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

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46 David A. Wardle <strong>and</strong> Duane A. Peltzer<br />

of terrestrial ecosystems (Vitousek et al. 1987, Lawton 1994, Chapin et al.<br />

1997), <strong>and</strong> feedbacks between the aboveground <strong>and</strong> belowground components<br />

are arguably among the most important of these biotic drivers (Wardle<br />

2002).<br />

Development of a thorough underst<strong>and</strong>ing of either primary or secondary<br />

ecological succession, <strong>and</strong> ecosystem restoration, requires specific consideration<br />

of both the aboveground <strong>and</strong> belowground subsystems, as well as the<br />

nature of feedbacks between them. Although primary successional development<br />

on newly created surfaces, or secondary succession following significant<br />

disturbance events, has usually been studied only with specific reference to the<br />

plant community <strong>and</strong> the availability of major soil nutrients (Bradshaw <strong>and</strong><br />

Chadwick 1980), the reality is that the aboveground <strong>and</strong> belowground communities<br />

develop in close concert with each other over successional time. Further,<br />

plant species replacement (such as occurs both during primary <strong>and</strong> secondary<br />

succession) has major effects on soil communities <strong>and</strong> the ecological processes<br />

that they control (Wardle et al. 1999, Porazinska et al. 2003, Belnap et al. 2005).<br />

Changes in soil communities in turn influence the direction <strong>and</strong> speed of both<br />

primary <strong>and</strong> secondary vegetation succession as well as ecosystem productivity<br />

(van der Putten et al. 1993, De Deyn et al. 2003). Knowledge about feedbacks<br />

between aboveground <strong>and</strong> belowground biota is also crucial to developing a better<br />

underst<strong>and</strong>ing of the principles of ecosystem restoration, because facilitating<br />

the recovery of ecosystems requires recognition of the role of these feedbacks<br />

in driving community- <strong>and</strong> ecosystem-level properties <strong>and</strong> processes.<br />

In this chapter, we will start by discussing vegetation succession <strong>and</strong> ecosystem<br />

development in the context of a combined aboveground–belowground approach.<br />

We will then present three case studies in which principles of succession<br />

studied through a combined aboveground–belowground approach are relevant<br />

to the goals of ecosystem restoration: (1) the consequences of human-induced<br />

changes in densities of browsing mammals, with particular reference to deer in<br />

New Zeal<strong>and</strong> rain forests; (2) the ecological impacts of fire in the long-term,<br />

with particular reference to boreal forests of northern Sweden; <strong>and</strong> (3) the belowground<br />

impacts of invasive, nonnative plant species <strong>and</strong> their feedbacks<br />

aboveground. These examples will be used to emphasize the importance of<br />

combined aboveground–belowground approaches to underst<strong>and</strong>ing vegetation<br />

succession, the ecological role of disturbance, <strong>and</strong> the restoration of ecological<br />

interactions <strong>and</strong> processes.<br />

3.2 <strong>Succession</strong>al Development <strong>and</strong> Aboveground–Belowground<br />

Linkages<br />

An important component of ecosystem development <strong>and</strong> succession is the<br />

changes that occur in the attributes of the dominant vegetation. As primary succession<br />

proceeds, there is a general shift in dominant plant species from those<br />

that are of small stature, often herbaceous, short-lived, have a high reproductive<br />

output, <strong>and</strong> produce litter with high quality with those that are increasingly<br />

larger, woody, long lived, more conservative in retaining nutrients, <strong>and</strong> produce<br />

foliage <strong>and</strong> litter of poorer quality (Grime 1979, Walker <strong>and</strong> Chapin 1987,<br />

Wardle 2002). Similar trends also occur in secondary succession, even though<br />

the starting point may be later in ecosystem development because of legacy

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