Predicting Richness of Native, Rare, and Exotic Plants in Response to Habitat and Disturbance Variables across a Variegated Landscape

Species richness of native, rare native, and exotic understorey plants was recorded at 120 sites in temperate grassy vegetation in New South Wales. Linear models were used to predict the effects of environment and disturbance on the richness of each of these groups. Total native species and rare native species showed similar responses, with richness declining on sites of increasing natural fertility of parent material as well as declining under conditions of water enrichment (resulting from human-induced changes in drainage characteristics, leading to increased run-off), severe livestock grazing, and soil disturbance. The response of rare native species to water enrichment, however, was significantly greater than that of all native species. Exotic species richness varied in reverse to that of native species with positive responses to water enrichment and soil disturbance. The contrasting behaviors are attributed to differences in the evolutionary history of native and exotic assemblages and their resulting preadaptations to a landscape recently subjected to agricultural settlement. It would appear that for exogenous disturbances, the intermediate disturbance hypothesis is not supported by our data. In the sampled region, pastures represent the major land-use in terms of area, but have relatively low densities of native and rare species compared with more lightly grazed areas. However, their management is considered to be essential to the maintenance of diversity on a regional scale.     

Modifying native and exotic species richness correlations: the influence of fire and seed addition.

An important goal in restoration is to increase the richness of native species while reducing exotic species. However, native species richness is often positively correlated with exotic species richness. In a grassland-savanna system in Michigan (USA), we show that management that focuses on changing the nature of the exotic-native richness relationship can be used to restore native communities. Native and exotic species richnesses were positively correlated, likely due to a shared coupling with aboveground live biomass (a surrogate for productivity). The addition of native seed shifted the exotic-native richness relationship from a linear positive to a monotonic relationship: in areas of intermediate levels of exotic species richness, seed addition increased native diversity without an associated effect on exotic diversity, but in areas of high or low exotic richness, it did not affect native species richness. Prescribed burning broke the correlation between native and exotic richness with no consistent effect on the richness of either group. However, when burning was combined with native-seed addition, the relationship between native and exotic richness was maintained and was shifted upwards, enhancing native recruitment. Although aboveground productivity was strongly related to species richness across the landscape, changes in productivity did not drive these shifts.     

Cattle Grazing Impacts on Annual Forbs and Vegetation Composition of Mesic Grasslands in California

Abstract:   Livestock grazing represents a major human alteration of natural disturbance regimes in grasslands throughout the world, and its impacts on plant communities have been highly debated. We investigated the impact of cattle grazing on the California coastal prairie plant community with a focus on native annual forbs, a number of which are of conservation concern. In spring 2000 and 2001, we surveyed the vegetation community composition, vegetation structure, and soil chemical parameters at 25 paired grazed and ungrazed sites over a 670-km range of the ecosystem. Native annual forb species richness and cover were higher in grazed sites, and this effect was concomitant with decreased vegetation height and litter depth. Soil properties explained less of the variation. Exotic annual grass and forb cover were higher in grazed sites. Native grass cover and species richness did not differ in grazed and ungrazed sites, but cover and species richness of native perennial forbs were higher in ungrazed sites. Our results suggest that cattle grazing may be a valuable management tool with which to conserve native annual forbs in the ecosystem we studied but that grazing differentially affects the various life-history guilds. Therefore, land managers must focus on creating a matrix of disturbance regimes to maintain the suite of species native to these mesic grasslands. The results of this and other studies highlight the importance of considering the adaptation of vegetation communities to disturbance in making recommendations for grazing management.     

High richness and dense seeding enhance grassland restoration establishment but have little effect on drought response

Restorations commonly utilize seed addition to formerly arable lands where the development of native plant communities is severely dispersal limited. However, variation in seed addition practices may profoundly affect restoration outcomes. Theory and observations predict that species-rich seed mixtures and seeding at high densities should enhance native plant community establishment, minimize exotic species cover, and may promote resistance and resilience to, and recovery from, environmental perturbations. We studied the post-seeding establishment of native plant communities in large grassland restoration plots, which were sown at two densities crossed with two levels of species richness on formerly arable land in Nebraska, USA, and their responses to drought. To evaluate drought resistance, recovery, and resilience of restored plant communities, we erected rainfall manipulation structures and tracked the response of seeded species cover and total plant biomass during experimental drought relative to controls and in the post-drought growing season. High seed richness and high-density seeding treatments resulted in greater richness and cover of native, seeded species per 0.5 m2 compared to low-richness and low-density treatments. Cover differences in response to seed mixture richness were driven by native forbs. Richness and cover of exotic species were lowest in high-richness and high-density treatments. We found little evidence of differential drought resistance, recovery, and resilience among seeding treatments. Increases in exotic species across years were restricted to drought subplots, and were not affected by seeding treatments. Grassland restoration was generally enhanced and exotic cover reduced both by the use of high-richness seed mixtures and high-density seeding. Given the lack of restoration treatment effects on the resistance, recovery, or resilience of seeded species exposed to drought, and the increases in exotic species following drought, other forms of active management may be needed to produce restored plant communities that are robust to climate change.     

