Exotic grasses and feces deposition by an exotic herbivore combine to reduce the relative abundance of native forbs

Increased resource availability can facilitate establishment of exotic plant species, especially when coincident with propagule supply. Following establishment, increased resource availability may also facilitate the spread of exotic plant species if it enhances their competitive abilities relative to native species. Exotic Canada geese (Branta canadensis) introduce both exotic grass seed and nutrients to an endangered plant community on the Gulf Islands of southwestern British Columbia, Canada. I used greenhouse experiments to assess the competitive advantage of the exotic grasses relative to native and exotic forbs in this community and to test the impacts of nutrient addition from goose feces on competitive outcomes. I grew experimental communities varying in their proportion of forbs versus exotic grasses, and added goose feces as a nutrient source. I found that both native and exotic forbs produced significantly more biomass in competition with conspecifics than in competition with the grasses, and that the proportional abundance of two out of three native forbs was lowest in the combined presence of exotic grasses and nutrient addition. In a second experiment, I found that in monoculture all species of forbs and grasses showed equal growth responses to nutrients. The exotic species did not convert additional nutrients into additional biomass at a higher rate, but did germinate earlier and grow larger than the native species regardless of nutrient availability. This suggests that the exotic species may have achieved their competitive advantage partly by pre-empting resources in community mixtures. Small and late-germinating native forbs may be particularly vulnerable to competitive suppression from exotic grasses and forbs and may be at an even greater disadvantage if their competitors are benefiting from early access to additional nutrients. In combination, the input of exotic propagules and additional nutrients by nesting geese may compromise efforts to maintain native community composition in this system.     

Post-pastoral changes in composition and guilds in a semi-arid conservation area, Central Otago, New Zealand

Changes in the vegetation of Flat Top Hill, a highly modified conservation area in semi;arid Central Otago, New Zealand, are described four years after the cessation of sheep and rabbit grazing. Unusually moist weather conditions coincide with the four-year period of change in response to the cessation of grazing. Between 1993 and 1997, the average richness and diversity (H') of species increased, and the average proportion of native species decreased significantly. The vegetation was significantly richer in exotic annual and perennial grass species, exotic perennial forbs, exotic woody species and native tussock grasses in 1997 than in 1993. Eight response guilds of species are identified. Most "remnant" native shrubs and forbs were stable, in that they remained restricted to local refugia and showed little change in local frequency. However, taller native grass species increased, some locally, and others over wide environmental ranges. Rare native annual forbs and several native perennial species from "induced" xeric communities decreased, and this may be a consequence of competition from exotic perennial grasses in the absence of grazing. The invasive exotic herb Sedum acre decreased in abundance between 1993 and 1997, but several other prominent exotic species increased substantially in range and local frequency over a wide range of sites. Exotic woody species, and dense, sward-forming grasses are identified as potential threats to native vegetation recovery.     

The roles of exotic grasses and forbs when restoring native species to highly invaded southern California annual grassland

Many semi-arid shrublands in the western US have experienced invasion by a suite of exotic grasses and forbs that have altered community structure and function. The effect of the exotic grasses in this area has been studied, but little is known about how exotic forbs influence the plant community. A 3-year experiment in southern California coastal sage scrub (CSS) now dominated by exotic grasses was done to investigate the influence of both exotic grasses (mainly Bromus spp.) and exotic forbs (mainly Erodium spp.) on a restoration seeding (9 species, including grasses, forbs, and shrubs). Experimental plots were weeded to remove one, both, or neither group of exotic species and seeded at a high rate with a mix of native species. Abundance of all species varied with precipitation levels, but seeded species established best when both groups of exotic species were removed. The removal of exotic grasses resulted in an increase in exotic and native forb cover, while removal of exotic forbs led to an increase in exotic grass cover and, at least in one year, a decrease in native forb cover. In former CSS now converted to exotic annual grassland, a competitive hierarchy between exotic grasses and forbs may prevent native forbs from more fully occupying the habitat when either group of exotics is removed. This apparent competitive hierarchy may interact with yearly variation in precipitation levels to limit restoration seedings of CSS/exotic grassland communities. Therefore, management of CSS and exotic grassland in southern California and similar areas must consider control of both exotic grasses and forbs when restoration is attempted.     

