Species richness and cover along a 60-year chronosequence in old-fields of southeastern Spain

We analyse changes in plant cover and species richness along a 60-year chronosequence in semi-arid Mediterranean old-fields of southeastern Spain. The objectives were: (i) to study patterns of species richness along the abandonment gradient in semi-arid conditions (e.g., to test the “humped-back model” in our system); (ii) to test whether different broad life forms (annuals, forbs, grasses and woody species) showed different patterns along the abandonment gradient, and (iii) to examine to what extent plants with different dispersal strategies dominate at different stages of succession. The explained variance of the regression relating species richness to years since abandonment is improved when considering different life forms. The results suggest that cover and richness of different functional groups show a non-linear unimodal (often positive-skewed) pattern along the gradient (age since abandonment). Maximum total richness is found at young stages of abandonment (<20 years), when most life forms and dispersal strategies coexist. Annuals and perennial forbs reached their maximum richness during the first 10 years of abandonment. About 45% of total woody species richness is reached at this time as a consequence of early colonization of zoochorous shrubs. While the results showed a tendency towards a life-form replacement sequence, the pattern is not so clear when looking at the different dispersal strategies. The results complement previous results in Mediterranean conditions and emphasise the importance of considering different functional types when studying successional patterns. This revised version was published online in June 2006 with corrections to the Cover Date.     

Effects of plant invasions on the species richness of abandoned agricultural land

While exotic plant invasions are thought to lead to declines in native species, the long-term impacts of such invasions on community structure are poorly known. Furthermore, it is unknown how exotic plant invasions compare to invasions by native species. We present data from 40 yr of continuous vegetation sampling of 10 fields released from agriculture to examine the effects of invasions on species richness. The effects of both exotic and native species invasions on species richness were largely driven by variations among fields with most species not significantly affecting species richness. However, invasion and dominance by the exotics Agropyron repens, Lonicera japonica. Rosa multiflora. Trifolium pratense and the native Solidago canadensis were associated with declines in richness. Invasions by exotic and native species during old field succession have similar effects on species richness with dominance by species of either group being associated with loss of species richness. Exotic species invasions tended to have stronger effects on richness than native invasions. No evidence was found of residual effects of invasions because the impact of the invasion disappeared with the decline of the invading population. When pooled across species, heavy invasion by exotic species resulted in greater loss o species richness than invasion by native species. Studies of invasion that utilize multiple sites must account for variability among sites. In our study, had we no included field as a factor we would have incorrectly concluded that invasion consistently resulted in changes in species richness.     

Remnant habitats for grassland species in an abandoned Swedish agricultural landscape

Questions: Which factors influence the persistence of vascular grassland plants in long‐abandoned (at least 50 yr) arable fields and meadows? What might be the implications of current levels of species richness on abandoned arable fields and meadows for future restoration? Location: Forested highlands of Kilsbergen, south central Sweden. Methods: The abundance of all vascular plant species was investigated in three habitat types: former arable fields, hay meadows and outlands (pastures) at 27 farms, abandoned for either approximately 50 yr or 90 yr. Time since abandonment, tree cover, soil depth, degree of soil podsol development, size of the infield area and two measures of connectivity were used as predictors for species richness and species composition. Results: Former outland had denser tree cover, fewer species and fewer grassland species than former arable fields and hay meadows, irrespective of time since abandonment. Former hay meadows and arable fields with a longer time since abandonment were less rich in species, more wooded and had greater podsolization than meadows and fields abandoned at a later stage. Species richness was higher in hay meadows and arable fields at farms with larger infield area and deeper soils compared with farms with smaller infield area and shallower soils. The greatest richness of species and most open habitat were former arable fields at larger farms abandoned 50 yr before the study. Former arable fields had the highest number of grassland species. Conclusion: After 50 yr of abandonment, former arable fields were the most important remnant habitats for grassland species and may be a more promising target for restoration than formerly managed grasslands.     