Invasion in a diversity hotspot: exotic cover and native richness in the Californian serpentine flora

Exotic species have been observed to be more prevalent in sites where the richness of native species is highest, possibly reflecting variation among sites in resources, propagule supply, heterogeneity, or disturbance. However, such a pattern leaves unclear whether natives at species-rich sites are subject to especially severe impacts from exotics as a result. We considered this question using path models in which relationships between exotic cover and native richness were evaluated in the presence of correlated environmental factors. At 109 sites on serpentine soils across California, USA, exotic cover was positively correlated with total native herbaceous richness and was negatively correlated with the richness of both serpentine-endemic and rare native herbs. However, in path models that accounted for the influences of soil chemistry, disturbance, overstory cover, and regional rainfall and elevation, we found no indication that exotic cover reduced any component of native herb richness. Rather, our results indicated similarities and differences in the conditions favoring exotic, native, endemic, and rare species. Our results suggest that, in spite of some localized impacts, exotic species are not exerting a detectable overall effect on the community richness of the unique native flora of Californian serpentine.     

Importance of Reserve Size and Landscape Context to Urban Bird Conservation

Abstract:  We tested whether reserve size, landscape surrounding the reserve, and their interaction affect forest songbirds in the metropolitan area of Seattle, Washington (U.S.A.), by studying 29 reserves of varying size (small, medium, large) and surrounding urbanization intensity (urban, suburban, exurban). Larger reserves contained richer and less even bird communities than smaller reserves. These size effects disappeared when we removed the positive correlation of shrub diversity with reserve size, suggesting that greater habitat diversity in large reserves supported additional species, some of which were rare. Standardizing the number of individuals detected among all reserve size classes reversed the effect of size on richness in exurban landscapes and reduced the magnitude of the effect in suburban or urban landscapes. The latter change suggested that richness increased with reserve size in most landscapes because larger areas also supported larger samples from the regional bird species pool. Most bird species associated with native forest habitat (native forest species) and with human activity (synanthropic species) were present in reserves larger than 42 ha and surrounded by >40% urban land cover, respectively. Thus, we recommend these thresholds as means for conserving the composition of native bird communities in this mostly forested region. Native forest species were least abundant and synanthropic species most abundant in urban landscapes, where exotic ground and shrub vegetation was most common. Therefore, control of exotic vegetation may benefit native songbird populations. Bird nests in shrubs were most dense in medium (suburban) and large reserves (urban) and tended to be most successful in medium (suburban) and large reserves (exurban), potentially supplying another mechanism by which reserve size increased retention of native forest species.     

Native and exotic plant species exhibit similar population dynamics during succession

A growing body of literature has led to the debate in invasion biology whether exotic species perform within communities differently than native taxa due to inherent advantages. To address this issue, the population dynamics of native and exotic plant species were assessed from a 48-year record of permanent plot data from the Hutcheson Memorial Forest Center (New Jersey, USA) to determine rate of increase, lag time, maximum frequency, and the year of peak frequency. Overall, native and exotic species exhibited very similar population dynamics. Rates of increase and length of lag times were similar between native and exotic taxa but were strongly influenced by plant life form. Short-lived species were characterized by rapid population growth rates and short lag times. Growth rates decreased and lag times increased with species longevity. Overall, correlations between population metrics were the same in native and exotic taxa, suggesting similar trade-offs in life history patterns. The one difference observed was that, in native species, peak frequency was negatively associated with the year of peak frequency (i.e., early-successional species tended to become more abundant), while there was no relationship in exotic species. These analyses show that exotic species behave in essentially the same way as native taxa within dynamic communities. This suggests that abundant native and exotic plant species are exploiting the same range of ecological strategies resulting in similar roles within communities.     

