Volatile chemicals from leaf litter are associated with invasiveness of a neotropical weed in Asia

Some invasive plant species appear to strongly suppress neighbors in their nonnative ranges but much less so in their native range. We found that in the field in its native range in Mexico, the presence of Ageratina adenophora, an aggressive Neotropical invader, was correlated with higher plant species richness than found in surrounding plant communities where this species was absent, suggesting facilitation. However, in two nonnative ranges, China and India, A. adenophora canopies were correlated with much lower species richness than the surrounding communities, suggesting inhibition. Volatile organic compound (VOC) signals may contribute to this striking biogeographical difference and the invasive success of A. adenophora. In controlled experiments volatiles from A. adenophora litter caused higher mortality of species native to India and China, but not of species native to Mexico. The effects of A. adenophora VOCs on seedling germination and growth did not differ between species from the native range and species from the nonnative ranges of the invader. Litter from A. adenophora plants from nonnative populations also produced VOCs that differed quantitatively in the concentrations of some chemicals than litter from native populations, but there were no chemicals unique to one region. Biogeographic differences in the concentrations of some volatile compounds between ranges suggest that A. adenophora may be experiencing selection on biochemical composition in its nonnative ranges.     

Comparative water use of native and invasive plants at multiple scales: a global meta-analysis

Ecohydrology and invasive ecology have become increasingly important in the context of global climate change. This study presents the first in-depth analysis of the water use of invasive and native plants of the same growth form at multiple scales: leaf, plant, and ecosystem. We reanalyzed data for several hundred native and invasive species from over 40 published studies worldwide to glean global trends and to highlight how patterns vary depending on both scale and climate. We analyzed all pairwise combinations of co-occurring native and invasive species for higher comparative resolution of the likelihood of an invasive species using more water than a native species and tested for significance using bootstrap methods. At each scale, we found several-fold differences in water use between specific paired invasive and native species. At the leaf scale, we found a strong tendency for invasive species to have greater stomatal conductance than native species. At the plant scale, however, natives and invasives were equally likely to have the higher sap flow rates. Available data were much fewer for the ecosystem scale; nevertheless, we found that invasive-dominated ecosystems were more likely to have higher sap flow rates per unit ground area than native-dominated ecosystems. Ecosystem-scale evapotranspiration, on the other hand, was equally likely to be greater for systems dominated by invasive and native species of the same growth form. The inherent disconnects in the determination of water use when changing scales from leaf to plant to ecosystem reveal hypotheses for future studies and a critical need for more ecosystem-scale water use measurements in invasive- vs. native-dominated systems. The differences in water use of native and invasive species also depended strongly on climate, with the greater water use of invasives enhanced in hotter, wetter climates at the coarser scales.     

Invasion through quantitative effects: intense shade drives native decline and invasive success.

The effects of invasive nonnative species on community composition are well documented. However, few studies have determined the mechanisms by which invaders drive these changes. The literature indicates that many nonnative plant species alter light availability differently than natives in a given community, suggesting that shading may be such a mechanism. We compared light quantity (photosynthetically active radiation, PAR) and quality (red: far-red ratio, R:Fr) in riparian reaches heavily invaded by a nonnative tree (Acer platanoides) to that in an uninvaded forest and experimentally tested the effects of our measured differences in PAR and R:Fr on the survival, growth, and biomass allocation of seedlings of the dominant native species and Acer platanoides. Light conditions representative of the understory of Acer platanoides-invaded forest decreased survival of the native maple Acer glabrum by 28%; Amelanchier alnifolia by 32%; Betula occidentalis by 55%; Elymus glaucus by 46%; and Sorbus aucuparia by 52%, relative to seedlings growing in PAR similar to that of native understories. In contrast, Acer platanoides and the native shrub Symphoricarpos albus were not affected by reductions in PAR. Acer platanoides seedlings and saplings are uniquely adapted to shade relative to native species. Acer platanoides was the only species tested that decreased allocation to roots relative to shoots in the invaded forest vs. the native forest light conditions. Therefore it was the only species to demonstrate an adaptive response to the particular light environment associated with Acer platanoides invasion as predicted by optimal partitioning theory. The profound change in light quantity associated with Acer platanoides canopies appears to act as an important driver of native suppression and conspecific success in invaded riparian communities. Further research is necessary to determine whether the effect of nonnative plant-driven changes on light quantity and quality is a widespread mechanism negatively affecting resident species and facilitating invasion by nonnatives.     

