Long-term dynamics of biotic and abiotic resistance to exotic species invasion in restored vernal pool plant communities

Invasion of native ecosystems by exotic species can seriously threaten native biodiversity, alter ecosystem function, and inhibit conservation. Moreover, restoration of native plant communities is often impeded by competition from exotic species. Exotic species invasion may be limited by unfavorable abiotic conditions and by competition with native species, but the relative importance of biotic and abiotic factors remains controversial and may vary during the invasion process. We used a long-term experiment involving restored vernal pool plant communities to characterize the temporal dynamics of exotic species invasion, and to evaluate the relative support for biotic and abiotic factors affecting invasion resistance. Experimental pools (n=256) were divided among controls and several seeding treatments. In most treatments, native vernal pool species were initially more abundant than exotic species, and pools that initially received more native seeds exhibited lower frequencies of exotic species over time. However, even densely seeded pools were eventually dominated by exotic species, following extreme climatic events that reduced both native and exotic plant densities across the study site. By the sixth year of the experiment, most pools supported more exotics than native vernal pool species, regardless of seeding treatment or pool depth. Although deeper pools were less invaded by exotic species, two exotics (Hordeum marinum and Lolium multiflorum) were able to colonize deeper pools as soon as the cover of native species was reduced by climatic extremes. Based on an information-theoretic analysis, the best model of invasion resistance included a nonlinear effect of seeding treatment and both linear and nonlinear effects of pool depth. Pool depth received more support as a predictor of invasion resistance, but seeding intensity was also strongly supported in multivariate models of invasion, and was the best predictor of resistance to invasion by H. marinum and L. multilorum. We conclude that extreme climatic events can facilitate exotic species invasions by both reducing abiotic constraints and weakening biotic resistance to invasion.     

Impact of Grazing Intensity during Drought in an Arizona Grassland

Abstract:  The ecological benefits of changing cattle grazing practices in the western United States remain controversial, due in part to a lack of experimentation. In 1997 we initiated an experimental study of two rangeland alternatives, cattle removal and high-impact grazing, and compared grassland community responses with those with more conventional, moderate grazing practices. The study was conducted in a high-elevation, semiarid grassland near Flagstaff, Arizona (U.S.A.). We conducted annual plant surveys of modified Whittaker plots for 8 years and examined plant composition shifts among treatments and years. High-impact grazing had strong directional effects that led to a decline in perennial forb cover and an increase in annual plants, particularly the exotic cheatgrass ( Bromus tectorum L.). A twofold increase in plant cover by exotic species followed a severe drought in the sixth year of the study, and this increase was greatest in the high-impact grazing plots, where native cover declined by one-half. Cattle removal resulted in little increase in native plant cover and reduced plant species richness relative to the moderate grazing control. Our results suggest that some intermediate level of cattle grazing may maintain greater levels of native plant diversity than the alternatives of cattle removal or high-density, short-duration grazing practices. Furthermore, episodic drought interacts with cattle grazing, leading to infrequent, but biologically important shifts in plant communities. Our results demonstrate the importance of climatic variation in determining ecological effects of grazing practices, and we recommend improving conservation efforts in arid rangelands by developing management plans that anticipate this variation.     

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.     

Effects of Cattle Grazing on Diversity in Ephemeral Wetlands

Abstract:  Cattle are usually thought of as a threat to biodiversity. In regions threatened by exotic species invasion and lacking native wild grazers, however, cattle may produce the type of disturbance that helps maintain diverse communities. Across 72 vernal pools, I examined the effect of different grazing treatments (ungrazed, continuously grazed, wet-season grazed and dry-season grazed) on vernal-pool plant and aquatic faunal diversity in the Central Valley of California. After 3 years of treatment, ungrazed pools had 88% higher cover of exotic annual grasses and 47% lower relative cover of native species than pools grazed at historical levels (continuously grazed). Species richness of native plants declined by 25% and aquatic invertebrate richness was 28% lower in the ungrazed compared with the continuously grazed treatments. Release from grazing reduced pool inundation period by 50 to 80%, making it difficult for some vernal-pool endemic species to complete their life cycle. My results show that one should not assume livestock and ranching operations are necessarily damaging to native communities. In my central California study site, grazing helped maintain native plant and aquatic diversity in vernal pools.     

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 .     

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.     

