Native weeds and exotic plants: relationships to disturbance in mixed-grass prairie

Disturbance frequently is implicated in the spread of invasive exotic plants. Disturbances may be broadly categorized as endogenous (e.g., digging by fossorial animals) or exogenous (e.g., construction and maintenance of roads and trails), just as weedy species may be native or exotic in origin. The objective of this study was to characterize and compare exotic and native weedy plant occurrence in and near three classes of disturbance – digging by prairie dogs (an endogenous disturbance to which native plants have had the opportunity to adapt), paved or gravel roads (an exogenous disturbance without natural precedent), and constructed trails (an exogenous disturbance with a natural precedent in trails created by movement of large mammals) – in three geographically separate national park units. I used plant survey data from the North and South Units of Theodore Roosevelt National Park and Wind Cave National Park in the northern mixed-grass prairie of western North and South Dakota, USA, to characterize the distribution of weedy native and exotic plants with respect to the three disturbance classes as well as areas adjacent to them. There were differences both in the susceptibility of the disturbance classes to invasion and in the distributions of native weeds and exotic species among the disturbance classes. Both exotic and native weedy species richness were greatest in prairie dog towns and community composition there differed most from undisturbed areas. Exotic species were more likely to thrive near roadways, where native weedy species were infrequently encountered. Exotic species were more likely to have spread beyond the disturbed areas into native prairie than were weedy native species. The response of individual exotic plant species to the three types of disturbance was less consistent than that of native weedy species across the three park units. This revised version was published online in June 2006 with corrections to the Cover Date.     

Evolutionary responses of native plants to novel community members

Both ecological and evolutionary processes can influence community assembly and stability, and native community members may respond both ecologically and evolutionarily as additional species enter established communities. Biological invasions provide a unique opportunity to examine these responses of native community members to novel species additions. Here, I use reciprocal transplant experiments among naturally invaded and uninvaded environments, along with experimental removals of exotic species, to determine whether exotic plant competitors and exotic insect herbivores evoke evolutionary changes in native plants. Specifically, I address whether the common native plant species Lotus wrangelianus has responded evolutionarily to a series of biological invasions by adapting to the presence of the exotic plant Medicago polymorpha and the exotic insect herbivore Hypera brunneipennis. Despite differences in selection regimes between invaded and uninvaded environments and the presence of genetic variation for traits relevant to the novel competitive and plant-herbivore interactions, these experiments failed to reveal evidence that Lotus has responded evolutionarily to the double invasion of Medicago followed by H. brunneipennis. However, when herbivory from H. brunneipennis was experimentally reduced, Lotus plants from source populations invaded by Medicago outperformed plants from uninvaded source populations when transplanted into heavily invaded destination environments. Therefore, Lotus showed evidence of adaptation to Medicago invasion but not to the newer invasion of an exotic shared herbivore. The presence of this exotic insect herbivore alters the outcome of evolutionary responses in this system and counteracts adaptation by the native Lotus to invasion by the exotic plant Medicago. This result has broad implications for the conservation of native communities. While native species may be able to adapt to the presence of one or a few exotics, a multitude of invasions may limit the ability of natives to respond evolutionarily to the novel and frequently changing selection pressures that arise with subsequent invasions.     

Site‐specific responses of native and exotic species to disturbances in a mesic grassland community

Abstract. Grassland communities are increasingly recognized as disturbance‐dependent ecosystems, yet there are few replicated, multi‐site studies documenting vegetation responses to varying frequencies and types of grassland disturbance. Even so, land managers frequently manipulate disturbance regimes in an attempt to favour native grassland plants over exotic species. We conducted a factorial experiment testing three frequencies of clipping combined with litter accumulation, litter removal, and soil disturbance within the highly threatened California coastal prairie plant community. We monitored the response of native/exotic, grass/forb plant guilds once a year for four years. More frequent clipping reduced cover of exotic grasses and favoured exotic forbs, whereas native species were largely unaffected by clipping frequency. Litter accumulation, litter removal, and soil disturbance did not affect vegetation composition. Effects of litter accumulation may take longer than our experiment allowed, and soil disturbance due to our treatments was not sufficiently strong to show consistent effects relative to mammalian soil disturbance. Treatment response of some plant guilds differed among sites, highlighting the importance of replicating experiments at several sites before recommending conservation management practices.     

