Exotic plant invasions over 40 years of old field successions: community patterns and associations

While exotic plant species often come to dominate disturbed communities, long-term patterns of invasion are poorly known. Here we present data from 40 yr of continuous vegetation sampling, documenting the temporal distribution of exotic plant species in old field succession. The relative cover of exotic species decreased with time since abandonment, with significant declines occurring ≥20 yr post-abandonment. The number of exotic species per plot also declined with time since abandonment while field-scale richness of exotics did not change. This suggests displacement occurring at small spatial scales. Life history types changed from short-lived herbaceous species to long-lived woody species for both native and exotic plant species. However, shrubs and lianas dominated woody cover of exotic plants while trees dominated native woody cover. The species richness of exotic and native species was positively correlated at most times. In abandoned hay fields, however, the proportion of exotic plant cover per plot was inversely related to total species richness. This relationship suggests that it is not the presence, but the abundance of exotic species that may cause a reduction in community diversity. While the development of closed-canopy forest appears to limit most introduced plant species, several shade-adapted exotic species are increasing within the fields. These invasions may cause a reversal of the patterns seen in the first 40 yr of succession and may result in further impacts on community structure.     

Invasion impacts local species turnover in a successional system

Exotic plant invasions are often associated with declines in diversity within invaded communities. However, few studies have examined the local community dynamics underlying these impacts. Changes in species richness associated with plant invasions must occur through local changes in extinction and/or colonization rates within the community. We used long‐term, permanent plot data to evaluate the impacts of the exotic vine Lonicera japonica. Over time, species richness declined with increasing L. japonica cover. L. japonica reduced local colonization rates but had no effect on extinction rates. Furthermore, we detected significant reductions in the immigration of individual species as invasion severity increased, showing that some species are more susceptible to invasion than others. These findings suggest that declines in species richness associated with L. japonica invasion resulted from effects on local colonization rates only and not through the competitive displacement of established species.     

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.     

Dominance As An Overlooked Measure of Invader Success

Recent multi-habitat studies across a range of spatial scales have shown that species-rich habitats are often highly invasible by exotic species. The primary measures of invasion in these and other studies are invader richness and the absolute cover or biomass of invaders. We argue that the relative biomass or cover of invaders (dominance) is an important but overlooked measure of plant invasion. We re-analyzed data presented in five previous studies to evaluate whether exotic relative abundance is positively correlated with native richness. There were either no relationships or negative relationships between native richness and relative exotic cover calculated from three spatial scales (1, 1000 and 4000 m²). Thus while the original studies reported high exotic richness or absolute cover in habitats rich in native species, native richness did not predict the degree to which exotics had become dominant or abundant relative to natives. Absolute measures of exotic cover reported in the original studies underestimated relative exotic cover in habitats with low native species richness. High exotic dominance in areas of low native richness may indicate that exotic richness and dominance are controlled by different factors. We conclude that it is useful for researchers to measure both invader richness and invader dominance when trying to understand the environmental factors that are associated with plant invasions.     

Grazing, Environmental Heterogeneity, and Alien Plant Invasions in Temperate Pampa Grasslands

Temperate humid grasslands are known to be particularly vulnerable to invasion by alien plant species when grazed by domestic livestock. The Flooding Pampa grasslands in eastern Argentina represent a well-documented case of a regional flora that has been extensively modified by anthropogenic disturbances and massive invasions over recent centuries. Here, we synthesise evidence from region-wide vegetation surveys and long-term exclosure experiments in the Flooding Pampa to examine the response of exotic and native plant richness to environmental heterogeneity, and to evaluate grazing effects on species composition and diversity at landscape and local community scales. Total plant richness showed a unimodal distribution along a composite stress/fertility gradient ranging several plant community types. On average, more exotic species occurred in intermediate fertility habitats that also contained the highest richness of resident native plants. Exotic plant richness was thus positively correlated with native species richness across a broad range of flood-prone grasslands. The notion that native plant diversity decreases invasibility was supported only for a limited range of species-rich communities in habitats where soil salinity stress and flooding were unimportant. We found that grazing promoted exotic plant invasions and generally enhanced community richness, whereas it reduced the compositional and functional heterogeneity of vegetation at the landscape scale. Hence, grazing effects on plant heterogeneity were scale-dependent. In addition, our results show that environmental fluctuations and physical disturbances such as large floods in the pampas may constrain, rather than encourage, exotic species in grazed grasslands.     

