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Principles of terrestrial ecosystem ecology.pdf

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Consumption Efficiency<br />

Consumption efficiency is determined primarily<br />

by food quality and secondarily by predation.<br />

Consumption efficiency is the proportion<br />

<strong>of</strong> the production at one trophic level that is<br />

ingested by the next trophic level (I n) (Fig.<br />

11.7).<br />

E<br />

I n<br />

od<br />

consump =<br />

Pr n-1<br />

(11.3)<br />

Unconsumed material eventually enters the<br />

detritus-based food chain as dead organic<br />

matter. On average, the quantity <strong>of</strong> food consumed<br />

by a given trophic level must be less than<br />

the production <strong>of</strong> the preceding trophic level,<br />

or the prey will be driven to extinction. In other<br />

words, the average consumption efficiency <strong>of</strong> a<br />

trophic link must be less than 100%. There may<br />

be times when the consumption by one trophic<br />

level exceeds that in the preceding level. Most<br />

vertebrate herbivores, for example, consume<br />

plants during winter, when there is no plant<br />

production. This is, however, typically <strong>of</strong>fset<br />

by other times, such as summer, when plants<br />

usually produce much more biomass than<br />

animals can consume. Situations in which consumption<br />

efficiency is greater than 100% for<br />

prolonged periods lead to dramatic <strong>ecosystem</strong><br />

changes. If predator control, for example, leads<br />

to a large deer population that consumes more<br />

plant biomass than is produced, the plant<br />

biomass will be reduced, altering plant species<br />

composition in ways that pr<strong>of</strong>oundly affect all<br />

<strong>ecosystem</strong> processes (see Chapter 12) (Pastor<br />

et al. 1988, Kielland and Bryant 1998, Paine<br />

2000). Sometimes this occurs naturally. Some<br />

herbivores, such as beavers, typically overexploit<br />

their local food supply and move to new<br />

areas when their food is depleted.<br />

The proportion <strong>of</strong> NPP consumed by herbivores<br />

varies at least 100-fold among <strong>ecosystem</strong>s,<br />

from less than 1% to greater than 40%<br />

(Table 11.1). The major factor accounting for<br />

this wide range in herbivore consumption efficiency<br />

is differences in plant allocation to structure.<br />

Herbivore consumption efficiency is<br />

generally lowest in forests (less than 1 to 5%),<br />

where there is a large plant allocation to wood.<br />

Herbivore consumption efficiencies are higher<br />

Plant-Based Trophic Systems 253<br />

Table 11.1. Consumption efficiency <strong>of</strong> the herbivore<br />

trophic level in selected <strong>ecosystem</strong> types.<br />

Consumption efficiency a<br />

Ecosystem type (% <strong>of</strong> aboveground NPP)<br />

Oceans 60–99<br />

Managed rangelands 30–45<br />

African grasslands 28–60<br />

Herbaceous old fields 5–15<br />

(1–7yr)<br />

Herbaceous old fields (30yr) 1.1<br />

Mature deciduous forests 1.5–2.5<br />

a Terrestrial estimates emphasize consumption by aboveground<br />

herbivores and may not accurately reflect the total<br />

<strong>ecosystem</strong>-scale consumption efficiency.<br />

Data from Wiegert and Owen (1971) and Detling (1988).<br />

in grasslands (10 to 60%), where most aboveground<br />

material is nonwoody, and highest<br />

(generally greater than 40%) in pelagic aquatic<br />

<strong>ecosystem</strong>s, where most plant (i.e., algal) biomass<br />

is cell contents rather than cell walls. In<br />

these <strong>ecosystem</strong>s, more algal biomass is <strong>of</strong>ten<br />

consumed by herbivores than dies and decomposes;<br />

this pattern contributes to inverted<br />

biomass pyramids (Fig. 11.8). In grasslands,<br />

consumption efficiencies are generally greater<br />

for <strong>ecosystem</strong>s dominated by large mammals<br />

(25 to 50%) than those dominated by insects<br />

and small mammals (5 to 15%) (Detling 1988).<br />

The toxic nature <strong>of</strong> some plant tissues (due to<br />

presence <strong>of</strong> plant secondary metabolites) and<br />

inaccessibility <strong>of</strong> other tissues (e.g., roots to<br />

aboveground herbivores) constrain the herbivore<br />

consumption efficiency <strong>of</strong> <strong>terrestrial</strong><br />

<strong>ecosystem</strong>s. Nematodes, one <strong>of</strong> the major<br />

belowground herbivores, consume 5 to 15%<br />

<strong>of</strong> belowground NPP in grasslands (Detling<br />

1988). Consumption efficiencies for belowground<br />

herbivores are not well documented,<br />

so whole-<strong>ecosystem</strong> estimates <strong>of</strong> consumption<br />

efficiencies almost always emphasize aboveground<br />

consumption. The highest consumption<br />

efficiencies in <strong>terrestrial</strong> <strong>ecosystem</strong>s are on<br />

grazing lawns, such as those found in some<br />

African savannas (McNaughton 1985) and<br />

arctic wetlands (Jefferies 1988). These highly<br />

productive grasslands are maintained as a lawn<br />

by repeated herbivore grazing. Nutrient inputs<br />

in urine and feces from these herbivores

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