Resolving whether botanic gardens are on the road to conservation or a pathway for plant invasions

A global conservation goal is to understand the pathways through which invasive species are introduced into new regions. Botanic gardens are a pathway for the introduction of invasive non‐native plants, but a quantitative assessment of the risks they pose has not been performed. I analyzed data on the living collections of over 3000 botanic gardens worldwide to quantify the temporal trend in the representation of non‐native species; the relative composition of threatened, ornamental, or invasive non‐native plant species; and the frequency with which botanic gardens implement procedures to address invasive species. While almost all of the world's worst invasive non‐native plants occurred in one or more living collections (99%), less than one‐quarter of red‐listed threatened species were cultivated (23%). Even when cultivated, individual threatened species occurred in few living collections (7.3), while non‐native species were on average grown in 6 times as many botanic gardens (44.3). As a result, a botanic garden could, on average, cultivate four times as many invasive non‐native species (20) as red‐listed threatened species (5). Although the risk posed by a single living collection is small, the probability of invasion increases with the number of botanic gardens within a region. Thus, while both the size of living collections and the proportion of non‐native species cultivated have declined during the 20th century, this reduction in risk is offset by the 10‐fold increase in the number of botanic gardens established worldwide. Unfortunately, botanic gardens rarely implement regional codes of conduct to prevent plant invasions, few have an invasive species policy, and there is limited monitoring of garden escapes. This lack of preparedness is of particular concern given the rapid increase in living collections worldwide since 1950, particularly in South America and Asia, and highlights past patterns of introduction will be a poor guide to determining future invasion risks.     

Plant invasions, generalist herbivores, and novel defense weapons

One commonly accepted mechanism for biological invasions is that species, after introduction to a new region, leave behind their natural enemies and therefore increase in distribution and abundance. However, which enemies are escaped remains unclear. Escape from specialist invertebrate herbivores has been examined in detail, but despite the profound effects of generalist herbivores in natural communities their potential to control invasive species is poorly understood. We carried out parallel laboratory feeding bioassays with generalist invertebrate herbivores from the native (Europe) and from the introduced (North America) range using native and nonnative tetraploid populations of the invasive spotted knapweed, Centaurea stoebe. We found that the growth of North American generalist herbivores was far lower when feeding on C. stoebe than the growth of European generalists. In contrast, North American and European generalists grew equally well on European and North American tetraploid C. stoebe plants, lending no support for an evolutionary change in resistance of North American tetraploid C. stoebe populations against generalist herbivores. These results suggest that biogeographical differences in the response of generalist herbivores to novel plant species have the potential to affect plant invasions.     

When there is no escape: the effects of natural enemies on native, invasive, and noninvasive plants

An important question in the study of biological invasions is the degree to which successful invasion can be explained by release from control by natural enemies. Natural enemies dominate explanations of two alternate phenomena: that most introduced plants fail to establish viable populations (biotic resistance hypothesis) and that some introduced plants become noxious invaders (natural enemies hypothesis). We used a suite of 18 phylogenetically related native and nonnative clovers (Trifolium and Medicago) and the foliar pathogens and invertebrate herbivores that attack them to answer two questions. Do native species suffer greater attack by natural enemies relative to introduced species at the same site? Are some introduced species excluded from native plant communities because they are susceptible to local natural enemies? We address these questions using three lines of evidence: (1) the frequency of attack and composition of fungal pathogens and herbivores for each clover species in four years of common garden experiments, as well as susceptibility to inoculation with a common pathogen; (2) the degree of leaf damage suffered by each species in common garden experiments; and (3) fitness effects estimated using correlative approaches and pathogen removal experiments. Introduced species showed no evidence of escape from pathogens, being equivalent to native species as a group in terms of infection levels, susceptibility, disease prevalence, disease severity (with more severe damage on introduced species in one year), the influence of disease on mortality, and the effect of fungicide treatment on mortality and biomass. In contrast, invertebrate herbivores caused more damage on native species in two years, although the influence of herbivore attack on mortality did not differ between native and introduced species. Within introduced species, the predictions of the biotic resistance hypothesis were not supported: the most invasive species showed greater infection, greater prevalence and severity of disease, greater prevalence of herbivory, and greater effects of fungicide on biomass and were indistinguishable from noninvasive introduced species in all other respects. Therefore, although herbivores preferred native over introduced species, escape from pest pressure cannot be used to explain why some introduced clovers are common invaders in coastal prairie while others are not.     

