J. Hall Cushman’s research while affiliated with University of Nevada, Reno and other places

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Publications (13)


Standing litter modifies top-down effects of large herbivores on a grassland plant community
  • Article

August 2023

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74 Reads

Ecosystems

Zhongnan Wang

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Zhiwei Zhong

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James F Cahill

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[...]

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Large herbivores can exert pronounced top-down effects on plant communities in grassland ecosystems. Previous studies highlighted the importance of the composition and traits of living plants in regulating the impact of herbivores on plant community. However, there has been little consideration of whether and how plant litter, a ubiquitous ''after-life'' plant component, affects the outcome of herbivore grazing on grasslands. Here, we conducted a large-scale field experiment in temperate grasslands of northeastern China to investigate how standing plant litter influenced top-down effects of large herbivores (sheep; Ovis aries) on plant species richness, evenness, community composition , and productivity. We found that, in the presence of standing litter, sheep grazing significantly reduced living biomass of forbs by 56%, but have no effects on biomass of the dominant grass, Leymus chinensis. However, in the absence of standing litter, sheep shifted their diet preference from forbs to the grass L. chinensis, leading to a 36% decrease in the biomass of L. chinensis and a 21% decrease in total biomass. Such changes in foraging pressure on plant species led to competitive release that in turn significantly altered plant community composition and increased species evenness. Synthesis and applications. Our results demonstrate that standing litter can alter foraging behaviors of large herbivores and modifying the outcome of their top-down effects on plant community properties in grasslands. These cryptic but perhaps ubiquitous interactions between litter and herbivores may help us better understand the organization and dynamics of plant communities in the grazed grasslands, with important implications for developing effective management and conservation plans.


Standing plant litter, primarily from the dominant L. chinensis grass, is common and is retained until the early growing seasons (for example, from late May (A) to early July (B)) of following year, potentially affecting the foraging behaviors of sheep (Ovis aries) and thus their top-down effects on plant community in semiarid grasslands of northeastern China. The two photographs were provided by Zhongnan Wang.
Combined effects of 3-year sheep grazing and standing litter removal treatments on A total plant biomass, BL. chinensis biomass, C forb biomass, and D biomass of other grasses in the 5 × 5 m subplots in August 2012. For A–C, an asterisk (*) between the bars indicates significant (P < 0.05) differences based on post-hoc comparisons, whereas ns indicates non-significant differences between treatments. No interactive effects were detected for biomass of other grasses (D); see Results for details. Error bars represent ± 1 SE.
Combined effects of 3-year sheep grazing and standing litter removal treatments on plant A species richness, and B species evenness in the 5 × 5 m subplots in August 2012. For B, an asterisk (*) between the bars indicates significant (P < 0.05) differences based on post-hoc comparisons, whereas ns indicates non-significant differences between treatments. No interactive effects were detected for species richness (A); see Results for details. Error bars represent ± 1 SE.
Principal component analysis (PCA) of variation in the plant community composition of the four treatments in the 5 × 5 m subplots in August 2012. n = 24. UG = Ungrazed, G = Grazed, LI = Litter intact, LR = Litter removed.
Effects of 3-year standing litter removal treatments on grazing frequency of sheep on AL. chinensis grasses, B forbs, and C other grasses in the 5 × 5 m subplots in August 2012. Asterisk (*) between the bars indicates significant differences, whereas ns indicates non-significant differences between treatments. Error bars represent ± 1 SE.
Standing Litter Modifies Top-Down Effects of Large Herbivores on a Grassland Plant Community
  • Article
  • Publisher preview available