Mechanisms underlying the impacts of exotic annual grasses in a coastal California meadow

Biological invasions can impact the abundance and diversity of native species, but the specific mechanisms remain poorly discerned. In California grasslands, invasion by European annual grasses has severely reduced the quality of habitat for native forb species. To understand how introduced grasses suppress native and exotic forbs, we examined the response of a Southern California grassland community to factorial removals of live grass and the litter produced in previous seasons. To examine the role that belowground competition for water plays in mediating the impact of grasses, we crossed grass and litter removal treatments with water addition. Our results show that forbs were almost equally suppressed by both competition from live grass and direct interference by litter. Water addition did not ameliorate the effect of grass competition, suggesting that water was not the resource for which plants compete. This evidence is consistent with the susceptibility of forbs to light limitation, especially considering that litter does not consume water or nutrients. Interestingly, despite different histories of co-occurrence with annual grass dominants, native and exotic forbs were comparably suppressed by exotic grasses. Our results indicate that suppression by both live and dead stems underlie the influence of exotic grasses on forb competitors.     

Effects of grazing intensity on growth,reproduction, and abundance of three palatable forbs in Kansas tallgrass prairie

The effects of different intensities of cattle grazing on theaboveground growth, reproduction, and abundances of three palatable forbs werestudied in native tallgrass prairie. Populations of Asterericoides, Ruellia humilis, andAmorpha canescens were sampled at peak flowering duringthe1993–1995 growing seasons in four annually-burned sites varying incattle stocking density [ungrazed, low, moderate, high]. The threeforbs exhibited reduced shoot growth and/or reproduction under moderate toheavy grazing, and in no case did grazing increase any measure of plantperformance. Ruellia showed reduced shoot height andbiomass, percentage of stems flowering, and reproductive biomass in response tograzing. Aster showed decreases in shoot biomass andheightwith grazing. Amorpha showed no change in shoot orreproductive biomass, but a decrease in percent of flowering stems and inreproductive allocation with grazing. Patterns in the percentage of stemsgrazedindicated generally high but variable palatability among these species. Bycontrast, the three species showed inconsistent population response to grazing.Abundance (frequency) of all three species indicated no short-term changebetween years in response to grazing intensity. Responses of these speciesdiffer considerably from those of most other perennial tallgrass prairie forbsthat are unpalatable, unconsumed, and increase in performance (e.g. size,abundance) due to release from competition from the dominant grasses underungulate grazing. The results demonstrate that immediate aboveground growth andreproductive responses of established adults to grazing are not good predictorsof grazer effects on population abundances in tallgrass prairie.     

Alpine road colonisation: An update

A wide road verge on the Bogong High Plains near Falls Creek (north‐eastern Victoria), stabilised with exotic species in the late 1950s, was monitored for species composition and cover between 1989 and 2010. Following the removal of cattle grazing in 1991, the cover and species richness of native shrubs, forbs and grasses increased on the verge. Changes were initially slow, but by 1999, they were profound. The cover of exotic species decreased over the same period, but the species richness of exotic species remained unchanged. A wildfire in 2003 did not alter the trajectories of change. Although native species are proving to be competitive, the exotic rhizomic grass Brown‐top Bent (Agrostis capillaris) has been very persistent and is still locally dominant. Feral horses and rabbits are future threats to the continued colonisation by native species.     

Reduced herbivory during simulated ENSO rainy events increases native herbaceous plants in semiarid Chile

