Ungulate populations are managed in shrubsteppe ecosystems around the world, but relatively few long-term datasets are available that test the impacts of these grazers on shrubsteppe structure and function. This study evaluated 8 exclosures in 4 shrubsteppe communities to determine the effects of deer and elk over 35 or 55 years on (1) plant biomass and species composition, (2) soil nitrogen (N) mineralization and net nitrification rates, Olsen extractable phosphorus (P), and C:N ratios, and (3) arthropod diversity and abundance. The site with deer, the highest ungulate densities, and coldest climate (Sinlahekin Wildlife Area, WA) had greater vegetative biomass and cover inside the exclosures than outside. The sites with elk, moderate ungulate densities, and intermediate climatic conditions (Oak Creek and Wenas Wildlife Areas, WA) had no effects of exclosures on vegetative biomass or cover. The site with elk, the lowest ungulate densities, and driest climatic conditions (L.T. Murray Wildlife Area, WA) had greater shrub cover outside of the exclosures. Ungulates significantly increased net N mineralization and nitrification rates, providing a potential explanation for compensatory growth by grazed plants. Our data suggest that arthropod herbivory does not substitute for ungulate herbivory. To the contrary, arthropod diversity reflected ungulate-induced changes in plant biomass. In addition to effects on standing biomass, ungulates were associated with increased exotic plant species richness at all 4 sites. Our results suggest that there may be a balance between indirect positive and direct negative effects of herbivory on plants that varies with ungulate densities and site characteristics, and that arthropod herbivory does not functionally replace ungulate herbivory.
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Vol. 67 • No. 1