August 2023

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145 Reads

Ecosystems

Large herbivores can exert pronounced top-down effects on plant communities in grassland ecosystems. Previous studies highlighted the importance of the composition and traits of living plants in regulating the impact of herbivores on plant community. However, there has been little consideration of whether and how plant litter, a ubiquitous “after-life” plant component, affects the outcome of herbivore grazing on grasslands. Here, we conducted a large-scale field experiment in temperate grasslands of northeastern China to investigate how standing plant litter influenced top-down effects of large herbivores (sheep; Ovis aries) on plant species richness, evenness, community composition, and productivity. We found that, in the presence of standing litter, sheep grazing significantly reduced living biomass of forbs by 56%, but have no effects on biomass of the dominant grass, Leymus chinensis. However, in the absence of standing litter, sheep shifted their diet preference from forbs to the grass L. chinensis, leading to a 36% decrease in the biomass of L. chinensis and a 21% decrease in total biomass. Such changes in foraging pressure on plant species led to competitive release that in turn significantly altered plant community composition and increased species evenness. Synthesis and applications. Our results demonstrate that standing litter can alter foraging behaviors of large herbivores and modifying the outcome of their top-down effects on plant community properties in grasslands. These cryptic but perhaps ubiquitous interactions between litter and herbivores may help us better understand the organization and dynamics of plant communities in the grazed grasslands, with important implications for developing effective management and conservation plans.

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Ungulate herbivores as drivers of aspen recruitment and understory composition throughout arid montane landscapes

September 2022

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63 Reads

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3 Citations

Herbivory by wild and domestic ungulates can influence tree recruitment and understory forest communities throughout the world. Herbivore‐driven declines in tree recruitment have been observed for quaking aspen (Populus tremuloides), a foundation species whose health and management is recognized as a critical priority throughout much of its range. Livestock fencing is commonly used to promote aspen regeneration, but its effectiveness is rarely assessed, especially across large spatial scales. Using a livestock‐reduction experiment, we evaluated the effects of ungulate herbivory on aspen in the Great Basin and southern Cascades, an expansive and environmentally heterogeneous region where aspen faces the interacting threats of climate change, conifer encroachment, and herbivory. We found that livestock fencing only reduced the intensity of herbivore browsing on aspen when wild ungulate abundance was low and did not increase stem densities of aspen recruits. Contrary to expectations, wild ungulate abundance was a strong driver of browsing intensity on juvenile aspen within fenced, but not unfenced, aspen stands, and when the abundance of these herbivores was high, browsing intensity in fenced stands exceeded that in unfenced stands. The density of aspen recruits decreased with browsing intensity on juvenile aspen and with the density of both adult aspen and conifers, suggesting that both herbivory and intra‐ and interspecific competition are important drivers of recruitment. Fire history was also an important driver of recruitment, with stands that burned 10–20 years ago having the greatest density of aspen recruits. Finally, in the stand understory, we found that livestock fencing decreased forb cover, increased shrub species richness, and increased the cover of exotic annual grasses, a group dominated by Bromus tectorum. This latter finding suggests that livestock fencing may not be appropriate in areas where controlling the spread of this invader is a priority. In sum, our findings indicate that aspen recruitment is limited by browsing by both wild and domestic ungulates, is mediated by competition with neighboring trees and fire history, and will require management actions beyond livestock fencing, as this approach does not control browsing by wild ungulates.


Large herbivores facilitate a dominant grassland forb via multiple indirect effects

March 2022

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275 Reads

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13 Citations

While large herbivores are critically important components of terrestrial ecosystems and can have pronounced top‐down effects on plants, our understanding of the underlying mechanisms driving these effects remains incomplete. Large herbivores can alter plant growth, reproduction, and abundance through direct effects (predominantly consumption) and through indirect effects via altered interactions with abiotic factors and other species. We know considerably less about these indirect effects than the direct effects. Here, we integrate medium‐ and small‐scale field experiments to investigate how a large vertebrate herbivore, cattle (Bos taurus), affects the aboveground biomass of a dominant forb species, Artemisia scoparia, via diverse direct and indirect pathways in a temperate grassland in northeast China. Although cattle consumed this forb, its biomass increased significantly in response to grazing, due to multiple indirect positive effects that outweighed the direct negative effects of consumption. Cattle preferentially consumed the competing grass Leymus chinensis, and altered Artemisia microhabitats by reducing total plant cover and litter biomass and by increasing the abundance of co‐occurring ant species (e.g., Formica spp. and Lasius spp.). This led to additional indirect positive effects on A. scoparia likely due to (1) increased light availability in understory layers and other limiting resources (e.g., soil nutrients and moisture) caused by removal of competitors and plant litter at the soil surface and (2) the changes in resource availability (e.g., soil nutrients and moisture) associated with ant colonies. Our results show that large herbivores can affect plant growth not only via direct consumption, but also via multiple indirect effects. Focusing on the causes and consequences of herbivore‐induced indirect effects will not only help us to better understand the influence of these animals in ecological systems, but will also lead to more effective land management and conservation practices in the regions they inhabit.