Plant invasions differentially affected by diversity and dominant species in native‐ and exotic‐dominated grasslands

Plant invasions are an increasingly serious global concern, especially as the climate changes. Here, we explored how plant invasions differed between native‐ and novel exotic‐dominated grasslands with experimental addition of summer precipitation in Texas in 2009. Exotic species greened up earlier than natives by an average of 18 days. This was associated with a lower invasion rate early in the growing season compared to native communities. However, invasion rate did not differ significantly between native and exotic communities across all sampling times. The predictors of invasion rate differed between native and exotic communities, with invasion being negatively influenced by species richness in natives and by dominant species in exotics. Interestingly, plant invasions matched the bimodal pattern of precipitation in Temple, Texas, and did not respond to the pulse of precipitation during the summer. Our results suggest that we will need to take different approaches in understanding of invasion between native and exotic grasslands. Moreover, with anticipated increasing variability in precipitation under global climate change, plant invasions may be constrained in their response if the precipitation pulses fall outside the normal growing period of invaders.     

Logging,exotic plant invasions,and native plant reassembly in a lowland tropical rain forest

Habitat modification and biological invasions are key drivers of global environmental change. However, the extent and impact of exotic plant invasions in modified tropical landscapes remain poorly understood. We examined whether logging drives exotic plant invasions and whether their combined influences alter understory plant community composition in lowland rain forests in Borneo. We tested the relationship between understory communities and local‐ and landscape‐scale logging intensity, using leaf area index (LAI) and aboveground biomass (AGB) data from 192 plots across a logging‐intensity gradient from primary to repeatedly logged forests. Overall, we found relatively low levels of exotic plant invasions, despite an intensive logging history. Exotic species were more speciose, had greater cover, and more biomass in sites with more local‐scale canopy loss. Surprisingly, though, exotic species invasion was not related to either landscape‐scale canopy loss or road configuration. Moreover, logging and invasion did not seem to be acting synergistically on native plant composition, except that seedlings of the canopy‐dominant Dipterocarpaceae family were less abundant in areas with higher exotic plant biomass. Current low levels of invasion, and limited association with native understory community change, suggest there is a window of opportunity to manage invasive impacts. We caution about potential lag effects and the possibly severe negative impacts of exotic plant invasions on the long‐term quality of tropical forest, particularly where agricultural plantations function as permanent seed sources for recurrent dispersal along logging roads. We therefore urge prioritization of strategic management plans to counter the growing threat of exotic plant invasions in modified tropical landscapes.     

Climate change and the invasion of California by grasses

Over the next century, changes in the global climate are expected to have major consequences for plant communities, possibly including the exacerbation of species invasions. We evaluated this possibility in the grass flora of California, which is economically and ecologically important and heavily invaded. We used a novel, trait‐based approach involving two components: identifying differences in trait composition between native and exotic components of the grass flora and evaluating contemporary trait–climate relationships across the state. The combination of trait–climate relationships and trait differences between groups allows us to predict changes in the exotic‐native balance under climate change scenarios. Exotic species are more likely to be annual, taller, with larger leaves, larger seeds, higher specific leaf area, and higher leaf N percentage than native species. Across the state, all these traits are associated with regions with higher temperature. Therefore, we predict that increasing temperatures will favor trait states that tend to be possessed by exotic species, increasing the dominance of exotic species. This prediction is corroborated by the current distribution of exotic species richness relative to native richness in California; warmer areas contain higher proportions of exotic species. This pattern was very well captured by a simple model that predicts invasion severity given only the trait–climate relationship for native species and trait differences between native and exotic species. This study provides some of the first evidence for an important interaction between climate change and species invasions across very broad geographic and taxonomic scales.     

Minimal Effects of an Invasive Flowering Shrub on the Pollinator Community of Native Forbs

Biological invasions can strongly influence species interactions such as pollination. Most of the documented effects of exotic plant species on plant-pollinator interactions have been observational studies using single pairs of native and exotic plants, and have focused on dominant exotic plant species. We know little about how exotic plants alter interactions in entire communities of plants and pollinators, especially at low to medium invader densities. In this study, we began to address these gaps by experimentally removing the flowers of a showy invasive shrub, Rosa multiflora, and evaluating its effects on the frequency, richness, and composition of bee visitors to co-flowering native plants. We found that while R. multiflora increased plot-level richness of bee visitors to co-flowering native plant species at some sites, its presence had no significant effects on bee visitation rate, visitor richness, bee community composition, or abundance overall. In addition, we found that compared to co-flowering natives, R. multiflora was a generalist plant that primarily received visits from generalist bee species shared with native plant species. Our results suggest that exotic plants such as R. multiflora may facilitate native plant pollination in a community context by attracting a more diverse assemblage of pollinators, but have limited and idiosyncratic effects on the resident plant-pollinator network in general.     

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.     