Herbivory by an introduced Asian weevil negatively affects population growth of an invasive Brazilian shrub in Florida

The enemy release hypothesis (ERH) is often cited to explain why some plants successfully invade natural communities while others do not. This hypothesis maintains that plant populations are regulated by coevolved enemies in their native range but are relieved of this pressure where their enemies have not been co-introduced. Some studies have shown that invasive plants sustain lower levels of herbivore damage when compared to native species, but how damage affects fitness and population dynamics remains unclear. We used a system of co-occurring native and invasive Eugenia congeners in south Florida (USA) to experimentally test the ERH, addressing deficiencies in our understanding of the role of natural enemies in plant invasion at the population level. Insecticide was used to experimentally exclude insect herbivores from invasive Eugenia uniflora and its native co-occurring congeners in the field for two years. Herbivore damage, plant growth, survival, and population growth rates for the three species were then compared for control and insecticide-treated plants. Our results contradict the ERH, indicating that E. uniflora sustains more herbivore damage than its native congeners and that this damage negatively impacts stem height, survival, and population growth. In addition, most damage to E. uniflora, a native of Brazil, is carried out by Myllocerus undatus, a recently introduced weevil from Sri Lanka, and M. undatus attacks a significantly greater proportion of E. uniflora leaves than those of its native congeners. This interaction is particularly interesting because M. undatus and E. uniflora share no coevolutionary history, having arisen on two separate continents and come into contact on a third. Our study is the first to document negative population-level effects for an invasive plant as a result of the introduction of a novel herbivore. Such inhibitory interactions are likely to become more prevalent as suites of previously noninteracting species continue to accumulate and new communities assemble worldwide.     

Diversity-invasibility across an experimental disturbance gradient in Appalachian Forests

Research examining the relationship between community diversity and invasions by nonnative species has raised new questions about the theory and management of biological invasions. Ecological theory predicts, and small-scale experiments confirm, lower levels of nonnative species invasion into species-rich compared to species-poor communities, but observational studies across a wider range of scales often report positive relationships between native and nonnative species richness. This paradox has been attributed to the scale dependency of diversity-invasibility relationships and to differences between experimental and observational studies. Disturbance is widely recognized as an important factor determining invasibility of communities, but few studies have investigated the relative and interactive roles of diversity and disturbance on nonnative species invasion. Here, we report how the relationship between native and nonnative plant species richness responded to an experimentally applied disturbance gradient (from no disturbance up to clearcut) in oak-dominated forests. We consider whether results are consistent with various explanations of diversity-invasibility relationships including biotic resistance, resource availability, and the potential effects of scale (1 m2 to 2 ha). We found no correlation between native and nonnative species richness before disturbance except at the largest spatial scale, but a positive relationship after disturbance across scales and levels of disturbance. Post-disturbance richness of both native and nonnative species was positively correlated with disturbance intensity and with variability of residual basal area of trees. These results suggest that more nonnative plants may invade species-rich communities compared to species-poor communities following disturbance.     

Potential Interference Between a Threatened Endemic Thistle and an Invasive Nonnative Plant

Although the establishment of nonnative plants is recognized as a threat to native ecosystems, there are few documented examples of an invasive species directly influencing a rare native plant. The Eurasian biennial Dipsacus sylvestris (teasel) is invading the central New Mexico habitat of Cirsium vinaceum , an endemic thistle that is federally listed as threatened. We documented changes in teasel distribution and abundance between 1989 and 1993 that suggest the potential for direct interactions with the native thistle. We then compared habitat characteristics, germination behavior, and performance in greenhouse and field competition trials to evaluate the potential outcome of interference between these two species. There were no significant differences in measured habitat characteristics between sites supporting C. vinaceum and those with D. sylvestris. Dipsacus was better able to germinate in low light than the thistle, suggesting that D. sylvestris might invade C. vinaceum populations but that thistle recruitment would be unlikely in dense stands of the nonnative plants. In the greenhouse growth of C. vinaceum rosettes was significantly reduced by the presence of Dipsacus , but the invader was unaffected by the thistle; results of a short-term field experiment were equivocal but suggestive of interference between the two. We suggest criteria for managers to use in determining whether invading species pose problems for specific rare native taxa, and we discuss the constraints on experimental work where protected taxa are involved.     