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.     

Exotic plant species in a C4-dominated grassland: invasibility, disturbance, and community structure

We used data from a 15-year experiment in a C₄-dominated grassland to address the effects of community structure (i.e., plant species richness, dominance) and disturbance on invasibility, as measured by abundance and richness of exotic species. Our specific objectives were to assess the temporal and spatial patterns of exotic plant species in a native grassland in Kansas (USA) and to determine the factors that control exotic species abundance and richness (i.e., invasibility). Exotic species (90% C₃ plants) comprised approximately 10% of the flora, and their turnover was relatively high (30%) over the 15-year period. We found that disturbances significantly affected the abundance and richness of exotic species. In particular, long-term annually burned watersheds had lower cover of exotic species than unburned watersheds, and fire reduced exotic species richness by 80–90%. Exotic and native species richness were positively correlated across sites subjected to different fire (r = 0.72) and grazing (r = 0.67) treatments, and the number of exotic species was lowest on sites with the highest productivity of C₄ grasses (i.e., high dominance). These results provide strong evidence for the role of community structure, as affected by disturbance, in determining invasibility of this grassland. Moreover, a significant positive relationship between exotic and native species richness was observed within a disturbance regime (annually burned sites, r = 0.51; unburned sites, r = 0.59). Thus, invasibility of this C₄-dominated grassland can also be directly related to community structure independent of disturbance. Received: 9 February 1999 / Accepted: 12 May 1999     

Comparison of native and exotic distribution and richness models across scales reveals essential conservation lessons

Comparing native and exotic plant species distribution and richness models can help to reveal the causes of invasive exotic species proliferation and provide recommendations for preserving native‐dominated ecosystems. However, models may have limited applicability if potentially divergent patterns across scales, spatial autocorrelation and correspondence with community‐wide patterns such as species richness are not considered. I modeled the distributions of 20 dominant native and 20 dominant exotic species among and within patches in a heavily‐invaded and threatened ecosystem in western North America, examining the roles of scale and species origin on variable selection, spatial autocorrelation and model accuracy to determine conditions that favour native over exotic dominants, and derive recommendations for effective management. I also compared distribution models with native and exotic species richness models, to determine the extent to which dominant native and exotic species were representative of synoptic community patterns. Predictability was lower for exotic dominants, possibly because they are environmental generalists, and was lower within than among patches. Predictors were generally shared between distribution and richness models; however, species‐specific differences were common within both native and exotic species groups. Predictors for individual species across scales were frequently different and sometimes opposing. Distribution and richness models suggest that management assuming environmental affiliation at one scale may be ineffective at another; that site prioritization to maximize native versus exotic richness may not preserve the habitat of some common native species; and that intensive management to reduce exotics may be difficult due to low predictability and shared affiliations with natives. Comparing native and exotic distribution and richness models at two scales enabled scale‐specific conservation recommendations and elucidated trade‐offs between management for richness and representation that distribution models at an individual scale would not have allowed.     

