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Browsing Faculty Publications by Author "Cooper, David J., author"
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Item Open Access Hydrologic regime and herbivory stabilize an alternative state in Yellowstone National Park(Colorado State University. Libraries, 2007) Wolf, Evan C., author; Cooper, David J., author; Hobbs, N. Thompson, author; Ecological Society of America, publisherA decline in the stature and abundance of willows during the 20th century occurred throughout the northern range of Yellowstone National Park, where riparian woody-plant communities are key components in multiple-trophic-level interactions. The potential causes of willow decline include climate change, increased elk browsing coincident with the loss of an apex predator, the gray wolf, and an absence of habitat engineering by beavers. The goal of this study was to determine the spatial and temporal patterns of willow establishment through the 20th century and to identify causal processes. Sampled willows established from 1917 to 1999 and contained far fewer young individuals than was predicted from a modeled stable willow population, indicating reduced establishment during recent decades. Two hydrologically distinct willow establishment environments were identified: fine grained beaver pond sediments and coarse-grained alluvium. Willows established on beaver pond sediment earlier in time, higher on floodplain surfaces, and farther from the current stream channel than did willows on alluvial sediment. Significant linear declines from the 1940s to the 1990s in alluvial willow establishment elevation and lateral distance from the stream channel resulted in a much reduced area of alluvial willow establishment. Willow establishment was not well correlated with climate-driven hydrologic variables, but the trends were consistent with the effects of stream channel incision initiated in ca. 1950, 20-30 years after beaver dam abandonment. Radiocarbon dates and floodplain stratigraphy indicate that stream incision of the present magnitude may be unprecedented in the past two millennia. We propose that hydrologic changes, stemming from competitive exclusion of beaver by elk over browsing, caused the landscape to transition from a historical beaver-pond and willow mosaic state to its current alternative stable state where active beaver dams and many willow stands are absent. Because of hydrologic changes in streams, a rapid return to the historical state may not occur by reduction of elk browsing alone. Management intervention to restore the historical hydrologic regime may be necessary to recover willows and beavers across the landscape.Item Open Access Relationships between groundwater use, water table, and recovery of willow on Yellowstone's northern range(Colorado State University. Libraries, 2011) Johnston, Danielle B., author; Cooper, David J., author; Hobbs, N. Thompson, author; Ecological Society of America, publisherExcessive levels of herbivory, incision of stream channels, and climate warming are believed to be responsible for the decline of woody deciduous plants in riparian zones in western North America, declines that are likely to be associated with diminished biological diversity. In the northern elk wintering range of Yellowstone National Park, USA, over browsing by elk (Cervus elaphus), lowered water tables resulting from stream incision, and loss of activity by beaver (Castor canadensis) have been implicated in the decline of willow (Salix sp.) communities. Reducing elk browsing appears sufficient for willow recovery in some areas, but where water table changes have been dramatic, recovery may be slow or absent. The importance of water table changes is disputed because experimental results demonstrate water table limitations, but water table depth has failed to explain variation in willow height at landscape scales. One explanation for this apparent discrepancy is that willows that have survived intensive browsing by elk have maintained access to groundwater despite declining water tables. Using stable isotopes of water, we examined the relationships between groundwater use, water table depth, and height of heavily browsed Salix geyeriana. Salix geyeriana groundwater use varied from 30% to 80%, and was higher later in the growing season, when soil water was less available and shoot water potentials were lower. Late season groundwater use explained 26% of the variation in total height of willows (P = 0.002), with taller plants using more groundwater. Water table depth explained only 8% of the variability in total height (P=0.051), with shorter willows having deeper water table depths. Groundwater use and water table depth were uncorrelated. Height recovery following a winter of heavy browsing was related to groundwater use, but not groundwater depth. We suggest that access to deeper water sources alleviates late season water stress, allowing for more rapid height recovery and higher total plant height. Variability in groundwater access may account for variability in height recovery at landscape scales.Item Open Access Water tables constrain height recovery of willow on Yellowstone's northern range(Colorado State University. Libraries, 2008) Bilyeu, Danielle M., author; Cooper, David J., author; Hobbs, N. Thompson, author; Ecological Society of America, publisherExcessive 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 (Castorcanadensis 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.