Afforestation causes changes in post‐fire regeneration in native shrubland communities of northwestern Patagonia,Argentina

Question: What are the effects of fire in native shrubland communities and in pine plantations established in these shrublands? Location: Northern Patagonia, Argentina. Methods: We surveyed four sites in Chall‐Huaco valley, located in northwest Patagonia. Each site was a vegetation mosaic composed of an unburned Pinus ponderosa plantation, a plantation burned in 1996, and an unburned matorral and a matorral burned by the same fire. We recorded the cover of all vascular plant species. We also analysed species richness, total cover, proportion of exotic species, abundance of woody species and herb species, cover of exotic species, abundance of woody and herb species and differences in composition of species. For both shrubs and tree species we recorded the main strategy of regeneration (by resprouting or by seed). Results: We found that fire had different effects on native matorral and pine plantations. Five years after fire, plantations came to be dominated by herbs and exotic species, showing differences in floristic composition. In contrast, matorral communities remained very similar to unburned matorral in terms of species richness, proportion of woody species, and herb species and proportion of exotics. Also, pine plantations were primarily colonized by seedlings, while matorrals were primarily colonized by resprouting. Conclusions: Matorrals are highly fire resilient communities, and the practice of establishing plantations on matorrals produces a strong reduction in the capacity of matorral to return to its original state. The elimination of shrubs owing to the effect of plantations can hinder regeneration of native ecosystems. Burned plantations may slowly develop into ecosystems similar to the native ones, or they may produce a new ecosystem dominated by exotic herbs. This study shows that plantations of exotic conifers affect native vegetation even after they have been removed, as in this case by fire.     

Old-field successional dynamics on the Inland Pampa,Argentina

Abstract. The first 10 yr of old-field successional dynamics on the Argentine Inland Pampa were studied on a series of adjacent plots established consecutively between 1978 and 1989. We examined differences in species abundance patterns among plots in order to detect the spatial and temporal variability of succession. Perennial grasses steadily increased in cover and replaced the dominant annual species after 5 yr. Pioneer dicots persisted in older seral stages with 20 — 23 species/plot. Overall, exotic species (mostly the grasses Lolium multiflorum and Cynodon dactylon) contributed much to the plant cover in these communities. Native grasses comprised 45 % of total cover at years 7 — 10 of succession, but occurred with less than 7 species/plot. Substantial variation was found in the successional pathway, which reflected the particular sequence from annual forbs to short-lived and perennial grasses in the various plots. The course of succession was apparently influenced by a 2-yr period of unusually high rainfall. Deyeuxia viridiflavescens, a native perennial grass virtually absent before the wet period, spread over the study area and dominated seral communities for 3 yr, irrespective of plot age. Climatic conditions thus affected the successional turnover of life forms by increasing the rate of colonization by perennial grasses. We further point out the constraints imposed on secondary succession by the life histories of ‘available’ species.     

Alternate pathways of succession on the Lake Erie Islands

In order to determine the effect of prior land use on secondary succession, woody plant species abundances were determined on the Lake Erie Islands in 21 sites 20–90 yr post-abandonment: 11 former vineyards, 8 former orchards, and 2 former quarries. Detrended Correspondence Analysis ordination was used to compare the relative abundances of woody plants in the 21 successional sites to each other and to 5 mature upland forests and 3 mature lowland forests. Vineyards 20–45 yr post-abandonment had closed canopies 2–4 m tall and were dominated by the same mix of species dominating mature forests at similar elevations. Abandoned orchards 20–45 yr post-abandonment supported species common in mature forest floras as well as many shade-intolerant, pioneer tree and shrub species not found in mature stands. These differences in successional patterns seem to be related to differences in the suite of colonization sites presented upon abandonment, with orchards being dominated by open, bare ground, and vineyards by shaded ground receiving ca. 20% of full sunlight. Newly abandoned vineyards had appreciable ‘advanced regeneration’ of saplings of shade-tolerant species 3–7 yr old growing among the grapes. While former quarries remained open, marginal habitats with sparse, open canopies dominated by species typical of rocky cliff edges, the community structure of former vineyards and orchards converged toward each other and toward the composition of mature forests at similar elevations. Thus patterns of secondary succession varied with prior land use and were determined by the suite of colonization sites available at the time of abandonment.     

Properties of native plant communities do not determine exotic success during early forest succession

Considerable research has been devoted to understanding how plant invasions are influenced by properties of the native community and to the traits of exotic species that contribute to successful invasion. Studies of invasibility are common in successionally stable grasslands, but rare in recently disturbed or seral forests. We used 16 yr of species richness and abundance data from 1 m² plots in a clearcut and burned forest in the Cascade Range of western Oregon to address the following questions: 1) is invasion success correlated with properties of the native community? Are correlations stronger among pools of functionally similar taxa (i.e. exotic and native annuals)? Do these relationships change over successional time? 2) Does exotic abundance increase with removal of potentially dominant native species? 3) Do the population dynamics of exotic and native species differ, suggesting that exotics are more successful colonists? Exotics were primarily annual and biennial species. Regardless of the measure of success (richness, cover, biomass, or density) or successional stage, most correlations between exotics and natives were non‐significant. Exotic and native annuals showed positive correlations during mid‐succession, but these were attributed to shared associations with bare ground rather than to direct biotic interactions. At peak abundance, neither cover nor density of exotics differed between controls and plots from which native, mid‐successional dominants were removed. Tests comparing nine measures of population performance (representing the pace, magnitude, and duration of population growth) revealed no significant differences between native and exotic species. In this early successional system, local richness and abundance of exotics are not explained by properties of the native community, by the presence of dominant native species, or by superior colonizing ability among exotics species. Instead natives and exotics exhibit individualistic patterns of increase and decline suggesting similar sets of life‐history traits leading to similar successional roles.     

