Response of native versus exotic plant guilds to cattle and elk herbivory in forested rangeland

Questions: Are exotic plant species favoured by non-native ungulate herbivores and disadvantaged by native herbivores in forested rangelands? Do the impacts of ungulates on exotic vs native plants depend on forest management activities such as prescribed fire and stand thinning? Location: Northeastern Oregon, USA. Methods: We recorded changes in richness and cover of different exotic and native plant life forms in experimental plots that were grazed only by cattle (a non-native herbivore), only by elk (a native herbivore) or not grazed by any ungulate over a 7-yr period at both managed (recently burned and thinned) and unmanaged (where no fire and thinning has occurred in >40 yr) forest stands. Results: There was a general decrease in exotic plant species richness and cover across all treatments. However, the decrease in exotic richness, particularly of exotic annual forbs, was slightly lower in plots grazed by elk than in ungrazed plots at managed stands. Managed stands also displayed a larger increase in native annual forb richness and exotic graminoid richness, and a larger decrease in native perennial graminoid cover with cattle grazing than elk grazing. At unmanaged stands, cover of woody native plants such as shrubs, sub-shrubs and trees as well as native perennial forbs decreased or remained relatively constant with elk grazing while increasing strongly at plots that were ungrazed or grazed by cattle. Conclusions: Cattle and elk have variable effects on different plant guilds at managed vs unmanaged forest stands. Overall, cattle grazing tends to have a larger impact on herbaceous plant guilds at managed stands, while elk grazing tends to have a larger impact on woody plant guilds at unmanaged stands. However, in contrast to findings from other ecosystems, grazing only has a minor impact on exotic plant dynamics in our study area, and cattle grazing does not favour exotic plants any more than grazing by elk or ungulate exclusion.Keywords: Non-Native Species, Conditioned Correspondence Analysis, Conifer Forest Management, Grazing, Ungulates, Stand Thinning, Prescribed Burnin

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

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

Impact of ungulate exclusion on understorey succession in relation to forest management in the Intermountain Western United States

QUESTIONS: Do successional trajectories in plant diversity, heterogeneity and dominance respond differently to ungulate exclusion in unmanaged forests vs managed forests that are thinned and burned? Is vegetation in recently thinned and burned stands more sensitive to changes in the grazing regime? LOCATION: Northeast OR, USA. METHODS: We evaluated changes in plant community composition, diversity, heterogeneity and dominance under herbivory by multiple ungulates (cattle, elk, deer) vs ungulate exclusion at sites where trees were recently thinned and a prescribed burn was applied (managed), and in sites that were not thinned or burned in over 40 yr (unmanaged). Plant species diversity was calculated with the Simpsons index and richness as the total number of plant species. We estimated changes in plant community heterogeneity using a measure of taxonomic dissimilarity. Plant dominance was measured as the relative evenness among different plant functional groups (annual and perennial forbs and graminoids, and shrubs, subshrubs and trees). RESULTS: As expected, managed sites displayed more early succession species, such as annual forbs and annual graminoids,while unmanaged sites were dominated by late-succession species such as shrubs, subshrubs and trees. Species richness, particularly of annuals, was strongly reduced when ungulates were excluded from managed sites, and to a lesser extent from unmanaged sites for some perennial plant species. Species diversity decreased to a slightly greater extent with ungulate exclusion at managed sites. Species dominance was not influenced by ungulate exclusion. The effect of ungulate exclusion on plant heterogeneity also depended on forest management. Heterogeneity increased at managed sites and decreased in unmanaged sites with ungulate exclusion. Overall, the change in vegetation composition over time increased with the exclusion of ungulates, particularly at managed sites. CONCLUSIONS: The strength and direction of specific vegetation and diversity responses to ungulate exclusion vary with forest management, and the influence of ungulate exclusion on plant succession is more pronounced in recently thinned and burned sites. Management of wild and domestic ungulates thus needs to account for forest management activities that alter vegetation seral stage and increase the sensitivity of vegetation to the ungulate grazing regime.Keywords: Grazing, Stand thinning, Cattle, Conifer forest, Elk and deer herbivory, Prescribed burning, Taxonomic dissimilarityKeywords: Grazing, Stand thinning, Cattle, Conifer forest, Elk and deer herbivory, Prescribed burning, Taxonomic dissimilarit

Impacts of increased fire frequency and aridity on eucalypt forest structure, biomass and composition in southwest Australia

Water stress and fire disturbance can directly impact stand structure, biomass and composition by causing mortality and influencing competitive interactions among trees. However, open eucalypt forests of southwest Australia are highly resilient to fire and drought and may respond differently to increased fire frequency and aridity than forests dominated by non-eucalypt species. We measured the variation in stem density, basal area, stand biomass, sapwood area, leaf area and litterfall across 16 mixed jarrah (Eucalyptus marginata) and marri (Corymbia calophylla) forest stands along an aridity gradient in southwest Australia that had variable fire histories. Fire frequency was defined as the total number of fires over a ~30-year period and aridity as the ratio of potential evapotranspiration to annual precipitation. Total stand biomass and sapwood area were predicted from diameter at breast height of individual jarrah and marri trees using allometric equations. Leaf area was estimated using digital cover photography. More arid and frequently burnt stands had higher stem density, especially of smaller trees, which were mainly jarrah. Overall, both standing biomass and leaf area decreased at more arid sites, while sapwood area was largely unaffected by aridity, suggesting that these stands respond to increased water limitation by decreasing their leaf area relative to their sapwood area. Biomass of marri was reduced at more arid and, to a lesser extent, at more frequently burnt stands. However, total stand biomass (jarrah and marri) and leaf area index did not vary with fire frequency, suggesting that less marri biomass (due to slower growth rates, higher mortality or less recruitment) was compensated by an increase in the density of jarrah trees (regeneration). We conclude that increased fire and drought shift tree species composition towards more fire-resistant species and result in denser stands of smaller trees. In contrast, total stand biomass declines with increasing aridity, but has no association with fire frequency

