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170<br />

M. Scherer-Lorenzen, H. Olde Venterink, and H. Buschmann<br />

leaching into groundwater (Schulze 2000; Nadelhoffer 2001). Ecosystems that<br />

are characterized by low natural levels of nitrogen availability are most vulnerable<br />

– for example, ombrothrophic bogs, heathlands, temperate and boreal<br />

forests, or nutrient-poor grasslands. Critical loads for acidifying and eutrophying<br />

nitrogen for these ecosystems lie in the range of 5–20 kg N ha –1 year –1 ,<br />

currently exceeded in many parts of the world (EEA 2005). These alterations<br />

of growing conditions also have direct consequences for the competitive success<br />

of species, associated with often dramatic changes in community composition<br />

and decrease of species richness, especially at N-limited oligo- and<br />

mesotrophic sites such as in northern temperate forests (Bobbink et al. 1998;<br />

Sala et al. 2000): slow-growing plants adapted to nutrient-poor soils are outcompeted<br />

by faster-growing species originating from nutrient-rich sites.<br />

Responsiveness of species to N deposition is thus highly related to plant traits<br />

typical for high-nutrient environments.<br />

Since invasive species are often associated with a particular suite of traits<br />

that are characteristic for plants of nutrient-rich sites (see Sect. 10.1), one<br />

might assume that N deposition could also influence the abundance patterns<br />

of native vs. invasive plant species. In addition, the invasibility of ecosystems<br />

is most often also related to the degree of resource availability (Davis et al.<br />

2000), and invasive species generally occur more frequently at nutrient-rich<br />

sites, as shown for the floras of Germany (Scherer-Lorenzen et al. 2000), and<br />

the Czech Republic (Pyšek et al. 1995). One could therefore hypothesize that<br />

invasive species are more successful, and therefore more abundant in areas of<br />

high atmospheric N deposition than in areas of low deposition.<br />

10.3.2 A Short Note on Mechanisms<br />

As Scherer-Lorenzen et al. (2000) have discussed, the mechanism of competitive<br />

exclusion of species adapted to low nutrient levels by faster-growing,<br />

more nitrophilic species is independent of whether the invading species is<br />

either alien or native. There are numerous examples of successful native<br />

species invading species-rich communities under high N deposition (Bobbink<br />

et al. 1998). Particularly well documented is the case of ombrotrophic<br />

bogs, an ecosystem type among the most sensitive to N enrichment because<br />

these bogs receive most of their nutrients from the atmosphere (e.g.,<br />

Tomassen et al. 2004). Other examples include community changes in wet<br />

heathlands (e.g.,Aerts and Berendse 1988), or after acidic deposition-induced<br />

forest dieback. In the latter case, native grasses responsive to nitrogen invade<br />

the understorey of the forests, replacing species adapted to less fertile soils.<br />

Examples from mountain spruce forests in Europe include the replacement of<br />

the dwarf shrubs Vaccinium myrtillus and Calluna vulgaris or the grass<br />

Deschampsia flexuosa by the grass Calamagrostis villosa (Scherer-Lorenzen<br />

et al. 2000). In all cases, combinations of specific traits, such as high relative

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