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Herbivores, the Functional Diversity of Plants Species, and the Cycling of Nutrients in Ecosystems

Numerous investigators have suggested that herbivores almost always increase rates of nutrient and energy flow through terrestrial ecosystems by returning to the soil fecal material and urine with faster turnover rate than shed plant litter. These previous theories and models always treat the producer compartment as a homogenous pool. Essentially, they assume that consumers feed through a pureed cream of vegetable soup. However, many field observations and experiments have shown that consumers feed selectively (i.e., in a cafeteria) and that consumer choice is made on the same chemical basis that determines decomposition rates. Plants that are preferred food sources often have higher nutrient content, higher growth rates, and faster decomposition rates. As consumption reduces dominance of these species in favor of unpreferred species with slower decomposition, rates of nutrient cycling and energy flow should therefore decline. We analyze a model in which the consumer is given a choice among producers that vary in nutrient uptake rates, rates of nutrient return to decomposers, and consumer preference, and which is parameterized for plants and consumers characteristic of boreal regions. In this model, in an open, well-mixed system with one consumer and two such producers, the nutrient/energy flow will not exceed that of a system without the consumer. If the consumer has a choice between two such producers, it must choose one plant over the other at a greater ratio than that between the two plants in uptake and decay rates. In contrast, in a closed system the consumer must be less selective to coexist with the two plants. The system behavior is determined by the level of nutrient return through the consumer and the differences between the plants in nutrient uptake rates and consumer preference. Species richness affects properties of this model system to the extent that species are functionally distinct (i.e., have different rate constants) in a multivariate space of life history traits (i.e., nutrient uptake and palatability). We suggest that the biochemical variability of plant tissues that simultaneously determines both consumer preference and decomposition rates is an essential feature of food webs that cannot be ignored. Thus, ecosystem models should, at minimum, consider more than one producer type with consumer preference.