Succession of Exotic and Native Species Assemblages within Restored Floodplain Forests: A Test of the Parallel Dynamics Hypothesis

Exotic plants pose a threat to restoration success in post‐agricultural bottomlands, but little information exists on their dynamics during succession of actively restored sites. We hypothesized that exotic assemblages would establish during succession and that their compositional trends during succession time would mirror those published for native species in other systems, with an early peak in herbaceous richness followed by a decline as woody species establish. In the summer of 2008, we sampled 16 sites across an 18‐year chronosequence of restored forests, with an additional four mature forest stands for comparison, within the Cypress Creek NWR, Illinois, U.S.A. We identified all vascular plant species and quantified canopy openness at three canopy strata, and soil texture and chemistry. Trends in exotic assemblages were significantly correlated with canopy openness at all strata. Richness of exotic and native herbaceous species was related to stand age and consistent with a Weibull regression model. Native and exotic herbaceous cover followed an exponential decay model. Woody native richness over time conformed to a logistic model; woody exotics exhibited no relationship with stand age and were present in sites of all ages. Our results indicate that although their rates of decline differ, herbaceous exotics and natives exhibit similar successional dynamics; therefore, herbaceous exotics may not pose a lasting threat to restoration success in reforested floodplains. Woody exotics can establish across a range of successional stages and persist under closed canopy conditions. Bottomland restorations are vulnerable to the invasion and expansion of exotic plant species even after canopy closure.     

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.     

Long-term grazing impacts on vegetation diversity,composition, and exotic species presence across an aridity gradient in northern temperate grasslands

Little is known about the specific role of exotic species on measures of grassland plant diversity, including how this may vary with climatic conditions or large mammal herbivory. This study examined vegetation responses to long-term livestock grazing, including plant richness and diversity, as well as the contribution of exotic species to these metrics, across a network of 107 northern temperate grasslands in Alberta, Canada, spanning a broad aridity gradient. Exposure to grazing modestly increased plant richness, but did not alter Shannon’s diversity, Simpson’s diversity, or evenness, suggesting stability in floral diversity relative to grazing. However, grazing did increase grass cover while reducing shrub cover, the latter of which was only apparent in mesic grasslands. Unlike total plant diversity, exotic species richness and cover, together with exotic plant contributions to diversity, varied jointly with grazing and aridity. While long-term grazing increased exotic species, this response was most apparent in wetter areas, and non-grazed grasslands remained more resistant to the presence of exotics. Several exotic species were positive indicators of grazing in wetter grasslands, and coincided with lower native species cover, indicating grazing may be facilitating a shift from native to exotic vegetation under these conditions. Overall, our results indicate that while long-term grazing has altered the composition and cover of certain functional groups, including favoring exotics and minimizing woody vegetation in mesic areas, overall changes to plant diversity were limited. Additionally, these findings suggest that semi-arid northern temperate grasslands remain relatively resistant to grazing effects, including their susceptibility to exotic plant encroachment. These results improve our understanding of how ongoing grazing exposure may impact grassland diversity, including efforts to conserve native vegetation, as well as the important role of climate in altering fundamental grassland responses to grazing.     

Impacts of mixed severity wildfire on exotic plants in a Colorado ponderosa pine–Douglas-fir forest

The 2002 Hayman Fire burned with mixed severity across 55,800 ha of montane Colorado forest, including pre-existing plots that were originally measured for understory plant composition and cover in 1997. We examined the influence of the Hayman Fire on exotic plants by remeasuring these plots annually from 2003 to 2007. We found that (1) exotic richness and cover generally increased as fire severity and time since fire increased; (2) the exotic species present in a plot before the fire were also largely present in the plot postfire, regardless of fire severity; (3) most of the new postfire species in a plot were present elsewhere in the study area before the fire, although some new species were truly new invaders that were not found in prefire surveys; (4) lightly burned riparian forests were not more susceptible to exotic invasion than surrounding uplands that burned with similar severity; and (5) native and exotic richness and cover were positively correlated or uncorrelated for all fire severities and years. Our findings indicate that exotics were stimulated by the Hayman Fire, especially in severely burned areas. However, exotic richness and cover remain low as of 2007, and correlations between native and exotic richness and cover suggest that exotics have not yet interfered with native understory development. Therefore, we conclude that exotic plants are not a major ecological threat at present, but recommend that monitoring be continued to evaluate if they will pose a threat in future years.     

