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Principles of terrestrial ecosystem ecology.pdf

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

enrich soils on hilltops where they bed down<br />

at night. Migrating salmon perform a similar<br />

nutrient-transport role in streams. They feed<br />

primarily in the open ocean and then return to<br />

small streams where they spawn, die, and<br />

decompose.The nutrients carried by the salmon<br />

from the ocean can sustain a substantial<br />

proportion <strong>of</strong> the algal and insect productivity<br />

<strong>of</strong> small streams. These nutrient subsidies<br />

can be transferred to adjoining <strong>terrestrial</strong><br />

habitats by bears and otters that feed on salmon<br />

or by predators <strong>of</strong> insects that emerge from<br />

streams.<br />

Nutrient Turnover<br />

Species differences in litter quality magnify site<br />

differences in soil fertility. Differences among<br />

plant species in tissue quality strongly influence<br />

litter decomposition rates (see Chapter 7).<br />

Litter from low-nutrient-adapted species<br />

decomposes slowly because <strong>of</strong> the negative<br />

effects on soil microbes <strong>of</strong> low concentrations<br />

<strong>of</strong> nitrogen and phosphorus and high concentrations<br />

<strong>of</strong> lignin, tannins, waxes, and other<br />

recalcitrant or toxic compounds. This slow<br />

decomposition <strong>of</strong> litter from species characteristic<br />

<strong>of</strong> nutrient-poor sites reinforces the low<br />

nutrient availability <strong>of</strong> these sites (Hobbie<br />

1992, Wilson and Agnew 1992). Species from<br />

high-resource sites, in contrast, produce rapidly<br />

decomposing litter due to its higher nitrogen<br />

and phosphorus content and fewer recalcitrant<br />

compounds, enhancing rates <strong>of</strong> nutrient<br />

turnover in nutrient-rich sites.<br />

Experimental planting <strong>of</strong> species on a<br />

common soil shows that species differences in<br />

litter quality can alter soil fertility quite quickly.<br />

Early successional prairie grasses, whose litter<br />

has a low C:N ratio, for example, causes an<br />

increase in the nitrogen mineralization rate <strong>of</strong><br />

soil within 3 years, compared to the same soil<br />

planted with late-successional species whose<br />

litter has a high C:N ratio (Wedin and Tilman<br />

1990) (Fig. 12.5).<br />

Seasonality <strong>of</strong> Resource Capture<br />

Phenological specialization could increase<br />

resource capture. Phenological specialization in<br />

the timing <strong>of</strong> plant activity can increase the<br />

Net N mineralization rate<br />

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

14<br />

7<br />

0<br />

(44)<br />

(56)<br />

(64)<br />

As Ar Pp Ss Ag<br />

Species<br />

(109)<br />

C:N ratios<br />

(122)<br />

Figure 12.5. Effects <strong>of</strong> prairie grass species on nitrogen<br />

mineralization when grown on soils with containing<br />

100gNm -2 (Wedin and Tilman 1990). Grasses<br />

range from early to late successional in the following<br />

order: Agrostis scabra (As), Agropyron repens<br />

(Ar), Poa pratensis (Pp), Schizochyrium scoparium<br />

(Ss), and Andropogon gerardi (Ag). Data are means<br />

±95% confidence interval (CI). Numbers are the C:<br />

N ratios <strong>of</strong> aboveground biomass.<br />

total time available for plants to acquire<br />

resources from their environment. This is most<br />

evident when coexisting species differ in the<br />

timing <strong>of</strong> their maximal activity. In mixed grasslands,<br />

for example, C4 species are generally<br />

more active in the warmer, drier part <strong>of</strong> the<br />

growing season than are C 3 species. Consequently<br />

C 3 species account for most early<br />

season production, and C 4 species account for<br />

most late-season production. Similarly, in the<br />

Sonoran desert, there is a different suite <strong>of</strong><br />

annuals that becomes active after winter than<br />

after summer rains. In both cases, phenological<br />

specialization probably enhances NPP and<br />

nitrogen cycling. In mixed-cropping agricultural<br />

<strong>ecosystem</strong>s, phenological specialization is<br />

more effective in enhancing production than<br />

are species differences in rooting depth<br />

(Steiner 1982).<br />

The <strong>ecosystem</strong> consequences <strong>of</strong> phenological<br />

specialization to exploit the extremes <strong>of</strong> the<br />

growing season are less clear. Evergreen<br />

forests, for example, have a longer photosynthetic<br />

season than deciduous forests, but most<br />

carbon gain occurs in midseason in both forest<br />

types, when conditions are most favorable

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