Variation in resource availability changes the impact of invasive thistles on native bunchgrasses.

The threat posed by invasive nonnative plants to native plant populations is one of the largest challenges facing both conservation biology and restoration ecology. California has been highly impacted by invaders, although many relict stands of native plants are found on shallow, rocky soils with limited resources. The abiotic conditions of these sites may strongly influence the performance of an invasive plant and its effect on resident native species. In addition, the maturity of native plants in these sites may modulate an invader's impact; larger, well-established plants may be better able to resist invaders. In this study we examined how the impact of an invasive thistle (Centaurea solstitialis) on a native perennial bunchgrass (Nassella pulchra) changed in response to variation in soil depth, soil water availability, and bunchgrass maturity. We measured plant performance in terms of survival, growth, reproduction, and predawn water potential. We found that soil depth, water availability, and bunchgrass maturity acted in concert to influence the impact of the invasive thistle on the native bunchgrass. Both species performed better in deep soils, especially during dry years. The combination of shallow soil and low water availability reduced C. solstitialis performance and ameliorated its negative effect on N. pulchra growth and reproduction. Higher water availability resulted in a stronger negative effect of C. solstitialis on N. pulchra in both shallow and deep soils. However, as N. pulchra matured and increased in size, we saw a steady decline in C. solstitialis growth and reproductive output. Higher water availability increased the performance of C. solstitialis in shallow soils. C. solstitialis may thus have a stronger impact on N. pulchra and be more able to invade relict stands of N. pulchra in shallow soils during high-rainfall years. However, established stands of N. pulchra appear to be more resistant to invasion by C. solstitialis as N. pulchra plants grow older and larger.     

Spatial heterogeneity influences native and nonnative plant species richness

Spatial heterogeneity may have differential effects on the distribution of native and nonnative plant species richness. We examined the effects of spatial heterogeneity on native and nonnative plant species richness distributions in the central part of Rocky Mountain National Park, Colorado, USA. Spatial heterogeneity around vegetation plots was characterized using landscape metrics, environmental/topographic variables (slope, aspect, elevation, and distance from stream or river), and soil variables (nitrogen, clay, and sand). The landscape metrics represented five components of landscape heterogeneity and were measured at four spatial extents (within varying radii of 120, 240, 480, and 960 m) using the FRAGSTATS landscape pattern analysis program. Akaike's Information Criterion adjusted for small sample size (AICc) was used to select the best models from a set of multiple linear regression models developed for native and nonnative plant species richness at four spatial extents and three levels of ecological hierarchy (i.e., landscape, land cover, and community). Both native and nonnative plant species richness were positively correlated with edge density, Simpson's diversity index and interspersion/juxtaposition index, and were negatively correlated with mean patch size. The amount of variation explained at four spatial extents and three hierarchical levels ranged from 30% to 70%. At the landscape level, the best models explained 43% of the variation in native plant species richness and 70% of the variation in nonnative plant species richness (240-m extent). In general, the amount of variation explained was always higher for nonnative plant species richness, and the inclusion of landscape metrics always significantly improved the models. The best models explained 66% of the variation in nonnative plant species richness for both the conifer land cover type and lodgepole pine community. The relative influence of the components of spatial heterogeneity differed for native and nonnative plant species richness and varied with the spatial extent of analysis and levels of ecological hierarchy. The study offers an approach to quantify spatial heterogeneity to improve models of plant biodiversity. The results demonstrate that ecologists must recognize the importance of spatial heterogeneity in managing native and nonnative plant species.     