Invasion of exotic earthworms into ecosystems inhabited by native earthworms

The most conspicuous biological invasions in terrestrial ecosystems have been by exotic plants, insects and vertebrates. Invasions by exotic earthworms, although not as well studied, may be increasing with global commerce in agriculture, waste management and bioremediation. A number of cases has documented where invasive earthworms have caused significant changes in soil profiles, nutrient and organic matter dynamics, other soil organisms or plant communities. Most of these cases are in areas that have been disturbed (e.g., agricultural systems) or were previously devoid of earthworms (e.g., north of Pleistocene glacial margins). It is not clear that such effects are common in ecosystems inhabited by native earthworms, especially where soils are undisturbed. We explore the idea that indigenous earthworm fauna and/or characteristics of their native habitats may resist invasion by exotic earthworms and thereby reduce the impact of exotic species on soil processes. We review data and case studies from temperate and tropical regions to test this idea. Specifically, we address the following questions: Is disturbance a prerequisite to invasion by exotic earthworms? What are the mechanisms by which exotic earthworms may succeed or fail to invade habitats occupied by native earthworms? Potential mechanisms could include (1) intensity of propagule pressure (how frequently and at what densities have exotic species been introduced and has there been adequate time for proliferation?); (2) degree of habitat matching (once introduced, are exotic species faced with unsuitable habitat conditions, unavailable resources, or unsuited feeding strategies?); and (3) degree of biotic resistance (after introduction into an otherwise suitable habitat, are exotic species exposed to biological barriers such as predation or parasitism, “unfamiliar” microflora, or competition by resident native species?). Once established, do exotic species co-exist with native species, or are the natives eventually excluded? Do exotic species impact soil processes differently in the presence or absence of native species? We conclude that (1) exotic earthworms do invade ecosystems inhabited by indigenous earthworms, even in the absence of obvious disturbance; (2) competitive exclusion of native earthworms by exotic earthworms is not easily demonstrated and, in fact, co-existence of native and exotic species appears to be common, even if transient; and (3) resistance to exotic earthworm invasions, if it occurs, may be more a function of physical and chemical characteristics of a habitat than of biological interactions between native and exotic earthworms.     

Life form and life history explain variation in population processes in a grassland community invaded by exotic plants and mammals

The existence of general characteristics of plant invasiveness is still debated. One reason we may not have found these characteristics is because we do not yet understand how processes underlying population dynamics contribute to community composition in invaded communities. Here I modify Ricker stock-recruitment models to parameterize processes important to community dynamics in an invaded grassland community: immigration, maximum intrinsic growth rate, self-regulation, and limitation by other species. I then used the parameterized models in a multi-species stochastic simulation to determine how processes affected long-term community dynamics. By parameterizing the models using the frequency of the 18 most common species in the grassland, I determined that life history and life form are stronger predictors of underlying processes than is native status. Immigration maintains exotic annual grasses and the dominant native perennial grass in the community. Growth rate maintains other perennial species. While the model mirrors the frequency of native species well, exotic species have lower observed than parameterized frequencies, suggesting that they are not reaching their potential frequency. These results, combined with results from past research, suggest that disturbance may be key to maintaining exotic species in the community. Here I showed that a continuous modified Ricker model fit discrete grassland frequency data well. This allowed me to model the dominant species in the community simultaneously and gain insight into the processes that determine community composition.     

Native versus exotic community patterns across three scales: Roles of competition,environment and incomplete invasion

Three fundamental, interrelated questions in invasion ecology are: (1) to what extent do exotic species outcompete natives; (2) are native and exotic communities functionally similar or different; and (3) are differences in biogeographic patterns in native and exotic communities due to incomplete invasions among exotics? These questions are analogous to general questions in community ecology regarding the relative roles of competition, environmental response and dispersal limitation in community assembly. We addressed each of these questions for plant communities in discrete meadow patches, using analyses at three scales ranging from the landscape to microsites. A weak positive relationship between native and exotic species richness in microsites, and a predominance of positive correlations in abundance among native and exotic species pairs suggest that competition has been less important than other factors in determining native versus exotic abundance and community composition. In contrast, models of species richness and community compositional change across scales suggest native versus exotic community patterns are largely determined by a mix of scale-dependent concordant (shared positive or negative) and discordant relationships with environmental variables. In addition, detailed analyses of species-area and species-abundance relationships suggest ongoing expansion of exotic species populations, indicating that the assembly of the exotic community is in its early stages. Thus, while competition does not appear to strongly affect native versus exotic abundances and compositions at present, it may intensify in the future. Our results indicate that synoptic patterns in native versus exotic richness that have been previously attributed to a single cause may in fact be due to a complex mix of concordant and discordant responses to environmental factors across scales. They also suggest that conservation efforts aimed at promoting natives and reducing exotics should focus on the factors and scales for which such a response (i.e., promotion of high native and low exotic richness) can be expected.     