Human impacts, plant invasion, and imperiled plant species in California.

Invasive species are one of the fastest growing conservation problems. These species homogenize the world's flora and fauna, threaten rare and endemic species, and impose large economic costs. Here, we examine the distribution of 834 of the more than 1000 exotic plant taxa that have become established in California, USA. Total species richness increases with net primary productivity; however, the exotic flora is richest in low-lying coastal sites that harbor large numbers of imperiled species, while native diversity is highest in areas with high mean elevation. Weedy and invasive exotics are more tightly linked to the distribution of imperiled species than the overall pool of exotic species. Structural equation modeling suggests that while human activities, such as urbanization and agriculture, facilitate the initial invasion by exotic plants, exotics spread ahead of the front of human development into areas with high numbers of threatened native plants. The range sizes of exotic taxa are an order of magnitude smaller than for comparable native taxa. The current small range size of exotic species implies that California has a significant "invasion debt" that will be paid as exotic plants expand their range and spread throughout the state.     

Estimating non-native plant richness with a species-accumulation model along roads

Monitoring non-native plant richness is important for biodiversity conservation and scientific research. The species-area model (SA model) has been used frequently to estimate the total species richness within a region. However, the conventional SA model may not provide robust estimations of non-native plant richness because the ecological processes associated with the accumulation of exotic and native plants may differ. Because roads strongly dictate the distributions of exotic plants, we propose a species-accumulation model along roads (SR model), rather than an SA model, to estimate the non-native plant richness within a region. Using 270 simulated data sets, we compared the differences in performance between the SR and SA models. A decision tree based on prediction accuracy was created to guide model application, which was validated using field data from 3 national nature reserves in 3 different provinces in China. The SR model significantly outperformed the SA model when non-native species were restricted to the roadsides and the proportion of uncommon exotic species was small. More importantly, the SR model accurately estimated the non-native plant richness in all field sites with an error of <1 species per site. We believe our new model meets the practical need to efficiently and robustly estimate non-native plant richness, which may facilitate effective biodiversity conservations and promote research on non-native plant invasion and vegetation dynamics.     

The invasion paradox: reconciling pattern and process in species invasions

The invasion paradox describes the co-occurrence of independent lines of support for both a negative and a positive relationship between native biodiversity and the invasions of exotic species. The paradox leaves the implications of native-exotic species richness relationships open to debate: Are rich native communities more or less susceptible to invasion by exotic species? We reviewed the considerable observational, experimental, and theoretical evidence describing the paradox and sought generalizations concerning where and why the paradox occurs, its implications for community ecology and assembly processes, and its relevance for restoration, management, and policy associated with species invasions. The crux of the paradox concerns positive associations between native and exotic species richness at broad spatial scales, and negative associations at fine scales, especially in experiments in which diversity was directly manipulated. We identified eight processes that can generate either negative or positive native-exotic richness relationships, but none can generate both. As all eight processes have been shown to be important in some systems, a simple general theory of the paradox, and thus of the relationship between diversity and invasibility, is probably unrealistic. Nonetheless, we outline several key issues that help resolve the paradox, discuss the difficult juxtaposition of experimental and observational data (which often ask subtly different questions), and identify important themes for additional study. We conclude that natively rich ecosystems are likely to be hotspots for exotic species, but that reduction of local species richness can further accelerate the invasion of these and other vulnerable habitats.     

Habitat as a mediator of mesopredator‐driven mammal extinction

A prevailing view in dryland systems is that mammals are constrained by the scarcity of fertile soils and primary productivity. An alternative view is that predation is a primary driver of mammal assemblages, especially in Australia, where 2 introduced mesopredators—feral cat (Felis catus) and red fox (Vulpes vulpes)—are responsible for severe declines of dryland mammals. We evaluated productivity and predation as drivers of native mammal assemblage structure in dryland Australia. We used new data from 90 sites to examine the divers of extant mammal species richness and reconstructed historic mammal assemblages to determine proportional loss of mammal species across broad habitat types (landform and vegetation communities). Predation was supported as a major driver of extant mammal richness, but its effect was strongly mediated by habitat. Areas that were rugged or had dense grass cover supported more mammal species than the more productive and topographically simple areas. Twelve species in the critical weight range (CWR) (35–5500 g) that is most vulnerable to mesopredator predation were extirpated from the continent's central region, and the severity of loss of species correlated negatively with ruggedness and positively with productivity. Based on previous studies, we expect that habitat mediates predation from red foxes and feral cats because it affects these species’ densities and foraging efficiency. Large areas of rugged terrain provided vital refuge for Australian dryland mammals, and we predict such areas will support the persistence of CWR species in the face of ongoing mammal declines elsewhere in Australia.     

