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Principles of terrestrial ecosystem ecology.pdf

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

Relative units<br />

200<br />

150<br />

100<br />

50<br />

0<br />

0<br />

Ca:Al and<br />

Mg:N ratios<br />

N mineralization<br />

NPP<br />

Nitrification<br />

cycling, trace gas fluxes, and carbon storage.<br />

Chronic nitrogen deposition initially reduces<br />

nitrogen limitation by increasing nitrogen<br />

cycling rates, foliar nitrogen concentrations,<br />

and NPP. Above some threshold, however, the<br />

<strong>ecosystem</strong> becomes saturated with nitrogen<br />

(Fig. 14.9) (Aber et al. 1998). As excess nitrate<br />

and sulfate leach from the soil, they carry with<br />

them cations to maintain charge balance, inducing<br />

calcium and magnesium deficiency in<br />

vegetation (Driscoll et al. 2001). In southern<br />

Sweden, for example, over half <strong>of</strong> the plantavailable<br />

cations have been lost from the upper<br />

70cm <strong>of</strong> soil in the past half century, probably<br />

due to chronic exposure to acid precipitation<br />

(Hallbacken 1992). The exchange complex<br />

becomes more dominated by aluminum and<br />

hydrogen ions, increasing soil acidity and the<br />

likelihood <strong>of</strong> aluminum toxicity. Together this<br />

suite <strong>of</strong> soil changes <strong>of</strong>ten enhances frost susceptibility,<br />

impairs root development, and promotes<br />

herbivory, leading to forest decline in<br />

many areas <strong>of</strong> Europe and the northeastern<br />

United States (Schulze 1989, Aber et al. 1998).<br />

The major surprise, however, has been how<br />

resilient many forests have been to acid rain,<br />

<strong>of</strong>ten retaining most <strong>of</strong> the nitrogen inputs<br />

within the <strong>ecosystem</strong> for as much as two<br />

decades. The resilience <strong>of</strong> <strong>ecosystem</strong>s depends<br />

in part on the magnitude <strong>of</strong> inputs (related to<br />

distance from pollution sources and amount <strong>of</strong><br />

precipitation received) and initial soil acidity,<br />

which in turn depends on parent material and<br />

species composition. Many <strong>ecosystem</strong>s, how-<br />

Foliar N<br />

Leaching<br />

1 2 3<br />

Stage <strong>of</strong> N saturation<br />

Figure 14.9. Changes hypothesized<br />

to occur as forests undergo longterm<br />

nitrogen deposition and nitrogen<br />

saturation. (Redrawn with<br />

permission from BioScience; Aber<br />

et al. 1998.)<br />

ever, now show clear signs <strong>of</strong> nitrogen saturation,<br />

resulting in forest decline, loss <strong>of</strong> acidneutralizing<br />

capacity in lakes, and increasing<br />

nitrogen inputs to streams (Aber et al. 1998,<br />

Carpenter et al. 1998, Driscoll et al. 2001).<br />

Nearly all research on the transport, deposition,<br />

and <strong>ecosystem</strong> consequences <strong>of</strong> anthropogenic<br />

nitrogen has been conducted in the<br />

temperate zone. Further increases in nitrogen<br />

deposition will, however, likely occur primarily<br />

in the tropics and subtropics (Galloway et al.<br />

1995), where plant and microbial growth are<br />

frequently limited by elements other than<br />

nitrogen. These <strong>ecosystem</strong>s might therefore<br />

show more immediate nitrogen loss in trace<br />

gases or leaching in response to nitrogen deposition<br />

(Matson et al. 1998). On the other hand,<br />

soil properties such as high clay content or<br />

cation exchange capacity may allow tropical<br />

soils to sequester substantial quantities <strong>of</strong> nitrogen<br />

before they become leaky.<br />

Biomass burning transfers nutrients directly<br />

from <strong>terrestrial</strong> pools to the atmosphere and<br />

then to down-wind <strong>ecosystem</strong>s. Biomass combustion<br />

releases a suite <strong>of</strong> gases that reflect the<br />

elemental concentrations in vegetation and fire<br />

intensity. About half <strong>of</strong> dry biomass consists <strong>of</strong><br />

carbon, so the predominant gases released are<br />

carbon compounds in various stages <strong>of</strong> oxidation,<br />

including carbon dioxide (CO2), methane<br />

(CH4), carbon monoxide (CO), and smaller<br />

quantities <strong>of</strong> nonmethane hydrocarbons. The<br />

atmospheric role <strong>of</strong> these gases varies. CO2 and<br />

CH4 are greenhouse gases, whereas carbon

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