Increasing native, but not exotic, biodiversity increases aboveground productivity in ungrazed and intensely grazed grasslands

Species-rich native grasslands are frequently converted to species-poor exotic grasslands or pastures; however, the consequences of these changes for ecosystem functioning remain unclear. Cattle grazing (ungrazed or intensely grazed once), plant species origin (native or exotic), and species richness (4-species mixture or monoculture) treatments were fully crossed and randomly assigned to plots of grassland plants. We tested whether (1) native and exotic plots exhibited different responses to grazing for six ecosystem functions (i.e., aboveground productivity, light interception, fine root biomass, tracer nitrogen uptake, biomass consumption, and aboveground biomass recovery), and (2) biodiversity-ecosystem functioning relationships depended on grazing or species origin. We found that native and exotic species exhibited different responses to grazing for three of the ecosystem functions we considered. Intense grazing decreased fine root biomass by 53% in exotic plots, but had no effect on fine root biomass in native plots. The proportion of standing biomass consumed by cattle was 16% less in exotic than in native grazed plots. Aboveground biomass recovery was 30% less in native than in exotic plots. Intense grazing decreased aboveground productivity by 25%, light interception by 14%, and tracer nitrogen uptake by 54%, and these effects were similar in native and exotic plots. Increasing species richness from one to four species increased aboveground productivity by 42%, and light interception by 44%, in both ungrazed and intensely grazed native plots. In contrast, increasing species richness did not influence biomass production or resource uptake in ungrazed or intensely grazed exotic plots. These results suggest that converting native grasslands to exotic grasslands or pastures changes ecosystem structure and processes, and the relationship between biodiversity and ecosystem functioning.     

Differentiation between native and exotic plant species from a dry grassland: fundamental responses to resource availability,and growth rates

Abstract Species from a mixed native/exotic dry grassland community in New Zealand were experimentally assessed for three ecophysiological parameters: nutrient response, water response and maximum relative growth rate (RGR_max). These are parameters that relate to factors proposed as important in structuring plant communities in dry environments. Native and exotic species did not differ consistently in water response. Exotic species tended to have a greater response to nutrients, but there was considerable overlap between native and exotic guilds. However, exotic species did have a higher intrinsic growth rate, and this effect was not attributable to differences in life histories. The results suggest that the exotic species are more competitive and more generalist than the native species. These traits are compatible with the concept of the ‘ideal invader’, and suggest the C‐R strategy of Grime’s theory. The native species showed characteristics consistent with stress tolerance (sensu Grime). The paucity of evidence for ecophysiological differentiation between the native and exotic guilds, except in intrinsic growth rate, indicates that the exotic species were able to invade not because they had superior adaptation to the physical environment, but because they possessed, by pre‐adaptation, the same ways of coping with that environment as the existing species. However, their ability to invade can be related to their growth rates.     

Rapid biodiversity declines in both ungrazed and intensely grazed exotic grasslands

Exotic-dominated ecosystems with low diversity are becoming increasingly common. It remains unclear, though, whether differences between native and exotic species (driver model), or changes in disturbances or resources (passenger model), allow exotics to become competitive dominants. In our field experiment, plant species origin (native or exotic), cattle grazing (ungrazed or intensely grazed once), and species composition treatments were fully crossed and randomly assigned to four-species mixtures and monocultures of grassland plants. We found that biodiversity declined more rapidly in exotic than in native species mixtures, regardless of our grazing disturbance treatment. Early declines in species evenness (i.e., increases in dominance) led to subsequent declines in species richness (i.e., local extinctions) in exotic mixtures. Specifically, Simpson’s diversity was 29% lower after 1 year, and species richness was 15% lower after 3 years, in exotic than in native mixtures. These rapid biodiversity declines in exotic mixtures were partly explained by decreased complementarity (i.e., niche partitioning and facilitation), presumably because exotic species lack the coevolutionary history that can lead to complementarity and coexistence in native communities. Thus, our results suggest that exotic species can drive biodiversity declines in the presence or absence of a grazing disturbance, partly because exotic species interactions differ from native species interactions. This implies that restoring plant biodiversity in grasslands may require removal of exotic species, in addition to disturbance management.     