Consequences of Buffelgrass Pasture Development for Primary Productivity,Perennial Plant Richness,and Vegetation Structure in the Drylands of Sonora,Mexico

Abstract: In large parts of northern Mexico native plant communities are being converted to non‐native buffelgrass (Pennisetum ciliare) pastures, and this conversion could fundamentally alter primary productivity and species richness. In Sonora, Mexico land conversion is occurring at a regional scale along a rainfall‐driven gradient of primary productivity, across which native plant communities transition from desert scrub to thorn scrub. We used a paired sampling design to compare a satellite‐derived index of primary productivity, richness of perennial plant species, and canopy‐height profiles of native plant communities with buffelgrass pastures. We sampled species richness across a gradient of primary productivity in desert scrub and thorn scrub vegetation to examine the influence of site productivity on the outcomes of land conversion. We also examined the influence of pasture age on species richness of perennial plants. Index values of primary productivity were lower in buffelgrass pastures than in native vegetation, which suggests a reduction in primary productivity. Land conversion reduced species richness by approximately 50% at local and regional scales, reduced tree and shrub cover by 78%, and reduced canopy height. Land conversion disproportionately reduced shrub species richness, which reflects the common practice among Sonoran ranchers of conserving certain tree and cactus species. Site productivity did not affect the outcomes of land conversion. The age of a buffelgrass pasture was unrelated to species richness within the pasture, which suggests that passive recovery of species richness to preconversion levels is unlikely. Our findings demonstrate that land conversion can result in large losses of plant species richness at local and regional scales and in substantial changes to primary productivity and vegetation structure, which casts doubt on the feasibility of restoring native plant communities without active intervention on the part of land managers.     

Biotic Impoverishment and Homogenization in Unfragmented Forest Understory Communities

Abstract:  Ecological change is often hard to document because of a lack of reliable baseline data. Several recent then-versus-now surveys of temperate forest and grassland communities demonstrate losses of local plant species, but most are based on data from a single site. We resurveyed understory communities in 62 upland forest stands in northern Wisconsin (U.S.A.) for which quantitative baseline data exist from 50 years ago. These stands are within a largely unfragmented region but vary in species composition and successional stage. We collected data on changes in (1) total and native species richness, (2) the ratio of exotic to native species, (3) the relative abundance of habitat generalists, and (4) community similarity among sites. We also compared how these rates of change varied over time. Over the past 50 years, native species density declined an average of 18.5% at the 20-m² scale, whereas the ratio of exotic species to native species increased at 80% of all sites. Habitat generalists increased, and habitat specialists declined, accounting in part for an 8.7% rise in average similarity in species composition among sites. Most of these changes cannot be related to succession, habitat loss, or invasion by exotic species. Areas without deer hunting showed the greatest declines in native species density, with parks and research natural areas faring no better than unprotected stands. Animal-pollinated and animal-dispersed species also declined, particularly at unhunted sites. These results demonstrate the power of quantitative multistand data for assessing ecological change and identify overabundant deer as a key driver of community change. Because maintaining forest habitats alone fails to preserve plant diversity at local scales, local biotic simplification seems likely to continue in the region unless active efforts are taken to protect diversity.     

Impact of Native Plants on Bird and Butterfly Biodiversity in Suburban Landscapes

Abstract:  Managed landscapes in which non-native ornamental plants are favored over native vegetation now dominate the United States, particularly east of the Mississippi River. We measured how landscaping with native plants affects the avian and lepidopteran communities on 6 pairs of suburban properties in southeastern Pennsylvania. One property in each pair was landscaped entirely with native plants and the other exhibited a more conventional suburban mixture of plants—a native canopy with non-native groundcover and shrubs. Vegetation sampling confirmed that total plant cover and plant diversity did not differ between treatments, but non-native plant cover was greater on the conventional sites and native plant cover was greater on the native sites. Several avian (abundance, species richness, biomass, and breeding-bird abundance) and larval lepidopteran (abundance and species richness) community parameters were measured from June 2006 to August 2006. Native properties supported significantly more caterpillars and caterpillar species and significantly greater bird abundance, diversity, species richness, biomass, and breeding pairs of native species. Of particular importance is that bird species of regional conservation concern were 8 times more abundant and significantly more diverse on native properties. In our study area, native landscaping positively influenced the avian and lepidopteran carrying capacity of suburbia and provided a mechanism for reducing biodiversity losses in human-dominated landscapes.     