a Spatial distribution of inventory plots (n = 3962) containing quaking aspen (P. tremuloides), colored by biome (Bailey 1998). The background gray color represents the geographic range of aspen (Little 1971). The six panels show modeled trends in mean (± 95% CI) annual mortality rate and relative change in live basal area (BA) of mature (dbh ≥ 12.7 cm) aspen stems by b, e biome, c, f initial mean diameter of aspen, and d, g initial relative abundance of aspen. Horizontal lines in e, f, and g represent no net change in live BA of mature aspen through time. Plots contained at least five live mature aspen stems and were remeasured 2–6 times between 1985–2018. For stem size- and composition-dependent models, a more recent subset of years (1999–2018) was used to ensure trends were representative of all biomes. Sample sizes: boreal (n = 859), montane coniferous (n = 874), prairie parkland (n = 122), mixed (n = 1846), and deciduous (n = 261)
a Total number of published studies by ecoregion (
adapted from Wiken et al. 2011) on the drivers of quaking aspen (P. tremuloides) mortality, growth and recruitment. b Total number of studies published per year by biome(s) over the past thirty years (1989–2019) on the drivers of quaking aspen mortality, growth and recruitment. All studies of deciduous forests also included other biomes, so these articles are instead included in the ‘multiple biomes’ study count. Ecoregions are labeled as follows: (b1) Alaska interior, (b2) boreal cordillera, (b3) boreal plains, (b4) boreal shield, (t1) taiga plains, (t2) western taiga shield, (t3) Hudson Plain, (t4) eastern taiga shield, (p1) prairie parkland, (c1) Canadian Rockies, (c2) Cascades, (c3) Idaho Batholith, (c4) Snake River Plain, (c5) Sierra Nevada, (c6) Great Basin, (c7) Arizona/New Mexico Mountains, (c8) Middle Rockies, (c9) Wasatch and Uinta Mountains, (c10) Colorado Plateaus, (c11) Southern Rockies, (m1) Northern Great Lakes and Southern Laurentians, (m2) Atlantic Highlands, (d1) interior and coastal lowlands, and (d2) Appalachian Plateau
Number of independent, published studies in the past thirty years (1989–2019) that reported a significant increase or decrease in a mortality, b growth, and c recruitment of quaking aspen (P. tremuloides) as a function of several abiotic and biotic stressors and endogenous factors
Modeled trends in mean (± 95% CI) a) stem density and b) basal area of live, mature (dbh ≥ 12.7 cm) trees of all species by biome from linear mixed models. Plots contained at least five live mature aspen stems and were remeasured 2–6 times between 1985–2018. Sample sizes: boreal (n = 859), montane coniferous (n = 874), prairie parkland (n = 122), mixed (n = 1846), and deciduous (n = 261)
Continent-wide synthesis of the long-term population dynamics of quaking aspen in the face of accelerating human impacts

September 2021

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152 Reads

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16 Citations

Oecologia

In recent decades, climate change has disrupted forest functioning by promoting large-scale mortality events, declines in productivity and reduced regeneration. Understanding the temporal dynamics and spatial extent of these changes is critical given the essential ecosystem services provided by forests. As the most widespread tree species in North America, quaking aspen (Populus tremuloides) is well suited for studying the dynamics of tree populations during a period of unprecedented climate change. Synthesizing continent-wide data, we show that mortality rates of mature aspen stems have increased over the past two-to-three decades, while relative gains in aspen basal area have decreased during the same period. Patterns were pervasive across multiple stand size classes and composition types in western North America biomes, suggesting that trends in demographic rates were not simply a reflection of stand development and succession. Our review of the literature revealed that increased aspen mortality and reduced growth rates were most often associated with hotter, drier conditions, whereas reduced recruitment was most often associated with herbivory. Furthermore, interactions between climate and competition, as well as climate and insect herbivory, had important, context-dependent effects on mortality and growth, respectively. Our analyses of aspen across its entire geographic range indicate that this important tree species is experiencing substantial increases in mortality and decreases in population growth rates across multiple biomes. If such trends are not accompanied by increased recruitment, we expect that the reduced dominance of aspen in forests will lead to major declines in the many essential ecosystem services it provides.