Species richness and stand stability in conifer forests of the Sierra Nevada

Theoretical and empirical studies have long suggested that stability and complexity are intimately related, but evidence from long-lived systems at large scales is lacking. Stability can either be driven by complex species interactions, or it can be driven by the presence/absence and abundance of a species best able to perform a specific ecosystem function. We use 64 years of stand productivity measures in forest systems composed of four dominant conifer tree species to contrast the effect of species richness and abundance on three stability measures. To perform this contrast, we measured the annual growth increments of > 900 trees in mixed and pure forest stands to test three hypotheses: increased species richness will (1) decrease stand variance, (2) increase stand resistance to drought events, and (3) increase stand resilience to drought events. In each case, the alternate hypothesis was that species richness had no effect, but that species composition and abundance within a stand drove variance, resistance, and resilience. In pure stands, the four species demonstrated significant differences in productivity, and in their resistance and resilience to drought events. The two pine species were the most drought resistant and resilient, whereas mountain hemlock was the least resistant and resilient, and red fir was intermediate. For community measures we found a moderately significant (P = 0.08) increase in the community coefficient of variation and a significant (P = 0.03) increase in resilience with increased species richness, but no significant relationship between species richness and community resistance, though the variance in community resistance to drought decreased with species richness. Community resistance to drought was significantly (P = 0.001) correlated to the relative abundance of lodgepole pine, the most resistant species. We propose that resistance is driven by competition for a single limiting resource, with negative diversity effects. In contrast resilience measures the capacity of communities to partition resources in the absence of a single limiting resource, demonstrating positive diversity effects.     

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.     

Recovery Patterns of Understory Herbs and Their Use as Indicators of Deciduous Forest Regeneration

Abstract: Habitat fragmentation has reduced the richness of native species of forests in northeastern North America. Despite recent large-scale increases in forest cover, studies indicate that understory herbaceous plant communities may take many decades to recover. We studied recovery patterns of vegetation following up to 35 years of forest regeneration in restored former cottage and road sites at Point Pelée National Park, Ontario, Canada, to assess the vulnerability of the understory herbaceous species. Overall, there were no significant differences in the diversity of native species between restored and relatively undisturbed reference sites. There was, however, significant among-site variation in the composition of the native species component of these plant communities. When only restored sites were examined, variation in native species composition was associated with time since site restoration, soil moisture, canopy cover, and distance to continuous forest. Native species were assigned vulnerability rankings according to their relative occurrence in reference and restored sites. Spring-flowering herbs, with ant- or gravity-dispersed seeds, were absent from restored sites and were defined as highly vulnerable. In contrast, summer- and fall-flowering herbs, with vertebrate- and wind-dispersed seeds, dominated restored sites and were less vulnerable. Species of low and intermediate vulnerability had colonized restored sites successfully, and the latter should function as indicators of recovery. In contrast, species with high vulnerability rankings had not recovered at all and, because of their limited dispersal ranges, may recolonize restored sites only if they are actively reintroduced.     

Effects of initial irrigation,seeding date and directlyhauled topsoil on mined land community composition

This study was implemented to determine the effects of one, two, or three months of establishment irrigation, after spring or summer seeding on the long-term composition of mined land plant associations. While the treatment plots were seeded with the same diverse mixture of native species, floristic differences among treatment combinations were apparent after five growing seasons. Summer seeding enhanced the development of seeded warm-season grasses by retarding the initial development of seeded cool-season grasses and by limiting competitive effects of nonseeded species through seedbed tillage. Supplemental irrigation was substantially more important to the development of warm- than cool-season grasses. Seeded perennial forbs performed poorly on all treatments, presumably due to low seed viability, low germination, and/or competition from more water-responsive species. Despite drought conditions during plant establishment, supplemental irrigation was not critical to the ultimate development of nonseeded plant species when seedbed tillage occurred shortly after topsoil was applied. However, irrigation became more important to the long-term development of these species when tillage was delayed. Most forbs encountered were perennial natives that had volunteered from the direct-haul topsoil.Atriplex canescens was the only successful seeded shrub and performed particularly well on the summer seeded plot that received one month of initial irrigation.Authors are research plant ccologist and director of research. This contribution submitted as Montana Agricultural Experiment Station Journal Series No. J-1560. This study was funded entirely by the Western Energy Company, Butte, Montana. The authors express their appreciation to Bill Schwarzkoph and Joe Cocnenberg of Western Energy Company, and former Reclamation Research Unit members Ed DePuit, Chet Skilbred, and Steve Young for their participation in the early stages of this study.     

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.     

