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Linking Restoration and Ecological Succession (Springer ... - Inecol

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phases <strong>and</strong> a domination by late-successional species; e.g., Elymus athericus<br />

(Bos et al. 2002) or Phragmites australis (Esselink et al. 2000). The overall<br />

result is a loss in biodiversity. <strong>Succession</strong> in salt marshes can be manipulated<br />

back to earlier stages by reopening existing embankments <strong>and</strong> flooding.<br />

In a survey of 70 restored flooded sites in northwestern Europe, the percentage<br />

of target species was as high as 70% in the best examples (Wolters et al.<br />

2005).<br />

6.5 The Role of Aliens<br />

6.5.1 <strong>Succession</strong>al Pattern: Repairing Function of Native Vegetation<br />

Recent attempts to bring the science of invasive species <strong>and</strong> succession together<br />

have proved profitable (Davis et al. 2000, 2005; see Chapter 3). Alien<br />

species (defined as those whose presence in a given area is due to intentional<br />

or unintentional human involvement, or which have arrived without the help<br />

of people from an area in which they are alien; see Pyˇsek et al. 2004b) are<br />

increasingly common in successional seres. For example, in 55 successional<br />

studies in central Europe <strong>and</strong> North America, 25% of the species (range 2–<br />

81%) were aliens. Aliens were most prevalent in ruderal habitats <strong>and</strong> old fields,<br />

<strong>and</strong> their representation declined during the successional process. The rate of<br />

this decrease was context-dependent: industrial habitats had a greater proportion<br />

of aliens at the start, but a faster decline with time than habitats associated<br />

with agricultural l<strong>and</strong>scapes. Alien species contributed more in terms of species<br />

number than cover reflecting that many of them are rare casuals (Richardson<br />

et al. 2000, Pyˇsek et al. 2004b). Those aliens classified as neophytes (species<br />

introduced after 1500AD) were most likely to become dominants (Pyˇsek et al.<br />

2004a).<br />

It is not known how much the pattern of decrease of alien species during<br />

succession is determined by the exposure of various successional stages to<br />

different propagule pressures of alien species. It is predicted that colonization<br />

by diaspores will be greater at the beginning of succession (Rejmánek 1989).<br />

Experimental studies on the invasibility of successional stages are rare, but<br />

Bastl et al. (1997) found that early, but not initial, successional stages were<br />

most prone to plant invasions. The establishment of aliens in the initial stages<br />

of succession was probably restricted by adverse abiotic conditions, whereas in<br />

later successional stages, intensive competition from resident species appeared<br />

more important. This seems to be a common pattern (Rejmánek 1989) <strong>and</strong><br />

should be considered in restoration practice. It has been suggested that the<br />

maximum cover <strong>and</strong> proportion of aliens are found in the initial stages of mesic<br />

succession (Rejmánek 1989). These results suggest support for the successional<br />

repairing function of native vegetation (Rejmánek 1989) indicating that during<br />

spontaneous succession alien species should disappear in time (Pyˇsek et al.<br />

2004a).<br />

6.5.2 Manipulating <strong>Succession</strong> in Invaded Sites<br />

The unassisted recovery of native vegetation during succession is of little practical<br />

use when dealing with those large-scale invasions where immediate action<br />

is needed. In such cases, the dominant alien species must be at least contained<br />

Chapter 6 Manipulation of <strong>Succession</strong> 137

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