Effects of human-altered landscapes on a reintroduced ungulate: Patterns of habitat selection at the rangeland-wildland interface

May 2021

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125 Reads

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13 Citations

Biological Conservation

Successful species reintroductions require land managers to balance the goal of viable wildlife populations with potential risks to human enterprise. Such risks are particularly acute at the wildland-agriculture interface, where native and domestic species are likely to come into contact. In a national park in northern California, we combined insights from three lines of evidence – long-term visual surveys, short-term GPS telemetry, and satellite remote sensing-based animal detections – to characterize spatial overlap between reintroduced tule elk (Cervus canadensis nannodes) and domestic cattle and to estimate the importance of multiple environmental features as predictors of habitat selection by elk. Our results indicate that, at large spatial scales (i.e., home-range level), cattle were the primary driver of habitat selection, with the occurrence of elk being negatively associated with cattle across all seasons. In addition, elk consistently selected for grasslands on gentle, south-facing slopes that occurred at high elevation and close to ponds. NDVI was a seasonally important, positive predictor of habitat selection, with a marked reversal when this resource was concentrated inside of fenced cow pastures during the dry summer months. By contrast, a novel analysis of satellite-derived animal locations yielded no evidence of avoidance of cattle by elk (within pasture areas commonly used by elk), indicating that this population has acclimated to the presence of cattle through spatial partitioning of resources. Thus, this once-imperiled native ungulate exhibits patterns of habitat selection that reduce the potential for grazing conflicts with cattle, even in cases where access to forage is limited.


Plant functional groups and species contribute to ecological resilience a decade after woodland expansion treatments

January 2021

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64 Reads

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21 Citations

Woody plant expansions are altering ecosystem structure and function, as well as fire regimes, around the globe. Tree‐reduction treatments are widely implemented in expanding woodlands to reduce fuel loads, increase ecological resilience, and improve habitat, but few studies have measured treatment outcomes over long timescales or large geographic areas. The Sagebrush Treatment Evaluation Project (SageSTEP) evaluated the ecological effects of prescribed fire and cut‐and‐leave treatments in sagebrush communities experiencing tree expansion in North American cold desert shrublands. We used 10 yr of data from the SageSTEP network to test how treatments interacted with pre‐treatment tree dominance, soil climate, and time since treatment to affect plant functional groups and dominant species. Non‐sprouting shrub (Artemisia spp.), sprouting shrub, perennial graminoid, and annual grass responses depended on tree dominance and soil climate, and responses were related to the dominant species' life‐history traits. Sites with warm and dry soils showed increased perennial graminoid but reduced Artemisia shrub cover across the tree dominance gradient after prescribed burning, while sites with cool and moist soils showed favorable post‐burn responses for both functional types, particularly at low to moderate tree dominance. Cut‐and‐leave treatments sustained or increased native perennial plant functional groups and experienced smaller increases in exotic annual plants in both soil climates across the tree dominance gradient. Both treatments reduced biocrust cover. Selecting appropriate tree‐reduction treatments to achieve desired long‐term outcomes requires consideration of dominant species, site environmental conditions, and the degree of woodland expansion. Careful selection of management treatments will reduce the likelihood of undesirable consequences to the ecosystem.