Effect of Invasive Plant Species on Temperate Wetland Plant Diversity

Abstract:  Invasive species are a major threat to global biodiversity and an important cause of biotic homogenization of ecosystems. Exotic plants have been identified as a particular concern because of the widely held belief that they competitively exclude native plant species. We examined the correlation between native and invasive species richness in 58 Ontario inland wetlands. The relationship between exotic and native species richness was positive even when we controlled for important covarying factors. In addition, we examined the relationship between the abundance of four native species (  Typha latifolia, T. angustifolia, Salix petiolaris, Nuphar variegatum ) and four invasive species (  Lythrum salicaria, Hydrocharis morsus-ranae, Phalaris arundinacea, Rhamnus frangula ) that often dominate temperate wetlands and native and rare native species richness. Exotic species were no more likely to dominate a wetland than native species, and the proportion of dominant exotic species that had a significant negative effect on the native plant community was the same as the proportion of native species with a significant negative effect. We conclude that the key to conservation of inland wetland biodiversity is to discourage the spread of community dominants, regardless of geographical origin.     

Windows of opportunity: historical and ecological controls on Berberis thunbergii invasions

Attempts to determine characteristics that render habitats invasible to nonnative species have met with limited success. This may be because most studies focus on modern habitat conditions and do not consider invasibility in the context of a historically dynamic landscape in which both the abundance of a species and the invasibility of a site may change. We surveyed 159 currently forested sites for the occurrence and abundance of Berberis thunbergii (Japanese barberry), an invasive, nonnative shrub in forests of the northeastern United States, relative to modern environmental conditions, contemporary logging activity, and two periods of historical land use. Berberis thunbergii occurred more frequently and was more abundant in post-agricultural forests than in continuously wooded sites. This relationship was stronger for agricultural sites that were abandoned and reforested after B. thunbergii was introduced to the region than for sites that reforested prior to B. thunbergii introduction. In contrast, recent forest harvesting did not influence the occurrence or abundance of B. thunbergii. Modern soil fertility explained a significant portion of the variation in B. thunbergii occurrence, whereas site history considerably improved predictions of population density and helped evaluate potential invasion mechanisms. While land-use history covaries with soil fertility and distance to putative seed sources, the strong relationship between modern abundance patterns and historical agriculture suggests that B. thunbergii colonized recently abandoned agricultural lands in the early 20th century and then persisted and spread locally during subsequent reforestation. Our results indicate that interpretations of both native community composition and modern plant invasions must consider the importance of historical landscape changes and the timing of species introduction along with current environmental conditions.     

Ranking Lepidopteran Use of Native Versus Introduced Plants

Abstract:  In light of the wide-scale replacement of native plants in North America with introduced, invasive species and noninvasive ornamental plants that evolved elsewhere, we compared the value of native and introduced plants in terms of their ability to serve as host plants for Lepidoptera. Insect herbivores such as Lepidoptera larvae are critically important components of terrestrial food webs and any reduction in their biomass or diversity due to the loss of acceptable host plants is predicted to reduce the production of the many insectivores in higher trophic levels. We conducted an exhaustive search of host records in the literature. We used the data we gathered to rank all 1385 plant genera that occur in the mid-Atlantic states of the United States by their ability to support Lepidoptera richness. Statistical comparisons were made with Welch's test for equality of means. Woody plants supported more species of moths and butterflies than herbaceous plants, native plants supported more species than introduced plants, and native woody plants with ornamental value supported more Lepidoptera species than introduced woody ornamentals. All these differences were highly significant. Our rankings provide a relative measure that will be useful for restoration ecologists, landscape architects and designers, land managers, and landowners who wish to raise the carrying capacity of particular areas by selecting plants with the greatest capacity for supporting biodiversity.     

