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Principles of terrestrial ecosystem ecology.pdf

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268 12. Community Effects on Ecosystem Processes<br />

the loss <strong>of</strong> one <strong>of</strong> a group <strong>of</strong> ecologically similar<br />

species because, in the latter case, the remaining<br />

species could continue to perform the relevant<br />

ecological functions <strong>of</strong> that functional<br />

type. The more species there are in a functional<br />

type, the less likely it is that a gain or loss <strong>of</strong> a<br />

single species from that functional type will<br />

have large <strong>ecosystem</strong> effects. Our challenge, as<br />

ecologists, is to identify the traits <strong>of</strong> organisms<br />

that have strong effects on <strong>ecosystem</strong>s; species<br />

with these traits are likely to be strong interactors<br />

in <strong>ecosystem</strong>s (Paine 2000).<br />

Species interactions govern the traits that are<br />

expressed most clearly in <strong>ecosystem</strong>s. The<br />

impact <strong>of</strong> a species on <strong>ecosystem</strong> processes<br />

depends on its interactions with other species.<br />

The impact <strong>of</strong> deer on <strong>terrestrial</strong> vegetation or<br />

the impact <strong>of</strong> Daphnia on algal biomass <strong>of</strong><br />

lakes, for example, depends on the density <strong>of</strong><br />

their predators. The mechanisms by which<br />

species diversity influences <strong>ecosystem</strong> processes<br />

<strong>of</strong>ten depend on species interactions<br />

such as competition, facilitation, and predation.<br />

Species Effects on<br />

Ecosystem Processes<br />

Species are most likely to have strong <strong>ecosystem</strong><br />

effects when they alter interactive controls,<br />

which are the general factors that directly<br />

regulate <strong>ecosystem</strong> processes. These controls<br />

include the supply <strong>of</strong> resources that are essential<br />

for primary production, climate, functional<br />

N inputs (g m -2 yr -1 )<br />

2.4<br />

1.2<br />

0<br />

Myrica<br />

absent<br />

Myrica<br />

present<br />

Net N mineralization rate<br />

(g N m -2 yr -1 )<br />

6<br />

3<br />

0<br />

Myrica<br />

absent<br />

types <strong>of</strong> organisms, disturbance regime, and<br />

human activities (see Chapter 1).<br />

Species Effects on Resources<br />

Resource Supply<br />

Species traits that influence the supply <strong>of</strong> limiting<br />

resources have major impacts. The supply<br />

<strong>of</strong> resources required for growth <strong>of</strong> primary<br />

producers is one <strong>of</strong> the interactive controls to<br />

which <strong>ecosystem</strong> processes are most sensitive<br />

(see Chapter 1). These resources include light,<br />

nutrients, and, on land, water. For this reason,<br />

species traits that alter the supply <strong>of</strong> limiting<br />

resources will substantially alter <strong>ecosystem</strong><br />

processes.<br />

The introduction <strong>of</strong> a strong nitrogen fixer<br />

into a community that lacks such species can<br />

substantially alter nitrogen availability and<br />

cycling. Invasion by the exotic nitrogen-fixing<br />

tree Myrica faya in Hawaii, for example,<br />

increased nitrogen inputs, litter nitrogen concentration,<br />

and nitrogen availability (Vitousek<br />

et al. 1987) (Fig. 12.3).A nitrogen-fixing invader<br />

is most likely to be successful in <strong>ecosystem</strong>s that<br />

are nitrogen limited; have no strong nitrogen<br />

fixers; and have adequate phosphorus, micronutrients,<br />

and light (Vitousek and Howarth 1991).<br />

Thus we expect large <strong>ecosystem</strong> changes from<br />

invasion <strong>of</strong> nitrogen-fixing species primarily in<br />

combinations <strong>of</strong> the following circumstances:<br />

(1) low nitrogen supply (early primary succession<br />

in the temperate zone and in other<br />

low-nitrogen environments), (2) distant from<br />

Myrica<br />

present<br />

Litter N concentration (%)<br />

1.4<br />

0.7<br />

0<br />

Myrica<br />

absent<br />

Myrica<br />

present<br />

Figure 12.3. Impact <strong>of</strong> the nitrogen-fixing tree Myrica faya on nitrogen inputs, litter nitrogen concentration,<br />

and nitrogen mineralization rate in a Hawaiian montane forest (Vitousek et al. 1987).

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