Mean (±1 SE) bulk density (a), compaction depth (b), infiltration rate (c), and pH (d) as a function of elk and soil formation. Letters above bars correspond to the results from Tukey multiple comparison tests
The effect of elk and soil moisture on nitrate delivery rate (a), the effect of elk and proportion of coarse sand (0.5–1 mm) on % soil moisture (b), and the effect of elk and bulk density on % soil moisture (c)
The effect of elk and proportion of very fine sand/silt/clay on ammonium delivery rate
The log response ratio of nitrate as a function of total dung area (a); the log response ratio of bulk density as a function of total dung area (data identified as outliers via interquartile range method in gray; b); log response ratio of nitrate as a function of the log response ratio of bulk density (outliers in gray; c); and the log response ratio of phosphate as a function of the log response ratio of bulk density (outliers in gray; d)
Context‐dependent effects of a reintroduced ungulate on soil properties are driven by soil texture, moisture, and herbivore activity

September 2020

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95 Reads

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7 Citations

Although there is considerable evidence that large mammalian herbivores influence ecosystem‐level processes, studies have reported such widely varying results that generalizations have remained elusive. Here, we use an 18‐year‐old exclosure experiment—stratified across a landscape heterogeneous with respect to soil texture, moisture and herbivore activity—to understand the variable effects of tule elk (Cervus canadensis nannodes), a native reintroduced herbivore, on soil properties along the coast of northern California. Elk significantly increased soil bulk density and created a compacted layer at shallow soil depth, while decreasing infiltration rate and pH. The effects of elk on bulk density, penetration resistance, and pH varied with soil type, being least pronounced in coarse, sandy loams, and greatest in loose sand. The effects of elk on nutrient availability varied along gradients of soil texture and moisture. In coarser soils, elk decreased ammonium availability, but increased it in finer soils. Elk also decreased soil moisture content, in part through their positive effect on bulk density, and this effect was most pronounced in coarser soils. Through decreasing soil moisture content, elk also decreased nitrate availability in coarser soils. At greater levels of elk activity (as measured by dung deposition), the elk effect on bulk density was amplified, and this had a corresponding negative effect on nitrate and phosphate availability. Our study has demonstrated that a better understanding of spatial variation in the effects of herbivores on ecosystems can emerge by evaluating their influences across gradients of soil texture, soil moisture, and herbivore activity. These data enabled us to evaluate several frameworks that have been developed to understand the variable effects of herbivores on ecosystems, which is a significant step in reconciling the many competing ideas put forth to explain the context‐dependent effects of large herbivores on grazed ecosystems.


Long‐term and interactive effects of different mammalian consumers on growth, survival, and recruitment of dominant tree species

July 2020

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109 Reads

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7 Citations

Throughout the world, numerous tree species are reported to be in decline, either due to increased mortality of established trees or reduced recruitment. The situation appears especially acute for oaks, which are dominant features of many landscapes in the northern hemisphere. Although numerous factors have been hypothesized to explain reductions in tree performance, vertebrate herbivores and granivores may serve as important drivers of these changes. Here, using data from 8- and 14-year-old exclosure experiments, we evaluated the individual and interactive effects of large and small mammalian herbivores on the performance of three widespread oak species in California-coast live oak (Quercus agrifolia), California black oak (Q. kelloggii), and Oregon white oak (Q. garryana). Although impacts varied somewhat by species and experiment, herbivory by black-tailed deer (Odocoileus hemionus columbianus) reduced the height and survival of juvenile coast live oaks and altered their architecture, as well as reduced the abundance of black oak seedlings, the richness of woody species and the cover of nonoak woody species. Small mammals (Microtus californicus and Peromyscus maniculatus) had even more widespread effects, reducing the abundance of black oak seedlings and the height and cover of all three oak species. We also detected numerous interactions between small mammals and deer, with one herbivore having positive or negative effects on oak abundance and cover when the other herbivore was either present or absent. For example, deer often had negative effects on seedling abundance only when, or even more so when, small mammals were present. In summary, mammalian consumers play crucial roles in limiting oak recruitment by reducing seedling abundance, maintaining trees in stunted states, and preventing them from reaching sapling stages and becoming reproductive. Interactions between large and small mammals can also alter the intensity and direction of their effects on trees.