Traits, not origin, explain impacts of plants on larval amphibians

Managing habitats for the benefit of native fauna is a priority for many government and private agencies. Often, these agencies view nonnative plants as a threat to wildlife habitat, and they seek to control or eradicate nonnative plant populations. However, little is known about how nonnative plant invasions impact native fauna, and it is unclear whether managing these plants actually improves habitat quality for resident animals. Here, we compared the impacts of native and nonnative wetland plants on three species of native larval amphibians; we also examined whether plant traits explain the observed impacts. Specifically, we measured plant litter quality (carbon : nitrogen : phosphorus ratios, and percentages of lignin and soluble phenolics) and biomass, along with a suite of environmental conditions known to affect larval amphibians (hydroperiod, temperature, dissolved oxygen, and pH). Hydroperiod and plant traits, notably soluble phenolics, litter C:N ratio, and litter N:P ratio, impacted the likelihood that animals metamorphosed, the number of animals that metamorphosed, and the length of larval period. As hydroperiod decreased, the likelihood that amphibians achieved metamorphosis and the percentage of tadpoles that successfully metamorphosed also decreased. Increases in soluble phenolics, litter N:P ratio, and litter C:N ratio decreased the likelihood that tadpoles achieved metamorphosis, decreased the percentage of tadpoles metamorphosing, decreased metamorph production (total metamorph biomass), and increased the length of larval period. Interestingly, we found no difference in metamorphosis rates and length of larval period between habitats dominated by native and nonnative plants. Our findings have important implications for habitat management. We suggest that to improve habitats for native fauna, managers should focus on assembling a plant community with desirable traits rather than focusing only on plant origin.     

Homogenization dynamics and introduction routes of invasive freshwater fish in the Iberian Peninsula.

Nonnative invasive species are one of the main global threats to biodiversity. The understanding of the traits characterizing successful invaders and invasion-prone ecosystems is increasing, but our predictive ability is still limited. Quantitative information on biotic homogenization and particularly its temporal dynamics is even scarcer. We used freshwater fish distribution data in the Iberian Peninsula in four periods (before human intervention, 1991, 1995, and 2001) to assess the temporal dynamics of biotic homogenization among river basins. The percentage of introduced species among fish faunas has increased in recent times (from 41.8% in 1991 to 52.5% in 2001), leading to a clear increase in the similarity of community composition among basins. The mean Jaccard's index increase (a measure of biotic homogenization) from the pristine situation to the present (17.1%) was similar to that for Californian fish but higher than for other studies. However, biotic homogenization was found to be a temporally dynamic process, with finer temporal grain analyses detecting transient stages of biotic differentiation. Introduced species assemblages were spatially structured along a latitudinal gradient in the Iberian Peninsula, with species related to sport fishing being characteristic of northern basins. Although the comparison of fish distributions in the Iberian Peninsula and France showed significant and generalized biotic homogenization, nonnative assemblages of northeastern Iberian basins were more similar to those of France than to those of the rest of the Iberian Peninsula, indicating a main introduction route. Species introduced to the Iberian Peninsula tended to be mainly piscivores or widely introduced species that previously had been introduced to France. Our results indicate that the simultaneous analysis of the spatial distribution of introduced assemblages (excluding native species that reflect other biogeographical patterns) and their specific traits can be an effective tool to detect introduction and invasion routes and to predict future invaders from donor regions.     

Effects of Climate Change,Invasive Species,and Disease on the Distribution of Native European Crayfishes

Climate change will require species to adapt to new conditions or follow preferred climates to higher latitudes or elevations, but many dispersal‐limited freshwater species may be unable to move due to barriers imposed by watershed boundaries. In addition, invasive nonnative species may expand into new regions under future climate conditions and contribute to the decline of native species. We evaluated future distributions for the threatened European crayfish fauna in response to climate change, watershed boundaries, and the spread of invasive crayfishes, which transmit the crayfish plague, a lethal disease for native European crayfishes. We used climate projections from general circulation models and statistical models based on Mahalanobis distance to predict climate‐suitable regions for native and invasive crayfishes in the middle and at the end of the 21st century. We identified these suitable regions as accessible or inaccessible on the basis of major watershed boundaries and present occurrences and evaluated potential future overlap with 3 invasive North American crayfishes. Climate‐suitable areas decreased for native crayfishes by 19% to 72%, and the majority of future suitable areas for most of these species were inaccessible relative to native and current distributions. Overlap with invasive crayfish plague‐transmitting species was predicted to increase. Some native crayfish species (e.g., noble crayfish [Astacus astacus]) had no future refugia that were unsuitable for the modeled nonnative species. Our results emphasize the importance of preventing additional introductions and spread of invasive crayfishes in Europe to minimize interactions between the multiple stressors of climate change and invasive species, while suggesting candidate regions for the debatable management option of assisted colonization. Efectos del Cambio Climático, Especies Invasoras y Enfermedades sobre la Distribución de Cangrejos de Río Europeos Nativos     