Effect of Vertebrate Grazing on Plant and Insect Community Structure

Abstract: We compared species diversity of plants and insects among grazed and ungrazed areas of Ponderosa pine–grassland communities in Arizona. Plant species richness was higher in two of three grassland communities that were grazed by native elk and deer and domestic cattle than in ungrazed areas inside a series of three large (approximately 40-ha) grazing exclosures. Similarly, plant species richness was higher in grazed areas relative to ungrazed areas at one of two series of smaller (approximately 25-m²) and short-term exclosure sites. Evenness of plant distribution, however, was greater inside ungrazed long-term exclosures but was reduced inside ungrazed short-term exclosures relative to grazed areas. Relative abundances of forbs, grasses, trees, and shrubs, and native and introduced plants did not differ between the long- and short-term grazing exclosures and their grazed counterparts. Relative abundances of some plant species changed when grazers were excluded, however. In contrast, insect species richness was not different between grazed and ungrazed habitats, although insect abundance increased 4- to 10-fold in ungrazed vegetation. Our results suggest that vertebrate grazing may increase plant richness, even in nutrient-poor, semi-arid grasslands, but may decrease insect abundances.     

A Two-Part Measure of Degree of Invasion for Cross-Community Comparisons

Abstract:  Invasibility is a critical feature of ecological communities, especially for management decisions. To date, invasibility has been measured in numerous ways. Although most researchers have used the richness (or number) of exotic species as a direct or indirect measure of community invasibility, others have used alternative measures such as the survival, density, or biomass of either a single or all exotic species. These different measures, even when obtained from the same communities, have produced inconsistent results and have made comparisons among communities difficult. Here, we propose a measure of the degree of invasion (DI) of a community as a surrogate for community invasibility. The measure is expressed as 2 independent components: exotic proportion of total species richness and exotic proportion of total species abundance (biomass or cover). By including richness and abundance, the measure reflects that the factors that control invasibility affect both of these components. Expressing exotic richness and abundance relative to the richness and abundance of all species in a community makes comparisons across communities of different sizes and resource availability possible and illustrates the importance of dominance of exotic species relative to natives, which is a primary management concern associated with exotic species.     

Exploring the internal and external wildlife gradients created by conservation fences

Spillover effects are an expansion of conservation benefits beyond protected areas through dispersal of species that reside within. They have been well documented in marine but not terrestrial systems. To understand the effects on wildlife created by conservation fences, we explored the internal and external gradients of activity in mammal, reptile, and bird species at a conservation reserve in arid Australia that is fenced to exclude invasive rabbits (Oryctolagus cuniculus), cats (Felis catus), and foxes (Vulpes vulpes). Two methods were used: counts of animal tracks along transects on sand dunes and captures at pitfall-trapping sites. In both cases, sites were spaced at different distances from the reserve fenceline inside and outside the reserve. We recorded a range of spillover, source-sink, step, and barrier effects that combined to create a zone within and around the reserve with fence-induced species-specific wildlife gradients. Two endemic rodents but none of the 4 mammal species reintroduced to the reserve showed positive spillover effects. Barrier effects, where activity was highest close to the fence, were recorded for the feral cat and native bettong (Bettongia lesueur), species that could not breach the fence. In comparison, some reptiles and native mammal species that could permeate the fence displayed source-sink effects; that is, their activity levels were reduced close to the fence likely due to constant emigration to the side with lower density. Activity of some reptiles was lowest at sites inside the reserve and gradually increased at outside sites with distance from the fence, a gradient likely related to trophic cascades triggered by predator exclusion. Our result shows that fenced reserves can create overlapping layers of species-specific gradients related to each species’ ability to permeate the fence and its varying susceptibility to threats. Managers should be aware that these gradients may extend for several kilometers either side of the fence and that not all contained species will increase in abundance. Creating wider conservation benefits may require increased fence permeability and threat reduction outside the fence.     