Negative native–exotic diversity relationship in oak savannas explained by human influence and climate

Recent research has proposed a scale-dependence to relationships between native diversity and exotic invasions. At fine spatial scales, native–exotic richness relationships should be negative as higher native richness confers resistance to invasion. At broad scales, relationships should be positive if natives and exotics respond similarly to extrinsic factors. Yet few studies have examined both native and exotic richness patterns across gradients of human influence, where impacts could affect native and exotic species differently. We examined native–exotic richness relationships and extrinsic drivers of plant species richness and distributions across an urban development gradient in remnant oak savanna patches. In sharp contrast to most reported results, we found a negative relationship at the regional scale, and no relationship at the local scale. The negative regional-scale relationship was best explained by extrinsic factors, surrounding road density and climate, affecting natives and exotics in opposite ways, rather than a direct effect of native on exotic richness, or vice versa. Models of individual species distributions also support the result that road density and climate have largely opposite effects on native and exotic species, although simple life history traits (life form, dispersal mode) do not predict which habitat characteristics are important for particular species. Roads likely influence distributions and species richness by increasing both exotic propagule pressure and disturbance to native species. Climate may partially explain the negative relationship due to differing climatic preferences within the native and exotic species pools. As gradients of human influence are increasingly common, negative broad-scale native–exotic richness relationships may be frequent in such landscapes.     

Beyond biodiversity: individualistic controls of invasion in a self-assembled community

Recent experimental and simulation results, and competition‐based ecological theory, predict a simple relationship between species richness and the invasibility of communities at small spatial scales – likelihood of invasion decreases with increasing richness. Here we show data from 42 continuous years of sampling old field succession that reveal quite different dynamics of plant invasion. Contrary to experimental studies, when richness was important in explaining invasion probability, it was typically positively associated with species invasion. Invasion of several species had a unimodal response to resident species richness, which appeared to be a mixture of compositional influences and a richness effect. Interestingly, invasions by native and exotic species did not fundamentally differ. Control of species invasion in this system is individualistic, caused by a variety of community‐level mechanisms rather than a single prevailing richness effect.     

Community-level consequences of invasion: impacts of exotic clonal plants on riparian vegetation

Biological invasions by exotic species are occurring at exceptional rates and spatial scales worldwide and are increasingly recognized as key forms of global environmental change. Despite this growing prominence, surprisingly few ecological studies have quantified the impacts of exotic taxa on the plant communities they invade, and this is especially evident in riparian ecosystems. Along the Russian River in northern California, we used both comparative and experimental studies to investigate the influence of two exotic clonal plant species—giant reed (Arundo donax) and blue periwinkle (Vinca major)—on the composition of riparian plant communities. Our results indicate that Arundo invasion was associated with significantly lower richness of native perennial plant species on stream banks and floodplains, whereas there was no relationship on gravel bars. Additional research showed that plots invaded by Arundo and Vinca, both individually and collectively, exhibited significantly lower native and exotic species richness and abundance of both established plants and seedlings than uninvaded plots. Finally, after 2 years, experimental reductions of Arundo biomass via cutting and herbicide resulted in significantly increased native plant species richness and abundances of both established plants and seedlings, while having no effects on other exotics. In summary, our results indicate that Arundo and Vinca have strongly negative effects on diverse components of a riparian plant community, which must be addressed via effective control and restoration efforts.     

Management strategies for plant invasions: manipulating productivity, disturbance, and competition

The traditional approach to understanding invasions has focused on properties of the invasive species and of the communities that are invaded. A well‐established concept is that communities with higher species diversity should be more resistant to invaders. However, most recently published field data contradict this theory, finding instead that areas with high native plant diversity also have high exotic plant diversity. An alternative environment‐based approach to understanding patterns of invasions assumes that native and exotic species respond similarly to environmental conditions, and thus predicts that they should have similar patterns of abundance and diversity. Establishment and growth of native and exotic species are predicted to vary in response to the interaction of plant growth rates with the frequency and intensity of mortality‐causing disturbances. This theory distinguishes between the probability of establishment and the probability of dominance, predicting that establishment should be highest under unproductive and undisturbed conditions and also disturbed productive conditions. However, the probability of dominance by exotic species, and thus of potential negative impacts on diversity, is highest under productive conditions. The theory predicts that a change in disturbance regime can have opposite effects in environments with contrasting levels of productivity. Manipulation of productivity and disturbance provides opportunities for resource managers to influence the interactions among species, offering the potential to reduce or eliminate some types of invasive species.     

