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

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

from the soil, <strong>and</strong> thus enhance NPP. This emphasizes the importance of underst<strong>and</strong>ing<br />

feedbacks between the aboveground <strong>and</strong> belowground subsystems<br />

when attempting to predict the ecological consequences of ecosystem restoration.<br />

Studies on these isl<strong>and</strong>s have also shown that the importance of species effects<br />

on ecosystem properties changes during successional time. An ongoing<br />

experiment set up in 1996 (first 7 years reported by Wardle <strong>and</strong> Zackrisson<br />

2005) involved experimental removals of various combinations of understory<br />

plant species on each of 30 isl<strong>and</strong>s across the sequence. This work revealed that<br />

two of the dominant understory species (V. myrtillus <strong>and</strong> V. vitis-idaea) drove<br />

belowground processes on the large isl<strong>and</strong>s but not the small ones. Removals of<br />

these species significantly reduced litter decomposition, soil microbial biomass,<br />

<strong>and</strong> soil respiration on the large isl<strong>and</strong>s to levels more characteristic of small<br />

isl<strong>and</strong>s. In contrast, species removals on the small isl<strong>and</strong>s had no detectable<br />

effects on these properties, probably because of the increasing relative importance<br />

of abiotic drivers. This points to species’ effects (<strong>and</strong> consequences<br />

of species losses) in ecosystems being context-dependent <strong>and</strong> of diminishing<br />

importance as retrogressive succession proceeds, as well as to the role of understory<br />

species in governing the effects of successional status on the functioning<br />

of the belowground subsystem. Any consequences of restoration effort for both<br />

the aboveground <strong>and</strong> belowground subsystems will therefore depend in a large<br />

part upon how the relative abundances of these understory species are influenced<br />

by frequency of fire regime.<br />

The isl<strong>and</strong> system provides evidence that reducing fire frequency, <strong>and</strong> the<br />

ecosystem retrogression that results, greatly influences ecosystem C sequestration<br />

(Wardle et al. 2003). As retrogressive succession proceeds (i.e., as isl<strong>and</strong>s<br />

become smaller), st<strong>and</strong>ing plant biomass <strong>and</strong> NPP are both impaired,<br />

in both the tree <strong>and</strong> understory shrub layers. This results in less C storage<br />

aboveground. Further, decomposition rates of plant litter are reduced, for three<br />

reasons: (1) because plant species that produce poorer litter quality (Picea <strong>and</strong><br />

Empetrum) begin to dominate; (2) because of phenotypic plasticity within plant<br />

species (i.e., plants of a given species producing poorer quality litter during retrogression);<br />

<strong>and</strong> (3) because of lower activity of the decomposer microflora.<br />

Further, during retrogression, decomposition is impaired before NPP, which<br />

results in a net gain of C to the soil. As such, the largest isl<strong>and</strong>s store less<br />

than 5 kg C/m 2 in the soil while some of the smallest ones store over 35<br />

kg C/m 2 . Because the belowground (rather than the aboveground) component<br />

stores the majority of C in these forests, there is net ecosystem C sequestration<br />

over time, in the order of 0.45 kg C/m 2 for every century without a<br />

major fire. These results point to fire suppression in forest ecosystems contributing<br />

significantly to C sequestration, <strong>and</strong> if these patterns are widespread<br />

elsewhere then this may play an important part in the global carbon cycle. They<br />

also point to the fact that ecosystem restoration involving the reintroduction<br />

or encouragement of natural fire regimes (<strong>and</strong> consequent reversion of retrogressive<br />

succession), would in turn alter the balance between aboveground <strong>and</strong><br />

belowground processes, <strong>and</strong> thereby greatly influence total ecosystem carbon<br />

storage.<br />

While there has been substantial fire suppression in the boreal forests of northern<br />

Sc<strong>and</strong>inavia over the past couple of centuries, there is recent recognition<br />

that the use of fire in these systems may have beneficial consequences from a

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