Introduced Birds and the Fate of Hawaiian Rainforests

Abstract:  The Hawaiian Islands have lost nearly all their native seed dispersers, but have gained many frugivorous birds and fleshy-fruited plants through introductions. Introduced birds may not only aid invasions of exotic plants but also may be the sole dispersers of native plants. We assessed seed dispersal at the ecotone between native- and exotic-dominated forests and quantified bird diets, seed rain from defecated seeds, and plant distributions. Introduced birds were the primary dispersers of native seeds into exotic-dominated forests, which may have enabled six native understory plant species to become reestablished. Some native plant species are now as common in exotic forest understory as they are in native forest. Introduced birds also dispersed seeds of two exotic plants into native forest, but dispersal was localized or establishment minimal. Seed rain of bird-dispersed seeds was extensive in both forests, totaling 724 seeds of 9 native species and 2 exotics with over 85% of the seeds coming from native plants. Without suitable native dispersers, most common understory plants in Hawaiian rainforests now depend on introduced birds for dispersal, and these introduced species may actually facilitate perpetuation, and perhaps in some cases restoration, of native forests. We emphasize, however, that restoration of native forests by seed dispersal from introduced birds, as seen in this study, depends on the existence of native forests to provide a source of seeds and protection from the effects of ungulates. Our results further suggest that aggressive control of patches of non-native plants within otherwise native-dominated forests may be an important and effective conservation strategy.     

Spatial and temporal patterns of seed dispersal: an important determinant of grassland invasion.

We measured spatial and temporal patterns of seed dispersal and seedling recruitment for 58 species in a grassland community to test whether seed dispersal could predict patterns of invasion after disturbance. For the 12 most abundant grasses, recruitment of native species was dependent on the propagule supply of both native and exotic species. Variability in seed rain on small spatial (1-10 m) and temporal (within season) scales led to qualitative differences in the outcome of disturbance colonization such that native species dominated disturbances when exotic seed supply was low but failed to establish when exotic seed supply was high. Local dispersal and spatial heterogeneity in species composition promoted coexistence of native and exotic species by creating refuges from high exotic seed supply within native dominated patches. Despite this, copious exotic seed production strongly limited recruitment of native species in exotic dominated patches. Most grasslands in California are presently dominated by exotic species, suggesting that competition at the seedling stage is a major barrier to native species restoration.     

Effects of Widespread Drought‐Induced Aspen Mortality on Understory Plants

Forest die‐off around the world is expected to increase in coming decades as temperature increases due to climate change. Forest die‐off will likely affect understory plant communities, which have substantial influence on regional biological diversity, ecosystem function, and land–atmosphere interactions, but how die‐off alters these plant communities is largely unknown. We examined changes in understory plant communities following a widespread, drought‐induced die‐off of trembling aspen (Populus tremuloides) in the western United States. We assessed shrub and herbaceous cover and volume in quadrats in 55 plots located across a wide range of levels of aspen mortality. We measured species richness and composition of herbaceous plant communities by recording species presence and absence in 12 sets of paired (1 healthy, 1 dying) aspen plots. Although understory composition in healthy and dying stands was heterogeneous across the landscape, shrub abundance, cover, and volume were higher and abundance of herbaceous species, cover, and volume were lower in dying aspen stands. Shrub cover and volume increased from 2009 to 2011 in dying stands, which suggests that shrub growth and expansion is ongoing. Species richness of herbs declined by 23% in dying stands. Composition of herbs differed significantly between dying and healthy stands. Richness of non‐native species did not differ between stand types. The understory community in dying aspen stands was not similar to other shrub‐dominated plant communities in the region and may constitute a novel community. Our results suggest that changes in understory plant communities as forests die off could be a significant indirect effect of climate change on biological diversity and forest communities. Efectos de la Mortalidad Extensiva de Álamos Inducida por Sequía sobre Plantas del Sotobosque     

Abiotic constraints eclipse biotic resistance in determining invasibility along experimental vernal pool gradients.

Effective management of invasive species requires that we understand the mechanisms determining community invasibility. Successful invaders must tolerate abiotic conditions and overcome resistance from native species in invaded habitats. Biotic resistance to invasions may reflect the diversity, abundance, or identity of species in a community. Few studies, however, have examined the relative importance of abiotic and biotic factors determining community invasibility. In a greenhouse experiment, we simulated the abiotic and biotic gradients typically found in vernal pools to better understand their impacts on invasibility. Specifically, we invaded plant communities differing in richness, identity, and abundance of native plants (the "plant neighborhood") and depth of inundation to measure their effects on growth, reproduction, and survival of five exotic plant species. Inundation reduced growth, reproduction, and survival of the five exotic species more than did plant neighborhood. Inundation reduced survival of three species and growth and reproduction of all five species. Neighboring plants reduced growth and reproduction of three species but generally did not affect survival. Brassica rapa, Centaurea solstitialis, and Vicia villosa all suffered high mortality due to inundation but were generally unaffected by neighboring plants. In contrast, Hordeum marinum and Lolium multiflorum, whose survival was unaffected by inundation, were more impacted by neighboring plants. However, the four measures describing plant neighborhood differed in their effects. Neighbor abundance impacted growth and reproduction more than did neighbor richness or identity, with growth and reproduction generally decreasing with increasing density and mass of neighbors. Collectively, these results suggest that abiotic constraints play the dominant role in determining invasibility along vernal pool and similar gradients. By reducing survival, abiotic constraints allow only species with the appropriate morphological and physiological traits to invade. In contrast, biotic resistance reduces invasibility only in more benign environments and is best predicted by the abundance, rather than diversity, of neighbors. These results suggest that stressful environments are not likely to be invaded by most exotic species. However, species, such as H. marinum, that are able to invade these habitats require careful management, especially since these environments often harbor rare species and communities.     