El Niño Southern Oscillation (ENSO) events have profound consequences for the dynamics of terrestrial ecosystems. Since increased climate variability is expected to favour the invasive success of exotic species, we conducted a field experiment to study the effects that simulated rainy ENSO events in combination with herbivores and shade have on the composition of a semiarid herbaceous community in north-central Chile. We hypothesized that water pulses, such as those associated with rainy ENSO events could trigger significant changes in the relative abundance of exotic and native herbaceous species. Specifically, we predicted an increase in native grasses and a reduction in the abundance of exotic species, especially prostrate forbs, if water pulses were combined with reduced herbivory. We found that herbivory by small mammals, especially introduced European rabbits (Oryctolagus cuniculus) and hares (Lepus europaeus), have an overwhelming effect on species abundance and composition in this semiarid herbaceous community. Herbivore exclusion produced an overall increase in herb density and biomass mostly due to the extraordinary growth of tall native grasses (especially Bromus berterianus) that outcompeted small prostrate forbs (both native and exotic ones), and small exotic grasses (Koeleria pleoides, Schismus arabicus). Our results suggest that it might be possible to enhance the recovery of native grasses by applying efficient herbivore control during rainy years such as those associated with ENSO events although a negative consequence would be the loss of small native forbs, which greatly contribute to the richness of herbaceous communities in semiarid ecosystems.     

Manipulating disturbance regimes and seeding to restore mesic Mediterranean grasslands

Question: Can managing disturbance regimes alone or in combination with seeding native species serve to shift the balance from exotic towards native species? Location: Central coast of California, USA. Methods: We measured vegetation composition for 10 yr in a manipulative experiment replicated at three sites. Treatments included no disturbance, grazing and clipping at three frequencies with and without litter removal. We seeded eight native species into clipped plots and compared cover in comparable plots with no seeding. Results: Regardless of frequency, clipping generally shifted community dominance from exotic annual grasses to exotic annual forbs, rather than consistently favoring native species. At one site, perennial grass cover decreased in no‐disturbance plots, but only after 4 yr. Litter removal had minimal impact on litter depth and plant community composition. Grazing had a highly variable effect on the abundance of different plant guilds across sites and years. Seeding increased abundance of only two of eight native species. Conclusions: Managing disturbance regimes alone is insufficient to restore native species guilds in highly‐invaded grasslands and seeding native species has highly variable success.     

Before Cattle: A Comparative Floristic Study of Eucalyptus Savanna Grazed by Macropods and Cattle in North Queensland,Australia1

Floristic differences between cattle-grazed and macropod-grazed sites (protected from domestic stock by enclosure within a Holocene lava flow) were examined in Eucalyptus savanna in north Queensland, Australia. While macropod-grazed sites have had a lower incidence of fire than the cattle-grazed sites, it seems probable that grazing impacts per se are the major differences between these treatments. One dominant (Capillipedium parviflorum) and another common (Sehima nervosa) native perennial grass in the macropod-grazed treatment were not recorded within the cattle-grazed treatment. There was generally a lower species richness and Shannon-Weaver diversity of perennial forbs in the cattle-grazed treatment compared to the macropod-grazed treatment. There was higher richness, diversity, and abundance of annual grasses in the cattle-grazed treatment compared to the macropod-grazed treatment. The relative richness of native and exotic species was not significantly affected by cattle grazing.     

Responses of Shrub Midstory and Herbaceous Layers to Managed Grazing and Fire in a North American Savanna (Oak Woodland) and Prairie Landscape

Worldwide, savanna remnants are losing acreage due to species replacement with shade-tolerant midstory forest species as a response to decades of fire suppression. Because canopy closes grasses and other easily ignitable fuels decline, therefore, fire, when reintroduced after years of absence, is not always effective at restoring the open structure original to these communities. Our study sought to determine if managed grazing is an alternative tool for reducing shrub densities and restoring savanna structure without the impacts on soils and native vegetation observed with unmanaged grazing. We compared effects of fire and managed grazing on shrub and herb composition within degraded oak savanna and tallgrass prairie of the U.S. Upper Midwest using a randomized complete block design. The vegetation response to treatments differed by species and by vegetation type. Total shrub stem densities declined 44% in grazed and 68% in burned paddocks within savanna and by 33% for both treatments within prairie. Within savanna, cattle reduced stem densities of Rubus spp. 97%, whereas fire reduced Ribes missouriense stems 96%. Both fire and grazing were effective at reducing stem numbers for several other shrub species but not to the same degree. Native forbs were suppressed in grazed savanna paddocks, as were native grasses in grazed prairie paddocks along with a minor increase of exotic forbs. We did not observe changes in soil bulk density. We conclude that managed grazing can serve as a valuable supplement but not as a replacement to fire for controlling shrubs in these systems.     

