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Principles of terrestrial ecosystem ecology.pdf

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occurs near their edges, rather than in the<br />

center, where the overlying air is so stable<br />

that it saturates rapidly and supports a relatively<br />

low evaporation rate. A swamp with<br />

interspersed patches <strong>of</strong> vegetation would<br />

have greater convective turbulence and overall<br />

evaporation than homogeneous water or moist<br />

vegetation. When <strong>ecosystem</strong> patches that differ<br />

strongly in energy partitioning are larger in<br />

diameter than the depth <strong>of</strong> the planetary<br />

boundary layer (greater than about 10km),<br />

they can modify mesoscale atmospheric circulations<br />

and cloud and precipitation patterns<br />

(Pielke and Avisar 1990, Weaver and Avissar<br />

2001).<br />

Seasonal Energy Exchange<br />

Over sufficiently long time scales, energy<br />

outputs are tightly coupled to inputs because<br />

there is little energy storage at Earth’s surface.<br />

Most <strong>ecosystem</strong>s have a limited capacity for<br />

long-term energy storage in vegetation and<br />

surface soils. Consequently, energy losses to the<br />

atmosphere closely track solar inputs on both a<br />

daily and a seasonal basis, although the form in<br />

which this energy is lost (sensible vs. latent heat<br />

flux) varies, depending on moisture availability.<br />

Ground heat fluxes are usually negligible when<br />

averaged over 24h (Oke 1987). Important<br />

exceptions to this generalization are water<br />

bodies such as lakes and oceans, in which the<br />

solar inputs <strong>of</strong>ten penetrate to depth, resulting<br />

in substantial warming <strong>of</strong> the water, and<br />

some high-latitude regions. Water bodies <strong>of</strong>ten<br />

absorb substantial energy in spring and early<br />

summer, when the solar angle is greatest,<br />

causing the water to warm. There is a net<br />

release <strong>of</strong> energy in the autumn, moderating<br />

the temperatures <strong>of</strong> adjacent <strong>terrestrial</strong> surfaces<br />

(see Chapter 2). This seasonal pattern <strong>of</strong><br />

energy exchange drives the annual or semiannual<br />

turnover <strong>of</strong> lakes (see Chapter 10). Permafrost<br />

contributes to a seasonal imbalance in<br />

energy absorption and release in cold climates.<br />

In the arctic, for example, 10 to 20% <strong>of</strong> the<br />

energy absorbed during summer is consumed<br />

by thawing <strong>of</strong> frozen soil. This energy is<br />

released back to the atmosphere the next<br />

winter, when the soil refreezes (Chapin et al.<br />

2000a).<br />

Water Inputs to Ecosystems 77<br />

Snow-covered surfaces experience threshold<br />

changes in energy exchange at the time<br />

<strong>of</strong> snowmelt (Liston 1999). The high albedo<br />

<strong>of</strong> snow-covered surfaces minimizes energy<br />

absorption until snowmelt occurs, at which<br />

time there is a dramatic increase in the energy<br />

absorbed by the surface and transferred to<br />

the atmosphere. This may result in abrupt<br />

increases in regional air temperature after<br />

snowmelt. Leaf out also alters energy exchange<br />

by both changing albedo and increasing evapotranspiration<br />

at the expense <strong>of</strong> sensible heat<br />

flux.<br />

Water Inputs to Ecosystems<br />

Precipitation is the major water input to most<br />

<strong>terrestrial</strong> <strong>ecosystem</strong>s. Global and regional controls<br />

over precipitation therefore determine the<br />

quantity and seasonality <strong>of</strong> water inputs to most<br />

<strong>ecosystem</strong>s (see Chapter 2). In <strong>ecosystem</strong>s that<br />

receive some precipitation as snow, however,<br />

the water contained in the snowpack does not<br />

enter the soil until snow melt, <strong>of</strong>ten months<br />

after the precipitation occurs. This causes the<br />

seasonality <strong>of</strong> water input to soils to differ from<br />

that <strong>of</strong> precipitation.<br />

Vegetation in some <strong>ecosystem</strong>s, particularly<br />

in riparian zones, accesses additional groundwater<br />

that flows laterally through the <strong>ecosystem</strong>.<br />

Desert communities <strong>of</strong> phreatophytes<br />

(deep-rooted plants that tap groundwater),<br />

for example, may absorb sufficient groundwater<br />

that the <strong>ecosystem</strong> loses more water<br />

in transpiration than it receives in precipitation.<br />

Lakes and streams also receive most<br />

<strong>of</strong> their water inputs from groundwater or<br />

run<strong>of</strong>f that drains from adjacent <strong>terrestrial</strong><br />

<strong>ecosystem</strong>s. Water inputs to freshwater <strong>ecosystem</strong>s<br />

are therefore only indirectly linked to<br />

precipitation.<br />

In <strong>ecosystem</strong>s with frequent fog, canopy<br />

interception <strong>of</strong> fog increases the water inputs to<br />

<strong>ecosystem</strong>s, when cloud droplets that might not<br />

otherwise precipitate are deposited on leaf surfaces<br />

and are absorbed by leaves or drip from<br />

the canopy to the soil. The coastal redwood<br />

trees <strong>of</strong> California, for example, depend on<br />

fog-derived water inputs during summer, when<br />

precipitation is low, but fog occurs frequently.

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