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

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154 Richard J. Hobbs, Anke Jentsch, <strong>and</strong> Vicky M. Temperton<br />

the onset of restoration measures, to help assess what the limiting factors for<br />

restoration are, <strong>and</strong> how one could alter the filter mesh to allow certain species to<br />

establish.<br />

This is where the issue of disturbance comes in. Disturbances modify community<br />

assembly in two ways. Firstly, disturbances change environmental filters<br />

such as nutrient availability, <strong>and</strong> secondly, they act on plant traits as filters of<br />

species (biotic) assembly in their own right (White <strong>and</strong> Jentsch 2004). Thus,<br />

disturbance can be not only the cause of degradation, but at appropriate scales<br />

<strong>and</strong> magnitude can also be a direct tool for restoration managers wanting to restore<br />

appropriate ecosystem dynamics. An example here would be the restoration<br />

of species-rich calcareous grassl<strong>and</strong>s on nutrient-poor sites, where active<br />

disturbance in the form of sheep grazing or mowing of the grassl<strong>and</strong>, keeps<br />

the ecosystem from accumulating nutrients <strong>and</strong> from undergoing succession to<br />

shrub- <strong>and</strong> woodl<strong>and</strong>. Another example would be the restoration of speciespoor,<br />

resource-limited inl<strong>and</strong> s<strong>and</strong> dunes, where, at least in some situations, anthropogenic<br />

disturbances in the form of military maneuvers or top-soil removal<br />

keep parts of the ground bare for seedling establishment <strong>and</strong> again prevent the<br />

ecosystem from accumulating nutrients <strong>and</strong> undergoing succession to shrub<strong>and</strong><br />

woodl<strong>and</strong> (Jentsch <strong>and</strong> Beyschlag 2003).<br />

7.4 <strong>Succession</strong>—Not One-Way<br />

Although there are various conceptual models in existence, succession is widely<br />

acknowledged to be a continuous, though often stepwise, process of species<br />

turnover with varying speeds <strong>and</strong> trajectories (see Chapter 1). Thus, the application<br />

of knowledge about temporal dynamics in ecosystems is the fundamental<br />

approach in successional theory that can be linked to restoration. We explore the<br />

idea that temporal dynamics in ecosystems are the product of two interacting<br />

factors: continuous versus discrete processes (Jentsch <strong>and</strong> White, unpublished<br />

data). Continuous processes include gradual accumulation of biomass <strong>and</strong> nutrients,<br />

as the system moves through progressive successional stages. Discrete<br />

processes include the occurrence of disturbance, which can cause rapid transitions<br />

between different ecosystem states or suddenly reset the successional<br />

clock. In addition, disturbance can change continuous processes such as colonization<br />

or extinction of indigenous species to sudden events such as rapid<br />

invasion of an alien species. Thus, restoration managers have two different<br />

options for modifying ecosystem dynamics at restoration sites: manipulating<br />

continuous processes (succession) or making use of discrete events (disturbance).<br />

While most successional sequences seem to follow a progressive accumulation<br />

of biomass, successional studies on very long-term chronosequences or<br />

pollen sequences indicate a decline phase in succession in the absence of a<br />

major disturbance (Iversen 1969, Wardle et al. 2004). This mirrors earlier considerations<br />

of vegetation dynamics as more cyclical than unidirectional (Watt<br />

1947, Remmert 1991, van der Maarel 1996), <strong>and</strong> is included in more recent<br />

conceptualizations of ecosystem dynamics (Holling 1986 <strong>and</strong> 1993). These<br />

ideas of cyclic dynamics implicitly include a decline phase which is followed<br />

by a recovery phase, either following disturbance or after the slow release of<br />

accumulated biomass <strong>and</strong> nutrients.

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