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Principles of terrestrial ecosystem ecology.pdf

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310 14. Landscape Heterogeneity and Ecosystem Dynamics<br />

Disturbance interval / Recovery interval [log scale]<br />

10.0<br />

5.0<br />

1.0<br />

0.5<br />

0.1<br />

0.05<br />

0.01<br />

Equilibrium<br />

or steady<br />

state<br />

Figure 14.2. Effect <strong>of</strong> disturbance size (relative to<br />

the size <strong>of</strong> the landscape) and disturbance frequency<br />

(relative to the time required for the <strong>ecosystem</strong> to<br />

recover) on the stability <strong>of</strong> landscape processes.<br />

Landscapes are close to equilibrium when disturbances<br />

are small (relative to the size <strong>of</strong> the study<br />

area) and when they are infrequent (relative to the<br />

time required for <strong>ecosystem</strong> recovery). As distur-<br />

Fire can also create large patches <strong>of</strong> a single<br />

successional stage on the landscape (Johnson<br />

1992). The 1988 fires in Yellowstone National<br />

Park that burned 3200km 2 <strong>of</strong> old pine forest<br />

altered large areas <strong>of</strong> the landscape. Fires <strong>of</strong><br />

this magnitude and intensity recur every few<br />

centuries (Romme and Knight 1982). Longterm<br />

human fire suppression has increased the<br />

proportion <strong>of</strong> late-successional communities in<br />

many areas. This results in a more homogeneous<br />

and spatially continuous, fuel-rich environment<br />

in which fires can burn large areas.<br />

Even large disturbed areas, however, are <strong>of</strong>ten<br />

internally quite patchy. Fires, for example,<br />

generate islands <strong>of</strong> unburned vegetation<br />

and patches <strong>of</strong> varying burn severity. These<br />

unburned islands act as seed sources for postfire<br />

succession and protective cover for wildlife,<br />

greatly reducing the effective size <strong>of</strong><br />

the disturbance (Turner et al. 1997). In many<br />

cases, these patches become less distinct as<br />

Stable, low<br />

variance<br />

Stable<br />

high<br />

variance<br />

Stable,<br />

very high<br />

variance<br />

Stable<br />

low<br />

variance<br />

0.25<br />

0.50<br />

Disturbance extent / Landscape extent [log scale]<br />

Unstable<br />

system,<br />

malfunction<br />

or crash<br />

0.75<br />

bances become more frequent or larger, the landscape<br />

becomes more heterogeneous and there is<br />

increasing probability that the individual patches<br />

may undergo a different successional trajectory.<br />

(Redrawn with permission from Landscape Ecology,<br />

Vol. 8 © 1993 Kluwer Academic Publishers; Turner<br />

et al. 1993.)<br />

succession proceeds, so spatial heterogeneity<br />

may decline with time in non–steady state<br />

mosaics.<br />

Human-induced disturbances alter the<br />

natural patterns and magnitude <strong>of</strong> landscape<br />

heterogeneity. Half <strong>of</strong> the ice-free <strong>terrestrial</strong><br />

surface has been transformed by human activities<br />

(Turner et al. 1990). We have cleared or<br />

selectively harvested forests, converted grasslands<br />

and savannas to pastures or agricultural<br />

systems, drained wetlands, flooded uplands, and<br />

irrigated dry lands. Isolated land use changes<br />

may augment landscape heterogeneity by creating<br />

small patches within a matrix <strong>of</strong> largely<br />

natural vegetation. As land use change<br />

becomes more extensive, however, the humandominated<br />

patches become the matrix in which<br />

isolated fragments <strong>of</strong> natural <strong>ecosystem</strong>s are<br />

embedded, causing a reduction in landscape<br />

heterogeneity. These contrasting impacts <strong>of</strong><br />

human actions on landscape heterogeneity

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