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Effects of unmanaged populations of large mammalian herbivores, especially elk (Cervus elaphus on vegetation is a concern in Yellowstone National Park, since wolves (Canis Lupus) are extirpated, ungulate migrations are altered by human activities and the disruption of natural process is possible. Stands of low, hedged (height-suppressed) willows (Salix spp.) are observed throughout the greater Yellowstone National Park area where high densities of wintering elk or moose (Alces alces) exist. The height of 47% of the willow stands surveyed on Yellowstone's northern winter range has been suppressed. Mean leader use of willows of all heights was (P < 0.05 in the winter of 1987-88, increased to 60% in winter 1988-89, following the drought and fires of 1988, then declined to 44% in 1989-90 and winter 1990-96. Height-suppressed willows (43 +/- 2 cm, mean +/- SE) were about one-half as tall as tall willows (83 +/- 4 cm). Percent twig use of suppressed willows in summer (25%) and winter (59%) was significantly more than for intermediate or tall stands (P < 0.05). Suppressed willows produced about one-fourth the aboveground annual biomass compared to taller willows; even after 27 or 31 years of protection, previously-suppressed willows produced only one-third the aboveground biomass of taller willows, suggesting suppressed willows grow on sites with lower growth potential. Growth conditions for willows on the northern winter range may have declined due to a warmer and drier climate this century, locally reduced water tables--because of the decline on beaver (Castor canadenis), or fire suppression may be responsible for the observed changes. Tall and intermediate-height willows contained higher concentrations of nitrogen and they exhibited more water stress than height-suppressed willows of the same species. More xeric growth conditions this century than last century, especially during the decades of the 1920's, 1930's, and 1980's, may explain the low growth rates and lower chemical defenses against ungulate herbivory for height-suppressed willows. We propose a more xeric climate and locally-reduced water tables likely contributed to the willow declines on the northern winter range, but that the proximate factor in the declines was herbivory by native ungulates.

Response to Comment: Ungulate Herbivory on Willows on Yellowstone's Northern Winter Range