When is an invasive not an invasive? Macrofossil evidence of doubtful native plant species in the Galápagos Islands

The Galápagos Islands are globally renowned for their ecological value and as a world symbol of scientific discovery; however the native biodiversity of this unique region is currently under threat. One of the primary concerns is the detrimental impact of approximately 750 nonnative plants introduced over the last 500 years of human presence in the archipelago. In addition to these known introduced species, there are an additional 62 vascular plants classified as "doubtful natives," where native status remains unclear. To help address the questions of provenance regarding these doubtfully native species and their impact on highland ecosystems over the past 500-1000 years, we analyzed plant macrofossils in sedimentary records. Appropriate species classification (native or introduced) was determined using baseline data of species presence on the islands. We confirmed that six plants (Ageratum conyzoides, Solanum americanum, Ranunculus flagelliformis, Brickellia diffusa, Galium canescens, and Anthephora hermaphrodita) once considered doubtful natives or introduced are actually native to the Galápagos flora. These results have relevance not just for the Galápagos but also many other oceanic islands in demonstrating the application of palaeobotanical data to conserving and restoring native biodiversity.     

Long-term population dynamics of seeded plants in invaded grasslands

In recent decades, dozens of studies have involved attempts to introduce native and desirable nonnative plant species into grasslands dominated by invasive weeds. The newly introduced plants have proved capable of establishing, but because they are rarely monitored for more than four years, it is unknown if they have a high likelihood of persisting and suppressing invaders for the long-term. Beyond invaded grasslands, this lack of long-term monitoring is a general problem plaguing efforts to reintroduce a range of taxa into a range of ecosystems. We introduced species from seed and then periodically measured plant abundances for nine years at one site and 15 years at a second site. To our knowledge, our 15-year data are the longest to date from a seeding experiment in invaded, never-cultivated grassland. At one site, three seeded grasses maintained high densities for three or more years, but then all or nearly all individuals died. At the second site, one grass performed similarly, but two other grasses proliferated and at least one greatly suppressed the dominant invader (Centaurea maculosa). In one study, our point estimate suggests that the seeded grass Thinopyrum intermedium reduced C. maculosa biomass by 93% 15 years after seeding. In some cases, data from three and fewer years after seeding falsely suggested that seeded species were capable of persisting within the invaded grassland. In other cases, data from as late as nine years after seeding falsely suggested seeded populations would not become large enough to suppress the invader. These results show that seeded species sometimes persist and suppress invaders for long periods, but short-term data cannot predict if, when, or where this will occur. Because short-term data are not predictive of long-term seeded species performances, additional long-term data are needed to identify effective practices, traits, and species for revegetating invaded grasslands.     

Native ecotypic variation and the role of host identity in the spread of an invasive herbivore, Cactoblastis cactorum

Environmental niche models (ENMs) have gained enormous popularity as tools to investigate potential changes in species distributions resulting from climate change and species introductions. Despite recognition that species interactions can influence the dynamics of invasion spread, most implementations of ENMs focus on abiotic factors as the sole predictors of potential range limits. Implicit in this approach is the assumption that biotic interactions are relatively unimportant, either because of scaling issues, or because fundamental and realized niches are equivalent in a species' native range. When species are introduced into exotic landscapes, changes in biotic interactions relative to the native range can lead to occupation of different regions of niche space and apparent shifts in physiological tolerances. We use an escaped biological control organism, Cactoblastis cactorum (Berg.), to assess the role of the environmental envelope as compared with patterns of host-herbivore associations based on collections made in the native range. Because all nonnative populations are derived from a single C. cactorum ecotype, we hypothesize that biotic interactions associated with this ecotype are driving the species' invasion dynamics. Environmental niche models constructed from known native populations perform poorly in predicting nonnative distributions of this species, except where there is an overlap in niche space. In contrast, genetic isolation in the native range is concordant with the observed pattern of host use, and strong host association has been noted in nonnative landscapes. Our results support the hypothesis that the apparent shift in niche space from the native to the exotic ranges results from a shift in biotic interactions, and demonstrate the importance of considering biotic interactions in assessing the risk of future spread for species whose native range is highly constrained by biotic interactions.