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      Ungulate Browsing Maintains Shrub Diversity in the Absence of Episodic Disturbance in Seasonally-Arid Conifer Forest

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          Abstract

          Ungulates exert a strong influence on the composition and diversity of vegetation communities. However, little is known about how ungulate browsing pressure interacts with episodic disturbances such as fire and stand thinning. We assessed shrub responses to variable browsing pressure by cattle and elk in fuels treated (mechanical removal of fuels followed by prescribed burning) and non-fuels treated forest sites in northeastern Oregon, US. Seven treatment paddocks were established at each site; three with cattle exclusion and low, moderate and high elk browsing pressure, three with elk exclusion and low, moderate and high cattle browsing pressure, and one with both cattle and elk exclusion. The height, cover and number of stems of each shrub species were recorded at multiple plots within each paddock at the time of establishment and six years later. Changes in shrub species composition over the six year period were explored using multivariate analyses. Generalized Linear Mixed Models were used to determine the effect of browsing pressure on the change in shrub diversity and evenness. Vegetation composition in un-browsed paddocks changed more strongly and in different trajectories than in browsed paddocks at sites that were not fuels treated. In fuels treated sites, changes in composition were minimal for un-browsed paddocks. Shrub diversity and evenness decreased strongly in un-browsed paddocks relative to paddocks with low, moderate and high browsing pressure at non-fuels treated sites, but not at fuels treated sites. These results suggest that in the combined absence of fire, mechanical thinning and ungulate browsing, shrub diversity is reduced due to increased dominance by certain shrub species which are otherwise suppressed by ungulates and/or fuels removal. Accordingly, ungulate browsing, even at low intensities, can be used to suppress dominant shrub species and maintain diversity in the absence of episodic disturbance events.

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          Divergent nonlinear responses of the boreal forest field layer along an experimental gradient of deer densities.

          The early responses of the field layer to changes in biotic and abiotic conditions are key determinants of the future composition and structure of forests where sustained heavy browsing pressure has depauperated the shrub understory. We investigated the relationships between white-tailed deer density and field layer plant community dynamics in boreal forests managed for wildlife and timber production. We hypothesized that the growth and reproduction of field layer plants are either: (H(1)) directly proportional to deer density, or (H(2)) related to deer density through nonlinear relationships or (H(3)) through nonlinear relationships with thresholds. We tested these hypotheses using data from a controlled browsing experiment involving a gradient of deer densities (0, 7.5, 15, 27 and 56 deer km(-2)) in interaction with timber harvesting conducted on Anticosti Island, Canada. In recent clearcuts, the dominant responses of the field layer plants were exponential recovery in growth and reproduction with decreasing deer densities. The abundance of browse-tolerant species such as grasses was positively related to deer density, suggesting an apparent competitive gain. These results support the prediction from our second hypothesis, although the presence of ecological thresholds should not be ruled out. Rapid changes in the early successional stages have potentially long-term consequences on successional patterns through processes such as the modulation of germination and early establishment success of seedlings from later successional species. Quantitative data as those presented here are essential for the development of ecosystem management prescriptions. On Anticosti Island, reduction of local deer densities to levels <15-7.5 deer km(-2) in the first 3 years following timber harvesting appears to be compatible with the regeneration dynamics of this system although lower levels of deer densities may be required for the conservation of browse-sensitive plant species.
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            Large herbivores influence the composition and diversity of shrub-steppe communities in the Rocky Mountains, USA.

            It is widely believed that wild and domestic herbivores have modified the structure and composition of arid and semi-arid plant communities of western North America, but these beliefs have rarely been tested in long-term, well-replicated studies. We examined the effects of removing large herbivores from semi-arid shrublands for 40-50 years using 17 fenced exclosures in western Colorado, USA. Shrub cover was greater (F=5.87, P=0.0020) and cover (F=3.01, P=0.0601) and frequency (F=3.89, P=0.0211) of forbs was less inside the exclosures (protected) relative to grazed plots. However, we found no significant effects (minimum P=0.18) of protection from grazing on cover or frequency of grasses, biotic crusts, or bare soil. Although mean species richness and diversity were similar between treatments, protected areas had much higher dominance by fewer species, primarily sagebrush. Exclusion of herbivores changed the relationship between species richness and evenness. Consistent with theoretical expectations, species evenness was positively correlated with richness in protected plots (r2=0.54). However, contrary to theory, evenness and richness were inversely related in grazed plots (r2adjacent=0.72, r2distant=0.84). We suggest that these differences resulted because grazing acts as a stressor promoting facilitative relationships between plant species that might compete for resources in the absence of grazing. We conclude that exclusion of grazing in the sites we studied caused minor changes in cover and diversity of herbaceous plants, but caused a clear increase in the cover of shrubs. Importantly, the exclusion of ungulates changed the relationship between evenness and richness.
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              Pervasive interactions between ungulate browsers and disturbance regimes promote temperate forest herbaceous diversity.