Evaluation of Central North American Prairie Management Based on Species Diversity, Life Form, and Individual Species Metrics

Abstract:  Reintroduction of fire and grazing, alone or in combination, has increasingly been recognized as central to the restoration of North American mixed-grass and tallgrass prairies. Although ecological studies of these systems are abundant, they have generally been observational, or if experimental, have focused on plant species diversity. Species diversity measures alone are not sufficient to inform management, which often has goals associated with life-form groups and individual species. We examined the effects of prescribed fire, light cattle grazing, and a combination of fire and grazing on three vegetation components: species diversity, groups of species categorized by life-form, and individual species. We evaluated how successful these three treatments were in achieving specific management goals for prairies in the Iowa Loess Hills (U.S.A.). The grazing treatment promoted the greatest overall species richness, whereas grazing and burning and grazing treatments resulted in the lowest cover by woody species. Burning alone best achieved the management goals of increasing the cover and diversity of native species and reducing exotic forb and (predominantly exotic) cool-season grass cover. Species-specific responses to treatments appeared idiosyncratic (i.e., within each treatment there existed a set of species attaining their highest frequency) and nearly half of uncommon species were present in only one treatment. Because all management goals were not achieved by any one treatment, we conclude that management in this region may need refining. We suggest that a mosaic of burning and grazing (alone and in combination) may provide the greatest landscape-level species richness; however, this strategy would also likely promote the persistence of exotic species. Our results support the need to consider multiple measures, including species-specific responses, when planning and evaluating management .     

Biodiversity Loss in Latin American Coffee Landscapes: Review of the Evidence on Ants,Birds, and Trees

Abstract: Studies have documented biodiversity losses due to intensification of coffee management (reduction in canopy richness and complexity). Nevertheless, questions remain regarding relative sensitivity of different taxa, habitat specialists, and functional groups, and whether implications for biodiversity conservation vary across regions. We quantitatively reviewed data from ant, bird, and tree biodiversity studies in coffee agroecosystems to address the following questions: Does species richness decline with intensification or with individual vegetation characteristics? Are there significant losses of species richness in coffee‐management systems compared with forests? Is species loss greater for forest species or for particular functional groups? and Are ants or birds more strongly affected by intensification? Across studies, ant and bird richness declined with management intensification and with changes in vegetation. Species richness of all ants and birds and of forest ant and bird species was lower in most coffee agroecosystems than in forests, but rustic coffee (grown under native forest canopies) had equal or greater ant and bird richness than nearby forests. Sun coffee (grown without canopy trees) sustained the highest species losses, and species loss of forest ant, bird, and tree species increased with management intensity. Losses of ant and bird species were similar, although losses of forest ants were more drastic in rustic coffee. Richness of migratory birds and of birds that forage across vegetation strata was less affected by intensification than richness of resident, canopy, and understory bird species. Rustic farms protected more species than other coffee systems, and loss of species depended greatly on habitat specialization and functional traits. We recommend that forest be protected, rustic coffee be promoted, and intensive coffee farms be restored by augmenting native tree density and richness and allowing growth of epiphytes. We also recommend that future research focus on potential trade‐offs between biodiversity conservation and farmer livelihoods stemming from coffee production.     

Influence of a Large Herbivore Reintroduction on Plant Invasions and Community Composition in a California Grassland

Abstract:  Despite many successful reintroductions of large mammalian herbivores throughout the world, remarkably little attention has focused on how these actions affect native and exotic vegetation at reintroduction sites. One such herbivore is tule elk ( Cervus elaphus nannodes ), which was on the brink of extinction in the mid 1800s, but now has numerous stable populations due to intensive reintroduction efforts. Here, we summarize results from a 5-year exclosure experiment that explored the effects of tule elk on a coastal grassland in northern California. Elk significantly altered the species composition of this community; the response of annual species (dominated heavily by exotic taxa) was dramatically different from perennial species. Elk herbivory increased the abundance and aboveground biomass of native and exotic annuals, whereas it either had no effect on or caused significant decreases in perennials. Elk also decreased the cover of native shrubs, suggesting that these herbivores play an important role in maintaining open grasslands. In addition, elk significantly reduced the abundance and biomass of a highly invasive exotic grass , Holcus lanatus, which is a major problem in mesic perennial grasslands. Our results demonstrate that the successful reintroduction of a charismatic and long-extirpated mammal had extremely complex effects on the plant community, giving rise to both desirable and undesirable outcomes from a management perspective. We suspect that these kinds of opposing effects are not unique to tule elk and that land managers will frequently encounter them when dealing with reintroduced mammals.