, such as increased costs for the management<br />

or purification of drinking water (Wamelink et al. 2005).<br />

The most direct ecological effect is a subtle, but continuous increase in<br />

nitrogen pools and fluxes, and hence in nitrogen availability for plant growth<br />

and productivity in N-limited ecosystems (e.g., Olde Venterink et al. 2002).<br />

The eutrophying influence of N deposition may be associated with an acidification<br />

of soils, mineral imbalances in plant nutrition resulting in reduced<br />

plant vitality and increased sensitivity against stress, and enhanced nitrate

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