Nutrient enrichment increases plant biomass and exotic plant proportional cover independent of warming in freshwater wetland communities

Anthropogenic environmental change can increase exotic species performance and reduce native biodiversity. Nutrient enrichment may favor exotic plants with higher growth rates. Warming may increase the performance of exotic species from warmer native ranges and/or decrease the performance of locally adapted native species. However, community level impacts of nutrient enrichment and warming may depend on their combined effects on individual species and species interactions. We conducted a factorial 11-month field experiment that manipulated 1) plant origin: native, exotic (species from warmer and nutrient rich habitats), or native-&-exotic; 2) nutrients: ambient or high; and 3) temperature: ambient, +1 °C, or +2 °C. Elevated nutrients increased biomass and exotic plant proportional cover. Exotic diversity was higher with elevated nutrients. Native and exotic biomass responses to elevated nutrients were smaller in native-&-exotic treatments. Elevated nutrients increased the relative abundance of two exotic and decreased one exotic and three native species in native-&-exotic treatments. The predicted exotic to native biomass ratio was higher than the observed ratio, indicating that native plants reduced the potential growth of exotic plants in native-&-exotic treatments. Warming had no effect on plant biomass or diversity. These results suggest that nutrient enrichment increases the performance of some exotic plants and that it is critical to consider native and exotic plant interactions when assessing anthropogenic factor and exotic plant effects on native plant communities.     

Invasibility of mature and 15-year-old deciduous forests by exotic plants

High species richness, resource availability and disturbance are community characteristics associated with forest invasibility. We categorized commonly measured community variables, including species composition, topography, and landscape features, within both mature and 15-year-old clearcuts in West Virginia, USA. We evaluated the importance of each variable for predicting the degree of forest invasion by early-establishing exotic invasive plants. Biotic variables, including overall richness (excluding exotic invasive species) and mutually exclusive native and exotic non-invasive species richness, were the strongest indicators of invasibility. Sites that were located on northeast-facing slopes, more mesic conditions, or in clearcuts were more likely to be invaded by exotic invasive plants. Invasion of clearcut sites was more dependent on available microsites (e.g., lower solar radiation, northeast-facing slopes, and lower elevations) within each site than on the condition of the surrounding landscape, whereas invasion into the mature forests was dependent more on the surrounding landscape (e.g., proximity to paved roads). Our results indicate that exotic invasive plant species in our study area respond similarly as other plant species to resource availability and that competitive interactions are relatively unimportant. Current invasion into this landscape is more likely to be a passive reaction to site conditions instead of a driver of change.     

Multiple drivers of apparent competition reduce re-establishment of a native plant in invaded habitats

Although rarely examined, apparent competition, whereby exotic plants increase consumer pressure on native plants, could play a significant role in affecting native plant establishment in invaded habitats. Moreover, although terrestrial consumer communities often contain many consumer species, little is known about which consumers may generate apparent competition, and whether the strength or mechanism of apparent competition differs among different members of the consumer community. Using consumer-specific experimental exclosures and seed additions in the invaded grasslands of California, we demonstrate that multiple mechanisms of apparent competition are capable of limiting the re-establishment of the native grass Nassella pulchra in the absence of direct competition with exotic plants. The effect of small mammalian consumers (mice and voles) and larger consumers (e.g. rabbits, squirrels, deer) decreased with distance to the exotic forb Brassica nigra , which varied from 0–33 meters from focal N. pulchra . The effect of larger consumers also depended upon characteristics of the plant community directly adjacent (i.e. approx. 1  m) from focal N. pulchra . The effect of large consumers also increased with the richness of the exotic plant community and the degree to which the exotic plant community was dominated by exotic grasses as opposed to exotic forbs. Our finding that apparent competition can be driven by different mechanisms, that the importance of each mechanism depends upon which consumers have access, and that each mechanism has a different spatial extent, suggests that the composition of both the consumer community and the exotic plant community may shape the spatial dynamics of reestablishment, the potential for restoration, and the need for conservation.     

