Water tables constrain height recovery of willow on Yellowstone's northern range.

Excessive levels of herbivory may disturb ecosystems in ways that persist even when herbivory is moderated. These persistent changes may complicate efforts to restore ecosystems affected by herbivores. Willow (Salix spp.) communities within the northern range in Yellowstone National Park have been eliminated or degraded in many riparian areas by excessive elk (Cervus elaphus L.) browsing. Elk browsing of riparian willows appears to have diminished following the reintroduction of wolves (Canis lupis L.), but it remains uncertain whether reduced herbivory will restore willow communities. The direct effects of elk browsing on willows have been accompanied by indirect effects from the loss of beaver (Castor canadensis Kuhl) activity, including incision of stream channels, erosion of fine sediments, and lower water tables near streams historically dammed by beaver. In areas where these changes have occurred, lowered water tables may suppress willow height even in the absence of elk browsing. We conducted a factorial field experiment to understand willow responses to browsing and to height of water tables. After four years of protection from elk browsing, willows with ambient water tables averaged only 106 cm in height, with negligible height gain in two of three study species during the last year of the experiment. Willows that were protected from browsing and had artificially elevated water tables averaged 147 cm in height and gained 19 cm in the last year of the experiment. In browsed plots, elevated water tables doubled height gain during a period of slightly reduced browsing pressure. We conclude that water availability mediates the rate of willow height gain and may determine whether willows grow tall enough to escape the browse zone of elk and gain resistance to future elk browsing. Consequently, in areas where long-term beaver absence has resulted in incised stream channels and low water tables, a reduction in elk browsing alone may not be sufficient for recovery of tall willow stands. Because tall willow stems are important elements of habitat for beaver, mitigating water table decline may be necessary in these areas to promote recovery of historical willow-beaver mutualisms.