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Principles of terrestrial ecosystem ecology.pdf

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92 4. Terrestrial Water and Energy Balance<br />

however, is <strong>of</strong>ten relatively small. Maximum<br />

stomatal conductance <strong>of</strong> individual leaves is<br />

relatively similar among natural <strong>ecosystem</strong>s<br />

(Körner 1994, Kelliher et al. 1995). Woody<br />

and herbaceous <strong>ecosystem</strong>s, for example, have<br />

similar stomatal conductance <strong>of</strong> individual<br />

leaves (Körner 1994) and similar surface conductance<br />

<strong>of</strong> entire <strong>ecosystem</strong>s (Kelliher et al.<br />

1995). Crops, however, which have about 50%<br />

higher stomatal conductance than does natural<br />

vegetation, also have about 50% higher surface<br />

conductance (Schulze et al. 1994, Kelliher et al.<br />

1995). There are currently insufficient data to<br />

know whether ecological variation in stomatal<br />

conductance associated with gradients in soil<br />

fertility causes similar variation in surface<br />

conductance and therefore evapotranspiration<br />

from <strong>ecosystem</strong>s.<br />

In summary, aerodynamic roughness, which<br />

depends on plant height and the number <strong>of</strong><br />

roughness elements, is the main way in which<br />

vegetation influences evapotranspiration from<br />

dry canopies under conditions <strong>of</strong> adequate<br />

water supply. Stomatal conductance exerts<br />

additional control in some <strong>ecosystem</strong>s. It also<br />

has an increasingly important control over<br />

evapotranspiration as soil moisture declines.<br />

In other words, stomatal conductance accounts<br />

for temporal variation in evapotranspiration in<br />

response to variation in soil moisture, but<br />

surface roughness is the major factor explaining<br />

differences in evapotranspiration among<br />

moist <strong>ecosystem</strong>s.<br />

Vegetation structure also influences the relative<br />

importance <strong>of</strong> different climatic variables<br />

in regulating evapotranspiration. In aerodynamically<br />

rough, well-mixed canopies such as<br />

open-canopied forests, the moisture content <strong>of</strong><br />

the air within the canopy is similar to that <strong>of</strong> the<br />

bulk air above the canopy. Under these wellcoupled<br />

conditions, evapotranspiration is determined<br />

more by the moisture content <strong>of</strong> the<br />

air (and the accompanying stomatal response)<br />

than by net radiation (Waring and Running<br />

1998). In short canopies, by contrast, the air<br />

adjacent to the leaves is mixed less readily with<br />

the bulk air above the canopy, allowing evapotranspiration<br />

to increase the moisture content<br />

within the canopy environment. In other words,<br />

the canopy air becomes decoupled from condi-<br />

Table 4.5. Decoupling coefficient <strong>of</strong> vegetation<br />

canopies in the field under conditions <strong>of</strong> adequate<br />

moisture supply.<br />

Vegetation Decoupling coefficient a<br />

Alfalfa 0.9<br />

Strawberry patch 0.85<br />

Permanent pasture 0.8<br />

Grassland 0.8<br />

Tomato field 0.7<br />

Wheat field 0.6<br />

Prairie 0.5<br />

Cotton 0.4<br />

Heathland 0.3<br />

Citrus orchard 0.3<br />

Forest 0.2<br />

Pine woods 0.1<br />

a A completely smooth surface has a decoupling coefficient<br />

<strong>of</strong> 1.0, and a canopy in which the air is identical to that in<br />

the atmosphere has a decoupling coefficient <strong>of</strong> zero.<br />

Data from Jarvis and McNaughton (1986) and Jones (1992).<br />

tions in the bulk atmosphere. In these smooth<br />

canopies, evapotranspiration is determined<br />

more by net radiation than by the moisture<br />

content <strong>of</strong> the bulk air, just as when canopies<br />

are wet.The decoupling coefficient is a measure<br />

<strong>of</strong> the degree to which a canopy is decoupled<br />

from the atmosphere (Table 4.5) (Jarvis and<br />

McNaughton 1986). It is determined primarily<br />

by canopy height. In summary, net radiation is<br />

the dominant environmental control over evapotranspiration<br />

in short canopies, whereas the<br />

vapor pressure deficit is the dominant control<br />

in tall canopies, when water is freely available<br />

(Waring and Running 1998).<br />

Changes in Storage<br />

Water inputs that exceed outputs replenish<br />

water that is stored in soil and groundwater.<br />

Water that enters the soil is retained until the<br />

soil reaches field capacity. Additional water<br />

moves downward to groundwater. In cold climates<br />

in winter, most <strong>of</strong> the precipitation input<br />

is stored above ground in the snowpack. The<br />

snowpack substantially increases the quantity<br />

<strong>of</strong> water that an <strong>ecosystem</strong> can store and<br />

the residence time <strong>of</strong> water in the <strong>ecosystem</strong>.<br />

Stored water supports evapotranspiration<br />

at times when evapotranspiration exceeds precipitation;<br />

the declines in soil moisture during

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