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Principles of terrestrial ecosystem ecology.pdf

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13<br />

Temporal Dynamics<br />

Ecosystem processes constantly change in response to variation in environment over<br />

all time scales. This chapter describes the major patterns and controls over the temporal<br />

dynamics <strong>of</strong> <strong>ecosystem</strong>s.<br />

Introduction<br />

Ecosystems are always recovering from past<br />

changes. In earlier chapters we emphasized<br />

<strong>ecosystem</strong> responses to the current environment.<br />

Ecosystems are, however, always responding<br />

to past changes that have occurred<br />

over all time scales (Holling 1973, Wu and<br />

Loucks 1995). These changes include relatively<br />

predictable daily and seasonal variations, less<br />

predictable changes in weather (e.g., passage<br />

<strong>of</strong> weather fronts, El Niño events, and glacial<br />

cycles), and occurrence <strong>of</strong> disturbances (e.g.,<br />

tree falls, herbivore outbreaks, fires, and volcanic<br />

eruptions). Consequently, the behavior <strong>of</strong><br />

an <strong>ecosystem</strong> is always influenced by both the<br />

current environment and many previous environmental<br />

fluctuations and disturbances.<br />

The global environment is changing more<br />

rapidly than it has for millions <strong>of</strong> years. These<br />

changes result from an exponentially rising<br />

human population that shows an everyincreasing<br />

technological capacity to alter<br />

Earth’s environment and <strong>ecosystem</strong>s. Perhaps<br />

the most urgent need in <strong>ecosystem</strong> <strong>ecology</strong> is to<br />

improve our understanding <strong>of</strong> factors governing<br />

the stability and change in ecological<br />

systems (Box 13.1). This understanding is critical<br />

to managing <strong>ecosystem</strong>s so they sustain<br />

their diversity and other important ecological<br />

attributes and so <strong>ecosystem</strong>s continue to<br />

produce the goods and services required by<br />

society. This chapter addresses the basis <strong>of</strong> the<br />

temporal dynamics <strong>of</strong> <strong>ecosystem</strong>s.<br />

Fluctuations in<br />

Ecosystem Processes<br />

Interannual Variability<br />

Ecosystem processes measured in 1 year are<br />

seldom representative <strong>of</strong> the long-term mean.<br />

Many <strong>ecosystem</strong> processes are sensitive to<br />

interannual variations in weather and to fluctuations<br />

in the internal dynamics <strong>of</strong> <strong>ecosystem</strong>s,<br />

such as outbreaks <strong>of</strong> herbivores or pathogens.<br />

The same <strong>ecosystem</strong>, for example, can change<br />

from being a carbon source in one year to a<br />

carbon sink in the next (Goulden et al. 1998).<br />

The production at a particular trophic level can<br />

change from being limited by food to being<br />

limited by predation. Some <strong>of</strong> the interannual<br />

variability in <strong>ecosystem</strong> dynamics reflects<br />

processes that are potentially predictable, such<br />

as the cyclic variation in climate. El Niño events<br />

are a consequence <strong>of</strong> large-scale oscillations in<br />

the global ocean-atmosphere system that recur<br />

every 2 to 10 years (see Chapter 2). These are<br />

associated with relatively repeatable climatic<br />

281

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