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

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

indigenous systems <strong>and</strong> un-manipulated systems. Similarly, do manipulations<br />

of animal herbivore densities, either through restoration of native species or<br />

the reduction of pest species, also restore soil communities? Models, long-term<br />

experiments, <strong>and</strong> observational studies (such as across well defined chronosequences)<br />

can be also used to determine what the long-term consequences of<br />

restoration treatments are for successional processes or ecosystem properties,<br />

although these approaches are probably most useful for systems that are highly<br />

predictable.<br />

Altering the aboveground or belowground community by removing herbivores<br />

or invasive plants may not always produce the desired result for succession<br />

or ecosystem properties (i.e., their impacts are not immediately reversed<br />

by removals) because of belowground legacies that may persist for a long time.<br />

For example, plant invaders interact strongly with belowground communities<br />

both directly through promoting NPP (higher C inputs), <strong>and</strong> influencing litter<br />

quality <strong>and</strong> nutrient inputs to the soil; <strong>and</strong> indirectly through interactions<br />

with native plants or species that show different plant–soil feedbacks. Many<br />

of these effects will have long-term implications for successional trajectories<br />

<strong>and</strong> ecosystem properties that cannot be mitigated by the removal of weeds,<br />

the most commonly used tool in restoration efforts. In addition to persistent<br />

effects or legacies of treatments, there will also be important lags in response<br />

to different components of the soil subsystem to aboveground manipulations.<br />

For example, if nutrient inputs into a system (e.g., from a N-fixing weed or<br />

atmospheric deposition) are offset by reducing these inputs or through soil impoverishment<br />

treatments, does the belowground system return to its previous<br />

state quickly or does it retain accumulated nutrients <strong>and</strong> switch to an alternative<br />

steady state dominated by early successional, nutrient-dem<strong>and</strong>ing species?<br />

Some components of the soil subsystem, such as the microbial biomass, will<br />

respond rapidly to aboveground treatments (i.e., weeks to months) <strong>and</strong> will be<br />

more sensitive indicators of changes in belowground processes to restoration<br />

treatments than will more slowly responding soil meso- <strong>and</strong> macrofauna (i.e.,<br />

months to years) or pools of soil organic matter <strong>and</strong> nutrients (i.e., months to<br />

decades). The key unresolved issue is which long-term ecosystem properties or<br />

community trajectories in restoration are predictable from short-term shifts in<br />

aboveground–belowground interactions.<br />

In summary, there are several critical areas for future research linking aboveground<br />

<strong>and</strong> belowground processes in succession to underst<strong>and</strong>ing restoration.<br />

First, are the shifts or trajectories in aboveground communities manipulated in<br />

restoration also mirrored belowground? Current studies document shifts in both<br />

aboveground <strong>and</strong> belowground communities, <strong>and</strong> clearly point to links between<br />

these components, but recognizing the importance of these links to succession,<br />

ecosystem development or restoration requires the inclusion of soil biologists<br />

alongside aboveground expertise. Examples of research that should provide<br />

insights into restoration include: (i) determining whether successful restoration<br />

aboveground also restores belowground communities, (ii) underst<strong>and</strong>ing<br />

whether soil mutualists or pathogens can be used to either promote native species<br />

or suppress nonnative species respectively, (iii) underst<strong>and</strong>ing in what situations<br />

soil manipulations (e.g., C additions) help or hinder ecosystem restoration, <strong>and</strong><br />

(iv) identifying to what extent shifts in belowground processes during primary<br />

<strong>and</strong> secondary succession from field studies can be used to determine restoration<br />

success. The implications of these topics for restoration are obvious: soils <strong>and</strong>

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