Effects of natural and anthropogenic gradients on native and exotic winter annuals in a southern California Desert

Native annual plant species constitute a large proportion of the plant diversity found in arid vegetation types within the southwestern United States; yet, little is known about controls on diversity patterns along natural and anthropogenic gradients. In this study we evaluated native species richness and exotic species cover across overlapping gradients of precipitation, wind, and N deposition in the Colorado Desert of southern California. Factors allowing native diversity to persist under high N deposition and high wind were also evaluated in a second, focused study at one end of the gradient. We found that gradients in precipitation, nitrogen deposition, and wind were the most important factors to native richness and exotic species cover across the landscape, while local heterogeneity in bare ground influenced richness and cover at the high deposition/windy, or high-disturbance, end of the gradient. Patterns of native diversity were evaluated across the gradients using non-metric multidimensional scaling, which showed diversity was split into two axes: one strongly correlated to precipitation and the other strongly correlated with disturbance factors. The disturbance factors were also positively associated with exotic grass and forb cover. In total, these results indicate that large-scale patterns in disturbance and exotic species cover negatively affect native annual plant species diversity but native species can also persist due to local heterogeneity.     

Differentiation between native and exotic plant species in a dry grassland: realized responses to perturbation,and comparison with fundamental responses

Abstract An area of dry grassland in New Zealand, comprising an equal mixture of native and exotic species, was subject to perturbations of irrigation, fertilization and cessation of grazing. The vegetation response was recorded for 3 years. Total cover, and the contribution of native species to that cover, fluctuated between years even in the control plots. Irrigation increased total cover, but decreased the cover of native species. Fertilization produced the same effects, only less strongly, and also reduced species richness, the loss being in native species. In spite of overall effects of treatments on native and exotic cover, when individual species’ responses to irrigation, fertilization or exclosure were calculated, there was no significant difference between the native and exotic plant guilds. Species differed in their responses, but the native and exotic guilds overlapped. When grouped by morphology, the only significant difference between the responses to perturbation was that forbs and graminoids responded more positively to irrigation than woody and cryptogamic species. The realized responses of the species to the perturbations described here showed little correlation with their physiological responses as determined in previous greenhouse experiments. It is suggested that the realized responses are strongly, and currently unpredictably, influenced by competition from the other species present. Soil nutrients and soil water were both important controls on the community. The relative similarity in the nature of the response to these two factors – nutrients and water – suggests that they affect species in similar ways, possibly because the greater growth rate of the exotic species mediates the short‐term response to both. Grazing has less effect on the current community than either nutrients or water, although it may have been historically important in shaping the species pool. From the poor predictability of field responses from morphological guilds or from ecophysiological responses, it is suggested that the ‘functional types’ approach, although conceptually attractive, lacks experimental support in these grasslands. It is concluded that the exotic species have invaded by being pre‐adapted to the environment with the same environmental responses as the natives, but with the advantage of generally higher growth rates.     

The distribution of native and exotic plants in a naturally fragmented sagebrush-steppe landscape

