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

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7.3 Assembly<br />

Chapter 7 <strong>Restoration</strong> as a Process of Assembly <strong>and</strong> <strong>Succession</strong> Mediated by Disturbance 153<br />

Concepts of succession <strong>and</strong> community assembly both address temporal dynamics<br />

within ecosystems. However, while succession focuses on the dynamics<br />

of a system following initial colonization of a denuded site (primary succession)<br />

or the dynamics of system regeneration after a disturbance (secondary<br />

succession), community assembly asks the question “How does the suite of<br />

species present at any particular location arrive <strong>and</strong> persist there, <strong>and</strong> how<br />

does that relate to the pool of species available within the region as a whole?”<br />

Hence, questions concerning community assembly, although inherently containing<br />

a dynamic component, are often spatially framed. Indeed, one of the<br />

main methodological approaches to assembly, the categorical approach, aims<br />

to study extant communities <strong>and</strong> provide a snapshot in time of the species,<br />

functional groups, or guilds present there. Different patterns of abundance of<br />

species in different functional groups that are found in a community are then explained<br />

via so-called assembly rules, which are often tested against null models<br />

of no interaction between organisms. An extension of this is the idea of guild<br />

proportionality, which suggests that, within a particular community type, the<br />

proportions of species of different guilds are almost constant across sites in<br />

different developmental stages (Wilson <strong>and</strong> Roxburgh 1994, Wilson 1999). As<br />

a consequence of the theory of guild proportionality, one would expect the<br />

nearest plant of a different species to belong to another guild or functional<br />

group.<br />

Applying such concepts of community assembly to restoration situations<br />

could prove difficult, unless one could unequivocally show, for a given system,<br />

that nonconstant proportions of species in different guilds was a sign of a<br />

highly degraded site compared to a reference ecosystem exhibiting clear guild<br />

proportionality. In this case, a lack of guild proportionality in a degraded site<br />

could be used as an indication of the system being stuck in a certain state, <strong>and</strong><br />

appropriate management measures (usually involving some kind of disturbance<br />

favoring a particular species) could be taken to move the system from the<br />

undesired stable state to a desired stable state. Although we do not have enough<br />

data on guild proportionality in different ecosystems (mainly in grassl<strong>and</strong>s <strong>and</strong><br />

deserts so far) to be able to apply such methods at this stage, it could be a<br />

promising venue for future research linking ecological theory with restoration.<br />

A promising community assembly approach for more direct application to<br />

ecological restoration is the concept of “filters,” whereby a species can only<br />

establish in an area if it can deal with the environmental conditions (i.e., the<br />

abiotic filters) as well as the other organisms it finds there (i.e., the biotic filter)<br />

(Kelt et al. 1995, Weiher <strong>and</strong> Keddy 1995, Zobel 1997, Díaz et al. 1999,<br />

but see Belyea 2004 for caveats). Various conceptualizations of environmental<br />

“filters,” for example very low or high nutrient levels in an ecosystem, also<br />

tend to indicate that the main effect of filters is to vary the species composition<br />

in relation to environmental (<strong>and</strong> hence spatial) variation (Díaz et al.<br />

1999; Hobbs 2004). On the other h<strong>and</strong>, the “response” or dynamic approach<br />

to assembly considers changes in the biotic community <strong>and</strong> the expression of<br />

community assembly “rules” over time, <strong>and</strong> recent treatments emphasize the<br />

dynamic nature of filters, which are likely to change over time as well as spatially<br />

(Fattorini <strong>and</strong> Halle 2004; Hobbs 2004). As discussed in Temperton <strong>and</strong><br />

Hobbs (2004), the dynamic filter approach could prove useful, at least before

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