Aboveground productivity and root–shoot allocation differ between native and introduced grass species

Plant species in grasslands are often separated into groups (C₄ and C₃ grasses, and forbs) with presumed links to ecosystem functioning. Each of these in turn can be separated into native and introduced (i.e., exotic) species. Although numerous studies have compared plant traits between the traditional groups of grasses and forbs, fewer have compared native versus introduced species. Introduced grass species, which were often introduced to prevent erosion or to improve grazing opportunities, have become common or even dominant species in grasslands. By virtue of their abundances, introduced species may alter ecosystems if they differ from natives in growth and allocation patterns. Introduced grasses were probably selected nonrandomly from the source population for forage (aboveground) productivity. Based on this expectation, aboveground production is predicted to be greater and root mass fraction to be smaller in introduced than native species. We compared root and shoot distribution and tissue quality between introduced and native C₄ grass species in the Blackland Prairie region of Central Texas, USA, and then compared differences to the more well-studied divergence between C₄ grasses and forbs. Comparisons were made in experimental monocultures planted with equal-sized transplants on a common soil type and at the same density. Aboveground productivity and C:N ratios were higher, on average, in native grasses than in native forbs, as expected. Native and introduced grasses had comparable amounts of shallow root biomass and tissue C:N ratios. However, aboveground productivity and total N were lower and deep root biomass and root mass fraction were greater in native than introduced grasses. These differences in average biomass distribution and N could be important to ecosystems in cases where native and introduced grasses have been exchanged. Our results indicate that native–introduced status may be important when interpreting species effects on grassland processes like productivity and plant N accumulation.     

Patch dynamics in grazed subtropical native pastures in south‐east Queensland

Abstract Patch formation is common in grazed grasslands but the mechanisms involved in the formation and maintenance of patches are not clear. To increase our knowledge on this subject we examined possible reasons for patch formation and the influence of management on changes between patch states in three experiments in native pasture communities in the Crows Nest district, south‐east Queensland. In these communities, small‐scale patches (tall grassland (dominated by large and medium tussock grasses), short swards (dominated by short tussock grasses and sedges), and lawns (dominated by stoloniferous and/or rhizomatous grasses)) are readily apparent. We hypothesized that the formation of short sward and lawn patches in areas of tall grassland was due to combinations of grazing and soil fertility effects. This was tested in Experiment 1 by applying a factorial combination of defoliation, nutrient application and transplants of short tussock and stoloniferous species to a uniform area of tall grassland. Total species density declined during the experiment, was lower with high nutrient applications, but was not affected by defoliation. There were significant changes in abundance of species that provided support for our hypotheses. With light defoliation and low nutrients, the tall grassland remained dominated by large tussock grasses and contained considerable amounts of forbs. With heavy defoliation, the pastures were dominated by medium tussock grasses and there were significant decreases in forbs and increases in sedges (mainly with low nutrients) and stoloniferous grasses (mainly with high nutrients). Total germinable seed densities and those of most species groups were significantly lower in the heavy defoliation than the light defoliation plots. Total soil seed numbers were not affected by nutrient application but there were fewer seeds of the erect forbs and more sedge seeds in plots with high nutrients. The use of resting from grazing and fire to manage transitions between patches was tested. In Experiment 2 , changes in species density and abundance were measured for 5 years in the three patch types with and without grazing. Experiment 3 examined the effects of fire, grazing and resting on short sward patches over 4 years. In Experiment 2 , total species density was lower in lawn than short sward or tall grassland patches, and there were more species of erect forbs than other plant groups in all patch types. The lawn patches were originally dominated by Cynodon spp. This dominance continued with grazing but in ungrazed patches the abundance of Cynodon spp. declined and that of forbs increased. In the short sward patches, dominance of short tussock grasses continued with grazing but in ungrazed plots their abundance declined while that of large tussock grasses increased. The tall grassland patches remained dominated by large and medium tussock species. In Experiment 3 , fire had no effect on species abundance. On the grazed plots the short tussock grasses remained dominant but where the plots were rested from grazing the small tussock grasses declined and the large tussock grasses increased in abundance. The slow and relatively small changes in these experiments over 4 or 5 years showed how stable the composition of these pastures is, and that rapid changes between patch types are unlikely.     