Assessing the Potential to Restore Historic Grazing Ecosystems with Tortoise Ecological Replacements

The extinction of large herbivores, often keystone species, can dramatically modify plant communities and impose key biotic thresholds that may prevent an ecosystem returning to its previous state and threaten native biodiversity. A potentially innovative, yet controversial, landscape‐based long‐term restoration approach is to replace missing plant‐herbivore interactions with non‐native herbivores. Aldabran giant (Aldabrachelys gigantea) and Madagascan radiated (Astrochelys radiata) tortoises, taxonomically and functionally similar to the extinct Mauritian giant tortoises (Cylindraspis spp.), were introduced to Round Island, Mauritius, in 2007 to control the non‐native plants that were threatening persistence of native species. We monitored the response of the plant community to tortoise grazing for 11 months in enclosures before the tortoises were released and, compared the cost of using tortoises as weeders with the cost of using manual labor. At the end of this period, plant biomass; vegetation height and cover; and adult, seedling, flower, and seed abundance were 3–136 times greater in adjacent control plots than in the tortoise enclosures. After their release, the free‐roaming tortoises grazed on most non‐native plants and significantly reduced vegetation cover, height, and seed production, reflecting findings from the enclosure study. The tortoises generally did not eat native species, although they consumed those native species that increased in abundance following the eradication of mammalian herbivores. Our results suggest that introduced non‐native tortoises are a more cost‐effective approach to control non‐native vegetation than manual weeding. Numerous long‐term outcomes (e.g., change in species composition and soil seed bank) are possible following tortoise releases. Monitoring and adaptive management are needed to ensure that the replacement herbivores promote the recovery of native plants. Estudiando el Potencial para Restaurar Ecosistemas Históricos de Forrajeo con Reemplazos Ecológicos de Tortugas Terrestres     

Multi-trophic invasion resistance in Hawaii: bioacoustics, field surveys, and airborne remote sensing

We used airborne imaging spectroscopy and scanning light detection and ranging (LiDAR), along with bioacoustic recordings, to determine how a plant species invasion affects avian abundance and community composition across a range of Hawaiian submontane ecosystems. Total avian abundance and the ratio of native to exotic avifauna were highest in habitats with the highest canopy cover and height. Comparing biophysically equivalent sites, stands dominated by native Metrosideros polymorpha trees hosted larger native avian communities than did mixed stands of Metrosideros and the invasive tree Morella faya. A multi-trophic analysis strongly suggests that native avifauna provide biotic resistance against the invasion of Morella trees and exotic birds, thus slowing invasion "meltdowns" that disrupt the functioning of native Hawaiian ecosystems.     

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.     

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.     

Aquatic plant community invasibility and scale-dependent patterns in native and invasive species richness

Invasive species richness often is negatively correlated with native species richness at the small spatial scale of sampling plots, but positively correlated in larger areas. The pattern at small scales has been interpreted as evidence that native plants can competitively exclude invasive species. Large-scale patterns have been understood to result from environmental heterogeneity, among other causes. We investigated species richness patterns among submerged and floating-leaved aquatic plants (87 native species and eight invasives) in 103 temperate lakes in Connecticut (northeastern USA) and found neither a consistently negative relationship at small (3-m2) scales, nor a positive relationship at large scales. Native species richness at sampling locations was uncorrelated with invasive species richness in 37 of the 60 lakes where invasive plants occurred; richness was negatively correlated in 16 lakes and positively correlated in seven. No correlation between native and invasive species richness was found at larger spatial scales (whole lakes and counties). Increases in richness with area were uncorrelated with abiotic heterogeneity. Logistic regression showed that the probability of occurrence of five invasive species increased in sampling locations (3 m2, n = 2980 samples) where native plants occurred, indicating that native plant species richness provided no resistance against invasion. However, the probability of three invasive species' occurrence declined as native plant density increased, indicating that density, if not species richness, provided some resistance with these species. Density had no effect on occurrence of three other invasive species. Based on these results, native species may resist invasion at small spatial scales only in communities where density is high (i.e., in communities where competition among individuals contributes to community structure). Most hydrophyte communities, however, appear to be maintained in a nonequilibrial condition by stress and/or disturbance. Therefore, most aquatic plant communities in temperate lakes are likely to be vulnerable to invasion.