              Disruptions to historic disturbance and herbivory regimes have altered plant assemblages in forests worldwide. An emerging consensus suggests that these disruptions often result in impoverished forest biotas. This is particularly true for eastern U.S. deciduous forests where large gaps and understory fires were once relatively common and browsers were far less abundant. Although much research has focused on how disturbance and browsers affect tree diversity, far less attention has been devoted to forest understories where the vast majority (>75%) of the vascular species reside. Here we test the hypothesis that the reintroduction of disturbances resembling historic disturbance regimes and moderate levels of ungulate browsing enhance plant diversity. We explore whether once-common disturbances and their interaction with the top-down influence of browsers can create conditions favorable for the maintenance of a rich herbaceous layer in a region recognized as a temperate biodiversity hotspot in West Virginia, U.S.A. We tested this hypothesis via a factorial experiment whereby we manipulated canopy gaps (presence/absence) of a size typically found in old-growth stands, low-intensity understory fire (burned/unburned), and deer browsing (fenced/unfenced). We tracked the abundance and diversity of more than 140 herb species for six years. Interactions among our treatments were pervasive. The combination of canopy gaps and understory fire increased herbaceous layer richness, cover, and diversity well beyond either disturbance alone. Furthermore, we documented evidence that deer at moderate levels of abundance promote herbaceous richness and abundance by preferentially browsing fast-growing pioneer species that thrive following co-occurring disturbances (i.e., fire and gaps). This finding sharply contrasts with the negative impact browsers have when their populations reach levels well beyond those that occurred for centuries. Although speculative, our results suggest that interactions among fire, canopy gaps, and browsing provided a variable set of habitats and conditions across the landscape that was potentially capable of maintaining much of the plant diversity found in temperate forests.
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                Author and article information

                Contributors
                Role: Editor
                Journal
                PLoS One
                PLoS ONE
                plos
                plosone
                PLoS ONE
                Public Library of Science (San Francisco, USA )
                1932-6203
                2014
                23 January 2014
                : 9
                : 1
                : e86288
                Affiliations
                [1 ]Division of Applied Plant Ecology, Institute for Conservation Research, San Diego Zoo Global, Escondido, California, United States of America
                [2 ]Pacific Northwest Research Station, USDA Forest Service, La Grande, Oregon, United States of America
                [3 ]Department of Forest Ecosystems and Society, Oregon State University, Corvallis, Oregon, United States of America
                University of Tartu, Estonia
                Author notes

                Competing Interests: The authors have declared that no competing interests exist.

                Conceived and designed the experiments: MJW BAE CGP. Performed the experiments: MJW BAE BJN. Analyzed the data: BKP. Contributed reagents/materials/analysis tools: BKP MJW BAE BJN CGP. Wrote the paper: BKP MJW BAE.

                Article
                PONE-D-13-30652
                10.1371/journal.pone.0086288
                3900502
                ae8a0ae6-1f6b-4c04-9bcf-f5b197bf0cde
                Copyright @ 2014

                This is an open-access article, free of all copyright, and may be freely reproduced, distributed, transmitted, modified, built upon, or otherwise used by anyone for any lawful purpose. The work is made available under the Creative Commons CC0 public domain dedication.

                History
                : 24 July 2013
                : 13 December 2013
                Page count
                Pages: 9
                Funding
                Research was funded by the United States Department of Agriculture (USDA) Forest Service Pacific Northwest Research Station, Oregon State University, and San Diego Zoo's Institute for Conservation Research. The funders had no role in study design, data collection and analysis, decision to publish, or preparation of the manuscript.
                Categories
                Research Article
                Agriculture
                Forestry
                Sustainable Agriculture
                Biology
                Ecology
                Ecological Metrics
                Maximum Sustainable Yield
                Optimum Sustainable Yield
                Plant Ecology
                Plant-Environment Interactions
                Biodiversity
                Plant Science
                Plant Ecology
                Plant-Environment Interactions
                Engineering
                Fire Engineering
                Fire Research

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                Uncategorized

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