Effects of bison and cattle on growth, reproduction, and abundances of five tallgrass prairie forbs

Forb populations were sampled on Kansas tallgrass prairie to examine the effects of native (bison) and domestic (cattle) ungulates on plant growth, reproduction, and species abundances. Five locally and regionally abundant native tallgrass prairie perennials, Baptisia bracteata, Oenothera speciosa, Vernonia baldwinii, Solidago missouriensis, and Salvia azurea, were selected for study. Replicate watershed-level treatments included three grazing regimes (ungrazed, grazed by cattle, and grazed by bison), and two spring fire frequencies (annually burned and burned at 4-yr intervals). The results show that forb responses to ungulates in tallgrass prairie are complex and vary significantly among plant species, ungulate species, fire regimes, and plant life history stages. Some forbs (e.g., B. bracteata, O. speciosa, and V. baldwinii) increased in growth and reproduction in grazed sites, indicating competitive release in response to selective grazing of the dominant warm-season matrix grasses. Forbs that reduced performance in grazed sites are likely negatively affected by disturbances generated by ungulate nongrazing activities, because none of the forbs studied were directly consumed by bison or cattle. Large grazers had no detectable effect on the frequency of plant damage by other herbivores or pathogens. Significant effects of grazers on patterns of flowering and seed production were not congruent with their effects on population densities, indicating that variation in sexual reproduction plays a minor role in regulating local population abundances. Furthermore, the native and domestic ungulates differ significantly in their effects on forb growth and reproduction.     

Clonal integration facilitates the proliferation of smooth brome clones invading northern fescue prairies

Predicting exotic invaders and reducing their impacts on the biodiversity and function of native ecosystems require understanding of the mechanisms that facilitate their success during key stages of invasion. We determined whether clonal growth, characteristic of the majority of successful invaders of natural areas, facilitates the proliferation of Bromus inermis (smooth brome), an exotic grass invading prairie ecosystems across the Great Plains. By manipulating the below-ground connections of proliferating rhizomes as well as the levels of soil nitrogen along the margins of clones invading northern fescue prairies in Manitoba, Canada, we hypothesized that physiological integration would most benefit ramets invading low resource environments. Severing clonal connections reduced the mass of smooth brome shoots invading native prairies and was exacerbated by the immobilization of soil nutrients with glucose. Clonal connections were equally important in the maintenance of smooth brome density and the horizontal proliferation of ramets. Our results demonstrate the role of physiological integration in the proliferation of a clonal exotic invader and may help explain the success of clonal invaders in other regions. Although integration among invading ramets suggests several possibilities for successful management, future research must continue to elucidate differences in the invasiveness of native versus exotic species as well as the persistence of clonal connections among exotic invaders.     

Strengthening Invasion Filters to Reassemble Native Plant Communities: Soil Resources and Phenological Overlap

Preventing invasion by exotic species is one of the key goals of restoration, and community assembly theory provides testable predictions about native community attributes that will best resist invasion. For instance, resource availability and biotic interactions may represent “filters” that limit the success of potential invaders. Communities are predicted to resist invasion when they contain native species that are functionally similar to potential invaders; where phenology may be a key functional trait. Nutrient reduction is another common strategy for reducing invasion following native species restoration, because soil nitrogen (N) enrichment often facilitates invasion. Here, we focus on restoring the herbaceous community associated with coastal sage scrub vegetation in Southern California; these communities are often highly invaded, especially by exotic annual grasses that are notoriously challenging for restoration. We created experimental plant communities composed of the same 20 native species, but manipulated functional group abundance (according to growth form, phenology, and N‐fixation capacity) and soil N availability. We fertilized to increase N, and added carbon to reduce N via microbial N immobilization. We found that N reduction decreased exotic cover, and the most successful seed mix for reducing exotic abundance varied depending on the invader functional type. For instance, exotic annual grasses were least abundant when the native community was dominated by early active forbs, which matched the phenology of the exotic annual grasses. Our findings show that nutrient availability and the timing of biotic interactions are key filters that can be manipulated in restoration to prevent invasion and maximize native species recovery.     