Impacts of Land Abandonment on Vegetation: Successional Pathways in European Habitats

Changes in traditional agricultural systems in Europe in recent decades have led to widespread abandonment and colonization of various habitats by shrubs and trees. We combined several vegetation databases to test whether patterns of changes in plant diversity after land abandonment in different habitats followed similar pathways. The impacts of land abandonment and subsequent woody colonization on vegetation composition and plant traits were studied in five semi-natural open habitats and two arable habitats in six regions of Europe. For each habitat, vegetation surveys were carried out in different stages of succession using either permanent or non-permanent plots. Consecutive stages of succession were defined on a physiognomic basis from initial open stages to late woody stages. Changes in vegetation composition, species richness, numbers of species on Red Lists, plant strategy types, Ellenberg indicator values of the vegetation, Grime CSR strategy types and seven ecological traits were assessed for each stage of the successional pathway. Abandonment of agro-pastoral land-use and subsequent woody colonization were associated with changes in floristic composition. Plant richness varied according to the different habitats and stages of succession, but semi-natural habitats differed from arable fields in several ecological traits and vegetation responses. Nevertheless, succession occurred along broadly predictable pathways. Vegetation in abandoned arable fields was characterized by a decreasing importance of R-strategists, annuals, seed plants with overwintering green leaves, insect-pollinated plants with hemi-rosette morphology and plants thriving in nutrient-rich conditions, but an increase in species considered as endangered according to the Red Lists. Conversely, changes in plant traits with succession within the initially-open semi-natural habitats showed an increase in plants thriving in nutrient-rich conditions, stress-tolerant plants and plants with sexual and vegetative reproduction, but a sharp decrease in protected species. In conclusion, our study showed a set of similarities in responses of the vegetation in plant traits after land abandonment, but we also highlighted differences between arable fields and semi-natural habitats, emphasizing the importance of land-use legacy.     

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.     

Exotic plants establish persistent communities

Many exotic plants utilize early successional traits to invade disturbed sites, but in some cases these same species appear able to prevent re-establishment of late-successional and native species. Between 2002 and 2004, I studied 25 fields that represent a 52-year chronosequence of agricultural abandonment in a shrub-steppe ecosystem in Washington State, USA, to determine if exotic plants behaved as early successional species (i.e., became less abundant over time) or if they established persistent communities. Exotics maintained dominance in tilled (73% of total cover) relative to never-tilled (6% of total cover) fields throughout the chronosequence. Exotic community composition, however, changed on annual and decadal timescales. Changes in exotic community composition did not reflect typical successional patterns. For example, some exotic perennial species (e.g., Centaurea diffusa and Medicago sativa) were less common and some exotic annual species (e.g., Sissymbrium loeselii and S. altissimum) were more common in older relative to younger fields. Exotics in the study area appeared to establish communities that are resistant to re-invasion by natives, resilient to losses of individual exotic species, and as a result, maintain total exotic cover over both the short- and long-term: exotics replaced exotics. Exotics did not invade native communities and natives did not invade exotic communities across the chronosequence. These results suggest that, in disturbed sites, exotic plants establish an alternative community type that while widely variable in composition, maintains total cover over annual and decadal timescales. Identifying alternative state exotic communities and the mechanisms that explain their growth is likely to be essential for native plant restoration.     

Effects of exotic annual grass litter and local environmental gradients on annual plant community structure

Exotic annual grasses have been introduced into many semi-arid ecosystems worldwide, often to the detriment of native plant communities. The accumulation of litter from these grasses (i.e. residual dry biomass) has been demonstrated to negatively impact native plant communities and promote positive feedbacks to exotic grass persistence. More targeted experiments are needed, however, to determine the relative impact of exotic grass litter on plant community structure across local environmental gradients. We experimentally added exotic grass litter to annual forb-dominated open woodland communities positioned along natural canopy cover gradients in southwest Western Australia. These communities are an important component of this region’s plant biodiversity hotspot and are documented to be under threat from exotic annual grasses. After a one-year treatment period, we measured the effects of exotic grass litter, soil properties, and canopy cover on native and exotic species richness and abundance, as well as common species’ biomass and abundances. Plant community structure was more strongly influenced by soil properties and canopy cover than by grass litter. Total plant abundances per plot, however, were significantly lower in litter addition plots than control plots, a trend driven by native species. Exotic grass litter was also associated with lower abundances of one very common native species: Waitzia acuminata. Our results suggest that exotic grass litter limits the establishment of some native species in this system. Over multiple years, these subtle impacts may contribute substantially to the successful advancement of exotic species into this system, particularly in certain microenvironments.     

