© 2006 The Authors 294 Journal compilation © 2006 The Royal Entomological Society Interactions between cottonwood and beavers

Abstract. 1. Cottonwood ( Populus spp.) are the dominant tree type in riparian forests of the western U.S.A. In these riparian forests, the beaver ( Castor canadensis) is a major ecosystem engineer that commonly browses cottonwood, resulting in distinct changes to plant architecture. Here the hypothesis that beaver herbivory indirectly affects the distribution of a keystone leaf-galling sawfly through architectural changes in cottonwood was examined. 2. It was found that: (a) beaver herbivory of cottonwood results in an increase in average shoot length over unbrowsed cottonwood; (b) sawfly galls were up to 7 – 14 times more abundant on browsed cottonwood than unbrowsed cottonwood; and (c) sawfly gall abundance was correlated positively with changes in shoot length after beaver herbivory. Together these data show that the individual and combined effects of cottonwood and beaver herbivory increase shoot length, positively affecting sawfly abundance. 3. Because herbivores are a ubiquitous component of most ecosystems, we argue that the indirect effects of herbivory on plant quality, and subsequently other herbivores, may be as important as environmental variation. Key words. Beaver, herbivores, Populus, sawfl y, trait-mediated indirect effects