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

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70 Joe Walker <strong>and</strong> Paul Reddell<br />

demonstrate the need for detailed process knowledge about succession when<br />

trying to restore old l<strong>and</strong>scapes following l<strong>and</strong>-use disturbances.<br />

4.2 Natural Retrogressive <strong>Succession</strong><br />

<strong>Succession</strong> theory in ecology had its beginnings in studies of vegetation changes<br />

across s<strong>and</strong> dune chronosequences (Warming 1895, Cowles 1895). A progressive<br />

build up to a maximum biomass <strong>and</strong> species richness were observed across<br />

the dunes, <strong>and</strong> these changes were accompanied by gradual changes in the<br />

physicochemical composition of the soils. The early vegetation work was contemporary<br />

with the idea of geographical cycles proposed by Davis (1899) to<br />

explain how l<strong>and</strong>forms evolved over time, <strong>and</strong> his schema included renewal<br />

<strong>and</strong> degradation. The vegetation observations led to the idea of a “climax community,”<br />

that is, the progressive development toward an optimum expression of<br />

vegetation for the climate <strong>and</strong> soils of a region. As ecological ideas developed,<br />

the climax state was seen to fluctuate but it was generally regarded as in a<br />

stable or dynamic equilibrium state. In terms of restoration actions, succession<br />

trajectories toward the original “climax” state have been viewed as desirable.<br />

Debate continued about the dynamics <strong>and</strong> trajectory of successional processes,<br />

but gradually it was realized that natural post-climax states exist. Early studies<br />

using pollen analysis in peat or mor layers in postglacial deposits (Iversen<br />

1964) showed examples of ecosystems with permanently reduced productivity.<br />

These so-called retrogressive successions were associated with leaching of the<br />

soils during pedogenesis (natural retrogression) <strong>and</strong> man-made disturbances<br />

(secondary retrogression). A clear demonstration of a post-climax natural retrogression<br />

was described for an intact aeolian s<strong>and</strong> dune chronosequences at<br />

Cooloola, Queensl<strong>and</strong>, Australia by Walker et al. (1981) <strong>and</strong> Thompson (1981<br />

<strong>and</strong> 1983). Dune building at Cooloola was episodic, <strong>and</strong> some parts of earlier<br />

dune systems were not buried by subsequent wind-blown deposits. These exposed<br />

parts were subjected to weathering, leaching, <strong>and</strong> erosion, <strong>and</strong> form an<br />

age sequence that stretches over some 750K years (Thompson 1992). These<br />

studies showed that for freely drained sites vegetation type, species richness,<br />

st<strong>and</strong>ing biomass, <strong>and</strong> soil carbon accumulation varied with dune age. After a<br />

period of biomass build up (progressive succession) seen in the youngest four<br />

dune systems, vegetation biomass <strong>and</strong> the store of organic material in the surface<br />

soils declined in the oldest three systems (retrogressive succession). Vitousek<br />

<strong>and</strong> Reiners (1975), Hedin et al. (2003) <strong>and</strong> Wardle et al. (2004) have shown<br />

for a series of very long-term chronosequences, that ecosystem decline is a<br />

widespread phenomenon. However, the mechanisms controlling retrogression<br />

vary between bioregions <strong>and</strong> with disturbance regimes <strong>and</strong> intensity.<br />

The hypothesis developed from the Cooloola study (Fig. 4.1) relevant to planning<br />

restoration actions is that pedologically young l<strong>and</strong>scapes when disturbed<br />

tend to recover toward the previous state, whereas old systems become “leaky”<br />

or cannot recover critical system functions <strong>and</strong> trend toward a new system state<br />

with lower biomass <strong>and</strong> complexity.<br />

Old l<strong>and</strong>scapes are common in Australia but also occur in most parts of the<br />

southern hemisphere <strong>and</strong> in tropical areas. At a global scale old l<strong>and</strong>scapes predominate,<br />

yet many of the developments in succession theory were carried out in<br />

pedologically young l<strong>and</strong>scapes, <strong>and</strong> perhaps the same is true of attempts to develop<br />

links with restoration activities. Like many areas in the tropics, Australia

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