Relationship between native and exotic plant species at multiple savannah sites

We tested whether the recently proposed two‐part measure of degree of invasion (DI) of a community relating exotic proportion of cover to exotic proportion of richness can characterize patterns of plant invasions at multiple savannah sites in Southern Africa. Regression analysis was performed on transformed data to assess how this two‐part measure of DI compares to other metrics of community invasibility. The results indicate that at the plot level, the absolute cover of exotics was not significantly related to native cover for three sites out of four assessed (R² ≤ 0.17; P > 0.05). Also, at all four sites, no significant relationships were detected between native and exotic plant richness at both the 1‐m² and 400‐m² plot scales. By contrast, significant (P < 0.05) positive linear relationships were observed between exotic proportion of richness and exotic proportion of cover at all sites (R² was as high as 0.67 and 0.97 for two sites). Our results indicate that the new two‐part measure of DI is able to characterize patterns of plant invasions across plant communities in African savannahs.     

Factors Affecting Revegetation of Oil Field Access Roads in Semiarid Grassland

We assessed vegetation recovery on access roads removed after well abandonment in an active oil‐producing region of northern Great Plains grasslands. We compared extant vegetation on 58 roads, restored 3–22 years previously, to records of species seeded on each and to adjacent, undisturbed prairie, to evaluate main differences between the restored and adjacent community and to explore patterns in the restored plant community over time. The restored plant community was dominated by low richness of seeded non‐native and native grasses and forbs, whereas adjacent prairie had numerous, abundant native graminoids and shrubs and higher richness of native forbs. Cover of seeded species on roads was double that of colonizing species. Disparity in cover of dominant native grasses between the adjacent community and relatively narrow restored roadway suggests that conditions for germination and survival in roadbeds are poor. This is at least partly due to persistence of seeded species. Differences in restored plant composition over time were best explained by changes in species seeded, from non‐natives to natives, and secondarily by successional shifts from ruderal to perennial non‐seeded species. Of the 30 species seeded at least once on these roads, only 10 were commonly used. The long‐term influence of seeding choices in grassland road restorations implies that improvements in these practices will be critical to reversing ecological impacts of roads.     

Size of the local species pool determines invasibility of a C4-dominated grassland

The size of the local species pool (i.e., species surrounding a community capable of dispersal into that community) and other dispersal limitations strongly influence native plant community composition. However, the role that the local species pool plays in determining the invasibility of communities by exotic plants remains to be evaluated. We hypothesized that the richness and abundance of exotic species would be greater in C4‐dominated grassland communities if the local species pool included a larger proportion of exotic species. We also predicted that an increase in the exotic species pool would increase the invasibility of sites thought to be resistant to invasion (annually burned grassland). To test these hypotheses, study plots were established within two long‐term (>20 yr) fire experiments at a tallgrass prairie preserve in NE Kansas (USA). Study plots were surrounded by either a small pool of exotic species (small species pool (SSP) plots; six species) or a larger exotic species pool (large species pool (LSP) plots; 18 species). We found that richness and absolute cover of exotic species was significantly (P<0.001) lower (～70 and 90%, respectively) in annually burned compared to unburned plots, regardless of the size of the exotic species pool. As predicted, exotic species richness was higher (P<0.001) for LSP plots (3.9 per 250 m2) than for SSP plots (0.7 per 250 m2); however, absolute cover was unaffected by the size of the exotic species pool. In the absence of fire, plots with a LSP had four times as many exotic species than SSP plots. An increase in the local exotic species pool also increased the invasibility of annually burned grassland. Indeed, richness of exotic plant species in annually burned LSP plots did not differ from unburned plots with a SSP, indicating that a larger pool of exotic species countered the negative effects of fire. These findings have important implications for predicting how the invasion of plant communities may respond to human‐induced global changes, such as habitat fragmentation. Community characteristics or factors such as frequent fires in grasslands may impart resistance to invasions by exotic species in large, intact ecosystems. However, when a large pool of exotic species is present, frequent fire may not be sufficient to limit the invasions of exotic plants in fragmented landscapes.