Cascading effects of mammalian herbivores on ground‐dwelling arthropods: Variable responses across arthropod groups, habitats and years

June 2019

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62 Reads

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15 Citations

Large mammalian herbivores are well known to shape the structure and function of ecosystems world‐wide, and these effects can in turn cascade through systems to indirectly influence other animal species. A wealth of studies has explored the effects of large mammals on arthropods, but to date they have reported such widely varying results that generalizations have been elusive. Three factors are likely drivers of this variability: the widely varying life‐history characteristics of different arthropod groups, the highly variable landscapes that mammalian herbivores commonly inhabit and temporal variation in environmental conditions. Here, we use an 18‐year‐old exclosure experiment stratified across three distinct coastal prairie habitats in northern California to address the effects of a reintroduced mammalian herbivore, tule elk (Cervus canadensis nannodes) on the composition, richness and abundance of ground‐dwelling arthropods over two years with very different precipitation regimes. We found that elk shifted the composition of arthropod communities, increasing the abundance of ants, beetles, spiders and mites, decreasing the abundance of woodlice and bristletails in some but not all habitats types, and having no effect on the abundance of bugs, crickets and springtails. Elk also increased richness and changed the composition of ant genera and beetle morpho‐species. Interestingly, the effects of elk on arthropod composition, richness and abundance varied little between years, despite very different precipitation levels, biomass accumulation and thatch height. Elk reduced shrub cover, above‐ground herbaceous biomass and thatch height and increased soil compaction, and these changes predicted the abundance and richness of arthropods, although taxonomic groups varied in their responses, presumably due to differences in environmental requirements. Synthesis. Our research highlights the importance of using long‐term experiments to assess the cascading effects of large herbivores on the composition of grounddwelling arthropod communities and to identify the mechanisms that indirectly shape arthropod responses to herbivores among variable habitats and years in order to develop a greater understanding of the variable responses of arthropods to large mammalian herbivores.


Citations (11)


... climate moisture index (CMI), which is calculated based on temperature, Wang et al. 2016] for the 5 years preceding when we sampled each plot (Table 2). We chose 5 years to be consistent with other studies that have assessed the inf luence of climate on juvenile aspen (Clement et al. 2019;Reikowski et al. 2022). In addition to climate variables obtained directly from ClimateNA, we calculated monsoon index (summer precipitation ÷ annual precipitation) and annual dryness index (annual degree-days above 5 • C ÷ annual precipitation) because of the importance of the monsoon system in Arizona and the important inf luence of precipitation, in general, on aspen occurrence, growth, and mortality (Rehfeldt et al. 2009;Worrall et al. 2013;Kane et al. 2014;Ireland et al. 2020). ...

Reference:

Sustainability and drivers of Populus tremuloides regeneration and recruitment near the southwestern edge of its range
Ungulate herbivores as drivers of aspen recruitment and understory composition throughout arid montane landscapes

... Moderate grazing is one of the most effective management strategies to maintain species diversity in grassland ecosystems (Oba et al., 2001;Bai et al., 2004;Wu et al., 2017;Zhan et al., 2020;Li et al., 2022). Large herbivores can exert profound effects on plant diversity directly by the selective consumption and trampling behaviors (Sun et al., 2018;Zhong et al., 2022) and indirectly by the excreta fertilization (Eldridge et al., 2016;Farji-Brener and Werenkraut, 2017;Zhong et al., 2021). Grazing exclusion often results in large standing plant cover which subsequently intensifies the interspecific competition for light and/or soil nutrient resources and subsequently leads to a loss of plant species diversity (Oba et al., 2001), while large herbivores could prevent competitive exclusion by direct removal of biomass of dominant species group (Harpole and Tilman, 2007;Borer et al., 2014). ...

Large herbivores facilitate a dominant grassland forb via multiple indirect effects

... The estimated 26% decline in aspen cover over the past four decades is consistent with observations from other parts of the western United States (Pierce and Taylor 2010, Estes 2016, Refsland and Cushman 2021.This trend is particularly concerning given the ecological importance of aspen in the Lake Tahoe Basin, where it has been identified as one of nine Ecologically Significant Areas that disproportionately support biodiversity relative to their area (Murphy et al. 2000). Several interacting factors may be contributing to this decline. ...