All is not loss: plant biodiversity in the anthropocene

Anthropogenic global changes in biodiversity are generally portrayed in terms of massive native species losses or invasions caused by recent human disturbance. Yet these biodiversity changes and others caused directly by human populations and their use of land tend to co-occur as long-term biodiversity change processes in the Anthropocene. Here we explore contemporary anthropogenic global patterns in vascular plant species richness at regional landscape scales by combining spatially explicit models and estimates for native species loss together with gains in exotics caused by species invasions and the introduction of agricultural domesticates and ornamental exotic plants. The patterns thus derived confirm that while native losses are likely significant across at least half of Earth's ice-free land, model predictions indicate that plant species richness has increased overall in most regional landscapes, mostly because species invasions tend to exceed native losses. While global observing systems and models that integrate anthropogenic species loss, introduction and invasion at regional landscape scales remain at an early stage of development, integrating predictions from existing models within a single assessment confirms their vast global extent and significance while revealing novel patterns and their potential drivers. Effective global stewardship of plant biodiversity in the Anthropocene will require integrated frameworks for observing, modeling and forecasting the different forms of anthropogenic biodiversity change processes at regional landscape scales, towards conserving biodiversity within the novel plant communities created and sustained by human systems.     

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.     

Differentiating between effects of invasion and diversity: impacts of aboveground plant communities on belowground fungal communities

Exotic plant species can affect soil microbial communities with the potential for community and ecosystem feedbacks. Yet, separating the effects of exotics from confounded changes in plant community diversity still remains a challenge. We focused on how plant diversity and native or exotic life history affected root fungi because of their significant roles in community and ecosystem processes. Specifically, we examined how fungi colonizing plant roots were affected by plant richness (one, two or four species) replicated across a range of plant community mixtures (natives, exotics, native-exotic mixtures). Fungal biomass inside roots was affected independently by plant richness and mixture, while root fungal community composition was affected only by plant richness. Extraradical networks also increased in size with plant richness. By contrast, plant biomass was a function of plant mixture, with natives consistently smaller than exotics and native-exotic mixtures intermediate. Plant invasions may have an impact on the belowground community primarily through their effects on diversity, at least in the short-term. Disentangling the effects of diversity and invasion on belowground microbial communities can help us to understand both the controllers of belowground resilience and mechanisms of successful colonization and spread of exotic plants.     

From plant neighbourhood to landscape scales: how grazing modifies native and exotic plant species richness in grassland

The interactive effect of grazing and soil resources on plant species richness and coexistence has been predicted to vary across spatial scales. When resources are not limiting, grazing should reduce competitive effects and increase colonisation and richness at fine scales. However, at broad scales richness is predicted to decline due to loss of grazing intolerant species. We examined these hypotheses in grasslands of southern Australia that varied in resources and ungulate grazing intensity since farming commenced 170 years ago. Fine-scale species richness was slightly greater in more intensively grazed upper slope sites with high nutrients but low water supply compared to those that were moderately grazed, largely due to a greater abundance of exotic species. At broader scales, exotic species richness declined with increasing grazing intensity whether nutrients or water supply were low or high. Native species richness declined at all scales in response to increasing grazing intensity and greater resource supply. Grazing also reduced fine-scale heterogeneity in native species richness and although exotics were also characterised by greater heterogeneity at fine scales, grazing effects varied across scales. In these grasslands patterns of plant species richness did not match predictions at all scales and this is likely to be due to differing responses of native and exotic species and their relative abundance in the regional species pool. Over the past 170 years intolerant native species have been eliminated from areas that are continually and heavily grazed, whereas transient, light grazing increases richness of both exotics and natives. The results support the observation that the processes and scales at which they operate differ between coevolved ungulate—grassland systems and those in transition due to recent invasion of herbivores and associated plant species.     

Rapid short-tussock grassland decline with and without grazing, Marlborough, New Zealand

Species abundance, species richness, and ground cover were measured over 10 years on nine paired grazed and exclosure plots in short-tussock grassland in the early stages of invasion by Hieracium species. With and without grazing, H. pilosella and H. caespitosum increased markedly and H. lepidulum increased locally. In contrast, 50% of all other common species and species groups, and total, native, and exotic species richness declined significantly. Exclusion increased or had no effect on rates of increase in Hieracium species and rates of decline in short tussocks, and did not reduce rates of decline in other species. Exclusion had no effect on decline in native species richness, but mainly accelerated declines in total and exotic richness. Declines in 13 key vegetation variables were significantly predicted by increase in Hieracium abundance, suggesting competitive exclusion. With or without grazing, Hieracium species will become more dominant and other species will continue to decline. The effects of large herbivores on plant species diversity can often be predicted from site productivity. Our results indicate the need also to account for species origin, spatial scale, time, and exotic invasion.