Effects of vertebrate herbivores and shrub characteristics on arthropod assemblages in a northern Arizona forest ecosystem

Abstract

Large herbivores have potential to affect invertebrate community structure through numerous processes, but little work has been done to evaluate the relative importance of direct and indirect factors. In this study, we measured arthropod community assemblages on Ceanothus fendleri A. Gray (Fendler's ceanothus) plants that were growing inside and outside of 4-m2 exclosures. We used univariate analyses and structural equation modeling (SEM) to evaluate relationships within this herbivore–plant–arthropod system in ponderosa pine (Pinus ponderosa Laws.) forests of northern Arizona, USA. Results showed that individual arthropod abundance, family diversity, family richness, and functional group richness were significantly greater on plots where C. fendleri plants were protected from large ungulate herbivores (e.g., mule deer (Odocoileus hemionus) and Rocky Mountain elk (Cervus elaphus nelsoni)) than on unprotected plots in each of the three study years. Results also indicated the following: (1) arthropod abundance was significantly greater on protected plants than unprotected plants; (2) rarefaction curves suggested arthropod family richness was similar between protected and unprotected plants in two of the three years when scaled by number of individuals but the estimated total richness was consistently higher on protected plants; (3) arthropod abundance was directly affected by protection from herbivores, plant stem length, and number of flowers; (4) arthropod family richness was related to the number of individuals collected and affected by stem length. Results from this study illustrate that arthropod communities are directly affected by foraging vertebrate herbivores as well as indirectly affected through complex plant-mediated factors in this model system. Protection of preferred forage plants such as C. fendleri from ungulate herbivores can potentially increase diversity of arthropod assemblages in these forests, help conserve biological diversity, and enhance ecosystem restoration efforts.