Continent-wide synthesis of the long-term population dynamics of quaking aspen in the face of accelerating human impacts

Oecologia

... We found that this effect continued in the three-week period after cattle left and did not rebound to the intensity of use recorded at those sites prior to cattle grazing. Our results are consistent with previous studies increased home range sizes during cattle grazing periods, and avoidance or reduced use of areas in which cattle were present [15,[26][27][28]. As we expected, elk were more affected by cattle presence than the other three species, likely because elk diet overlap more with cattle than the other ungulates. ...

Effects of human-altered landscapes on a reintroduced ungulate: Patterns of habitat selection at the rangeland-wildland interface
  • Citing Article
  • May 2021

Biological Conservation

... Our study reinforces the durable, positive effect of conifer removal on perennial forbs and grasses consistently demonstrated in previous studies. Field research shows that tree removal increases water and nutrient availability ( Bates et al., 20 0 0 ;Young et al., 2013 ;Roundy et al., 2014b ;Bates and Davies, 2017 ), generally resulting in increased perennial herbaceous vegetation and shrub cover ( Ross et al., 2012 ;Roundy et al., 2014aRoundy et al., , 2014bMiller et al., 2014 ;Stephens et al., 2016 ;Havrilla et al., 2017 ;Ernst-Brock et al., 2019 ;Freund et al., 2021 ;Nigro et al., 2023 ;Williams et al., 2020Williams et al., , 2019Williams et al., , 2017. Positive treatment effects on herbaceous vegetation may take several growing seasons to emerge (e.g., Havrilla et al., 2017 ), but the effects are often persistent. ...

Plant functional groups and species contribute to ecological resilience a decade after woodland expansion treatments

... Red deer were expected to significantly affect soil compaction since big animals are known to increase soil penetration resistance and bulk density, changing also soil texture (Whalley et al. 1995;Lobry & Kingston 1997;Hamza & Anderson 2005;Eldridge et al. 2017;Navarro-Castilla et al. 2017;Shah et al. 2017;Dodge et al. 2020). ...

Context‐dependent effects of a reintroduced ungulate on soil properties are driven by soil texture, moisture, and herbivore activity

... The browsed leafless seedlings were found both concentrated in one space and as dispersed single individuals growing within the same subplot among (1-4) other leafy specimens, which resulted from a different browsing strategy of herbivores. Some of them can forage leisurely in one place, which results in local damage to a higher number of seedlings, while others gnaw on seedlings when they wander (Cushman et al., 2020;Götmark et al., 2005;Oswalt et al., 2006, and references therein). ...

Long‐term and interactive effects of different mammalian consumers on growth, survival, and recruitment of dominant tree species

... Up to date, it has been reported that 60% of the sampled rural families have reduced the expectation on the product output from family gardens, including animal-feeding and vegetable plantation for their own safety when they realized the dangerous characteristics of agrochemical over-input [21]. The public seems to gradually believe that grassland based meat production provides a healthier procedure for animal feeding than factory farming, because it allows more physical practice to animals and requires fewer chemical inputs during the feeding period [31][32]. If the linkage between rural and urban becomes closer and valuable, the degrading rural ecosystem can be restored and sustained via the sustainable maintenance of grassland ecosystems with the concerns or support from urban citizens. ...

Effects of native bryophytes on exotic grass invasion across an environmental gradient

... The individual number, genus number and biomass of Carabidae are positively correlated with plant aboveground biomass (AGB) ( Table 3). The plant AGB in the mowing plots is not determined at the same time of arthropod sampling in this study, however, early studies suggest that autumn mowing can reduce the accumulation of litter on the surface, thus give some visual predators more open space to hunt (Cecil et al. 2019). Therefore, mowing tends to increase the arthropod biomass compared with CK. ...

Cascading effects of mammalian herbivores on ground‐dwelling arthropods: Variable responses across arthropod groups, habitats and years

... Evidence for ecosystem engineering focuses primarily on contributions from large-bodied herbivores. For example, terrestrial herbivores in grassland ecosystems contribute to the formation of bare ground and soil compaction, which results in landscapes that promote high biodiversity and increased plant community function and productivity 57,58 . Many species also contribute to the distribution and cycling of key nutrients and resources, such as nitrogen, phosphorus and carbon 57,58 . ...

Synthesizing the effects of large, wild herbivore exclusion on ecosystem function