Effects of grazing intensity in grasslands of the Espinal of central Chile

Question: How is grazing intensity associated with species and morpho‐functional traits (MFTs) composition, productivity and richness of annual dominated grasslands? Have native and exotic species similar associations to this gradient? Location: Anthropogenic grassland in the Espinal vegetation in the sub‐humid area of the mediterranean type climate region of Chile (35°58’ S, 72°17’ W). Methods: Data were obtained from a long‐term (eight years) experiment with six stocking rates (1 to 3.5 sheep/ha). Detrended Correspondence Analysis (DCA) and regression analysis were used to determinate the relationship between grazing intensity and biomass, richness, abundance and traits of the species. Results: The first DCA axis was related to grazing intensity and explained most of the floristic variation (69.3%); the abundance of some non‐native species, e.g. Vulpia megalura were highly correlated with this axis. In the DCA for MFTs the first axis explained 87% of the variance and was also related to grazing intensity; the abundance of small size plants and shallow roots increased with grazing intensity. The relative abundance of grasses and composites, but not of legumes, changed with stocking rate: as grazing intensity increased composites became the predominant species to the detriment of grasses. The above‐ground biomass measured in exclusion cages declined with increasing grazing pressure. The richness of exotic species was greater compared to native ones at low stocking rates, but they converge to similar values at higher stocking rates. However, the relative abundance of exotic species was greater than 75% in all stocking rates. Conclusions: Grazing intensification has large effects in the structure of grassland in central Chile. With grazing intensities greater than 1 sheep/ha species characteristics change; evolving in a few years (6–8) towards a similar community regardless of the stocking rate. The overgrazed community has more native than exotic species richness, possibly due to greater defence traits against herbivory of this group of species.     

Temporal Changes in Native and Exotic Vegetation and Soil Characteristics following Disturbances by Feral Pigs in a California Grassland

Invasive species that increase prevailing disturbance regimes can profoundly alter the composition and structure of ecosystems they invade. Using both comparative and manipulative approaches, we investigated how native and exotic vegetation and soil characteristics at a coastal grassland site in northern California changed through time following disturbances by feral pigs (Sus scrofa). We quantified these successional changes by comparing pig disturbances of varying ages (2, 14, 26+, and 60+ months) during the spring and early summer of 2001. Our results indicate that species richness of native plants increased slowly but steadily through time following disturbances, whereas richness of exotic species rebounded much more rapidly. Percent cover of native perennial grasses also increased steadily through time after pig disturbance, whereas the cover of exotic perennial grasses, annual grasses and forbs initially increased rapidly after disturbance and then remained the same or subsided slightly with time. The cover of native forbs and bulbs either increased weakly through time following disturbance or did not change substantially. Pools of ammonium and nitrate in the soil did not change greatly through time following pig disturbance. Net mineralization rates for ammonium and nitrate also varied little with age since disturbance, although we did find that nitrate mineralization was greater at intermediate ages in one study. Neither organic matter content or particle size varied significantly with disturbance age. In summary, we have shown that native and exotic plants from different functional groups vary greatly in how they recovered from pig disturbances. Exotic taxa were generally able to rapidly colonize and persist in pig disturbances, whereas native taxa usually exhibited a slow but steady rebounding following pig disturbance. Given our results, and those of others from nearby sites, we suggest that the health of coastal grasslands may be enhanced substantially by eliminating or greatly reducing the size of feral pig populations.     