We tested two general hypotheses for the diversity of native and exotic plants in an undisturbed, naturally fragmented sagebrush-steppe landscape in SE Idaho, USA, evaluating whether the MacArthur–Wilson hypothesis of island biogeography or a suite of environmental variables explained the distributions of native and exotic plants. We also tested a third hypothesis, which incorporated assumptions about the origin of exotic plants and their interaction with native plants. Of the three hypotheses we tested, the hypothesis that included exotic species best explained the diversity of the native plant community. The MacArthur–Wilson model of island biogeography did not explain the diversity of native (R ² = 0.13) or exotic plants well (R ² = 0.11), and the model fit the data poorly. A model of environmental variables better explained the diversity of native (R ² = 0.48) and exotic plants (R ² = 0.57), but it also fit the data poorly. Instead, proximity to a railroad explained the cover (R ² = 0.59) and richness of exotic plants (R ² = 0.63), which then explained the species richness of native plants (R ² = 0.34), and the model fit was adequate and had the lowest AIC value. This suggests that the transportation corridor had a significant, though indirect, effect on the native plant community, even in this undisturbed area. Moreover, explained variance, model fit, and the AIC model selection criteria favored the model with the railroad and exotic species over the M–W and environmental models. Since the habitat patches we studied were largely undisturbed by people and their activities, our results further suggest that the transportation corridor influenced the distribution of exotic plants by serving as a vector for colonization, rather than as a source of disturbance. Additionally, the results suggest that exotic plant species have had a negative effect on the diversity of the native plant community and have changed its composition. The results also support the inference that the nascent exotic plant community is influenced by source-sink (Pulliam in Am Nat 132:652–661, 1988) and assembly dynamics. In contrast, the native plant community appears to be more strongly influenced by environmental conditions associated with an elevational gradient, but there is evidence that the native community also has undergone directional change in species composition, associated with the invasion by non-native species.     

The role of plant fidelity and land-use changes on island exotic and indigenous canopy spiders at local and regional scales

Understanding the processes that lead to successful invasions is essential for the management of exotic species. We aimed to assess the comparative relevance of habitat (both at local and at regional scale) and plant features on the species richness of local canopy spiders of both indigenous and exotic species. In an oceanic island, Azores archipelago, we collected spiders in 97 transects belonging to four habitat types according to the degree of habitat disturbance, four types of plants with different colonisation origin (indigenous vs. exotic), and four types of plants according to the complexity of the vegetation structure. Generalised linear mixed models and linear regressions were performed separately for indigenous and exotic species at the local and regional landscape scales. At the local scale, habitat and plant origin explained the variation in the species richness of indigenous spiders, whereas exotic spider richness was poorly correlated to habitat and plant structure. The surrounding landscape matrix substantially affected indigenous spiders, but did not affect exotic spiders, with the exception of the negative effect exerted by native forests on the richness of exotic species. Our results revealed that the local effect of habitat type, plant origin and plant structure explain variations in the species richness observed at a regional scale. These results shed light on the mechanistic processes behind the role of habitat types in invasions, i.e., plant fidelity and plant structure are revealed as key factors, suggesting that native forests may act as physical barriers to the colonisation of exotic spiders.     

Landscape-scale patterns of biological invasions in shoreline plant communities

Little is known about the patterns and dynamics of exotic species invasions at landscape to regional spatial scales. We quantified the presence (identity, abundance, and richness) and characteristics of native and exotic species in estuarine strandline plant communities at 24 sites in Narragansett Bay, Rhode Island, USA. Our results do not support several fundamental predictions of invasion biology. Established exotics (79 of 147 recorded plant species) were nearly indistinguishable from the native plant species (i.e. in terms of growth form, taxonomic grouping, and patterns of spatial distribution and abundance) and essentially represent a random sub-set of the current regional species pool. The cover and richness of exotic species varied substantially among quadrats and sites but were not strongly related to any site-level physical characteristics thought to affect invasibility (i.e. the physical disturbance regime, legal status, neighboring habitat type, and substrate characteristics). Native and exotic cover or richness were not negatively related within most sites. Across sites, native and exotic richness were positively correlated and exotic cover was unrelated to native richness. The colonization and spread of exotics does not appear to have been substantially reduced at sites with high native diversity. Furthermore, despite the fact that the Rhode Island strandline system is one of the most highly-invaded natural plant communities described to date, exotic species, both individually and as a group, currently appear to pose little threat to native plant diversity. Our findings are concordant with most recent, large-scale investigations that do not support the theoretical foundation of invasion biology and generally contradict small-scale experimental work.