Plant functional traits along environmental gradients in seasonally dry and fire-prone ecosystem

This paper investigates the questions “How does the abundance and richness of plant assemblages with different functional (regeneration and nutrient acquisition) traits vary with fire regime, moisture availability and substrate fertility? What is the role of different functional traits in maintaining plant diversity under changing environmental conditions in seasonally dry and fire-prone environments?

Response of plant species and life form diversity to variable fire histories and biomass in the Jarrah forest of south-west Australia

Frequent fires reduce the abundance of woody plant species and favour herbaceous species. Plant species richness also tends to increase with decreasing vegetation biomass and cover due to reduced competition for light.We assessed the influence of variable fire histories and site biomass on the following diversity measures: woody and herbaceous species richness, overall species richness and evenness, and life form evenness (i.e. the relative abundance or dominance among six herbaceous and six woody plant life forms), across 16 mixed jarrah (Eucalyptus marginata) and marri (Corymbia calophylla) forest stands in south-west Australia. Fire frequency was defined as the total number of fires over a 30-year period. Overall species richness and species evenness did not vary with fire frequency or biomass. However, there were more herbaceous species (particularly rushes, geophytes and herbs) where there were fewer shrubs and low biomass, suggesting that more herbaceous species coexist where dominance by shrubs is low. Frequently burnt plots also had lower number and abundance of shrub species. Life form evenness was also higher at both high fire frequency and low biomass sites. These results suggest that the impact of fire frequency and biomass on vegetation composition is mediated by local interactions among different life forms rather than among individual species. Our results demonstrate that measuring the variation in the relative diversity of different woody and herbaceous life forms is crucial to understanding the compositional response of forests and other structurally complex vegetation communities to changes in disturbance regime such as increased fire frequency

Plant diversity is linked to nutrient limitation of dominant species in a world biodiversity hotspot

This paper investigates the question “What is the role of nutrient limitation of dominant species in maintaining plant species diversity at small spatial scales, particularly in biodiversity hotspots?”

Ungulate browsing maintains shrub diversity in the absence of episodic disturbance in seasonally-arid conifer forest.

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

EndressBryanAnimalRangelandSciResponseNativeVersus.pdf

Questions: Are exotic plant species favoured by non-native ungulate herbivores and disadvantaged by native herbivores in forested rangelands? Do the impacts of ungulates on exotic vs native plants depend on forest management activities such as prescribed fire and stand thinning? Location: Northeastern Oregon, USA. Methods: We recorded changes in richness and cover of different exotic and native plant life forms in experimental plots that were grazed only by cattle (a non-native herbivore), only by elk (a native herbivore) or not grazed by any ungulate over a 7-yr period at both managed (recently burned and thinned) and unmanaged (where no fire and thinning has occurred in >40 yr) forest stands. Results: There was a general decrease in exotic plant species richness and cover across all treatments. However, the decrease in exotic richness, particularly of exotic annual forbs, was slightly lower in plots grazed by elk than in ungrazed plots at managed stands. Managed stands also displayed a larger increase in native annual forb richness and exotic graminoid richness, and a larger decrease in native perennial graminoid cover with cattle grazing than elk grazing. At unmanaged stands, cover of woody native plants such as shrubs, sub-shrubs and trees as well as native perennial forbs decreased or remained relatively constant with elk grazing while increasing strongly at plots that were ungrazed or grazed by cattle. Conclusions: Cattle and elk have variable effects on different plant guilds at managed vs unmanaged forest stands. Overall, cattle grazing tends to have a larger impact on herbaceous plant guilds at managed stands, while elk grazing tends to have a larger impact on woody plant guilds at unmanaged stands. However, in contrast to findings from other ecosystems, grazing only has a minor impact on exotic plant dynamics in our study area, and cattle grazing does not favour exotic plants any more than grazing by elk or ungulate exclusion.Keywords: Non-Native Species, Conifer Forest Management, Conditioned Correspondence Analysis, Prescribed Burning, Grazing, Stand Thinning, Ungulate

Mixed model results showing effect of ungulate browsing on shrub species diversity and evenness at non-fuels treated and fuels treated sites.

<p>The intercept (<i>Int.</i>), coefficients, estimate (<i>Est.</i>) and standard error (<i>SE</i>), and P-value (<i>P</i>) are shown for each model. The degrees of freedom is 4 for non-fuels treated sites and 8 for fuels treated sites. P-values indicate significance of difference in the mean response between each browsing pressure group from non-browsed paddocks as illustrated in <a href="http://www.plosone.org/article/info:doi/10.1371/journal.pone.0086288#pone-0086288-g004" target="_blank">Figure 4</a>.</p