Summer irrigation,grazing and seed addition differentially influence community composition in an invaded serpentine grassland

Community assembly theory predicts that resource availability, biotic interactions, and dispersal dynamics will determine community composition. Recent work has demonstrated that manipulating these processes or “filters” to exclude exotic species may assist in restoring invaded plant communities. In this study, we began by manipulating an abiotic filter, summer water availability, on the theory that irrigation prior to the growing season could trigger the germination of exotic species during unfavorable environmental conditions. First, we performed a greenhouse experiment to assess the germination traits of 23 native and exotic species at low (16°C, spring) and high (30°C, summer) temperatures. At summer temperatures, we found high emergence of many exotic and native grasses and low emergence of native forbs suggesting that summer irrigation may help deplete the exotic seed bank. In a second experiment, we established field plots to test the efficacy of summer irrigation and simultaneously manipulated a biotic and a dispersal filter, subjecting some plots to grazing and/or native seed addition. Summer irrigation and seed addition had no effect on percent cover or species richness while grazing reduced native cover but increased native species richness and soil nitrogen content. Our data suggest that manipulating grazing (a biotic filter) may be more effective than altering abiotic or dispersal filters when restoring invaded serpentine grassland. However, summer irrigation may also be effective, if applied at lower temperatures or for longer periods.     

Multi-scale responses of vegetation to removal of horse grazing from Great Basin (USA) mountain ranges

Although free-roaming equids occur on all of the world’s continents except Antarctica, very few studies (and none in the Great Basin, USA) have either investigated their grazing effects on vegetation at more than one spatial scale or compared characteristics of areas from which grazing has been removed to those of currently grazed areas. We compared characteristics of vegetation at 19 sites in nine mountain ranges of the western Great Basin; sites were either grazed by feral horses (Equus caballus) or had had horses removed for the last 10–14 years. We selected horse-occupied and horse-removed sites with similar aspect, slope, fire history, grazing pressure by cattle (minimal to none), and dominant vegetation (Artemisia tridentata). During 1997 and 1998, line-intercept transects randomly located within sites revealed that horse-removed sites exhibited 1.1−1.9 times greater shrub cover, 1.2–1.5 times greater total plant cover, 2–12 species greater plant species richness, and 1.9–2.9 times greater cover and 1.1–2.4 times greater frequency of native grasses than did horse-occupied sites. In contrast, sites with horses tended to have more grazing-resistant forbs and exotic plants. Direction and magnitude of landscape-scale results were corroborated by smaller-scale comparisons within horse-occupied sites of horse-trail transects and (randomly located) transects that characterized overall site conditions. Information-theoretic analyses that incorporated various subsets of abiotic variables suggested that presence of horses was generally a strong determinant of those vegetation-related variables that differed significantly between treatments, especially frequency and cover of grasses, but also species richness and shrub cover and frequency. In contrast, abiotic variables such as precipitation, site elevation, and soil erodibility best predicted characteristics such as forb cover, shrub frequency, and continuity of the shrub canopy. We found species richness of plants monotonically decreased across sites as grazing disturbance increased, suggesting that either the bell-shaped diversity-disturbance curve of the intermediate-disturbance hypothesis does not apply in this system or that most sites are already all on the greater-disturbance slope of the curve. In our study, numerous vegetation properties of less-grazed areas and sites differed notably from horse-grazed sites at local and landscape scales during a wetter and an average-precipitation year. Electronic Supplementary Material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Severe plant invasions can increase mycorrhizal fungal abundance and diversity

Invasions by non-native plants can alter ecosystem functions and reduce native plant diversity, but relatively little is known about their effect on belowground microbial communities. We show that invasions by knapweed (Centaurea stoebe) and leafy spurge (Euphorbia esula, hereafter spurge)—but not cheatgrass (Bromus tectorum)—support a higher abundance and diversity of symbiotic arbuscular mycorrhizal fungi (AMF) than multi-species native plant communities. The higher AMF richness associated with knapweed and spurge is unlikely due to a co-invasion by AMF, because a separate sampling showed that individual native forbs hosted a similar AMF abundance and richness as exotic forbs. Native grasses associated with fewer AMF taxa, which could explain the reduced AMF richness in native, grass-dominated communities. The three invasive plant species harbored distinct AMF communities, and analyses of co-occurring native and invasive plants indicate that differences were partly driven by the invasive plants and were not the result of pre-invasion conditions. Our results suggest that invasions by mycotrophic plants that replace poorer hosts can increase AMF abundance and richness. The high AMF richness in monodominant plant invasions also indicates that the proposed positive relationship between above and belowground diversity is not always strong. Finally, the disparate responses among exotic plants and consistent results between grasses and forbs suggest that AMF respond more to plant functional group than plant provenance.