Among-population variation in drought responses is consistent across life stages but not between native and non-native ranges.

Understanding how widespread species adapt to variation in abiotic conditions across their ranges is fundamental to ecology. Insight may come from studying how among-population variation (APV) in the common garden corresponds with the environmental conditions of source populations. However, there are no such studies comparing native vs non-native populations across multiple life stages. We examined APV in the performance and functional traits of 59 Conyza canadensis populations, in response to drought, across large aridity gradients in the native (North America) and non-native (Eurasia) ranges in three experiments. Our treatment (dry vs wet) was applied at the recruitment, juvenile, and adult life stages. We found contrasting patterns of APV in drought responses between the two ranges. In the native range, plant performance was less reduced by drought in populations from xeric than mesic habitats, but such relationship was not apparent for non-native populations. These range-specific patterns were consistent across the life stages. The weak adaptive responses of non-native populations indicate that they can become highly abundant even without complete local adaptation to abiotic environments and suggest that long-established invaders may still be evolving to the abiotic environment. These findings may explain lag times in invasions and raise concern about future expansions.

Solidago gigantea plants from nonnative ranges compensate more in response to damage than plants from the native range.

Resistance and tolerance are two ways that plants cope with herbivory. Tolerance, the ability of a plant to regrow or reproduce after being consumed, has been studied less than resistance, but this trait varies widely among species and has considerable potential to affect the ecology of plant species. One particular aspect of tolerance, compensatory responses, can evolve rapidly in plant species; providing insight into interactions between consumers and plants. However, compensation by invasive species has rarely been explored. We compared compensatory responses to the effects of simulated herbivory expressed by plants from seven Solidago gigantea populations from the native North American range to that expressed by plants from nine populations from the nonnative European range. Populations were also collected along elevational gradients to compare ecotypic variation within and between ranges. Solidago plants from the nonnative range of Europe were more tolerant to herbivory than plants from the native range of North America. Furthermore, plants from European populations increased in total biomass and growth rate with elevation, but decreased in compensatory response. There were no relationships between elevation and growth or compensation for North American populations. Our results suggest that Solidago gigantea may have evolved to better compensate for herbivory damage in Europe, perhaps in response to a shift to greater proportion of attack from generalists. Our results also suggest a possible trade-off between rapid growth and compensation to damage in European populations but not in North American populations.

Do exotic plants lose resistance to pathogenic soil biota from their native range? A test with Solidago gigantea.

Native plants commonly suffer from strong negative plant-soil feedbacks. However, in their non-native ranges species often escape from these negative feedbacks, which indicates that these feedbacks are generated by at least partially specialized soil biota. If so, introduced plants might evolve the loss of resistance to pathogens in their former native range, as has been proposed for the loss of resistance to specialized herbivores. We compared the magnitude of plant-soil feedbacks experienced by native and exotic genotypes of the perennial forb, Solidago gigantea. Feedbacks were assessed in soil collected across 14 sites sampled across the western part of Solidago's native range in the US. Both native and exotic genotypes of Solidago suffered consistently negative and broadly similar plant-soil feedbacks when grown in North American soil. Although there was substantial variation among soils from different sites in the strength of feedbacks generated, the magnitude of feedbacks generated by North American genotypes of S. gigantea were strongly correlated with those produced in the same soil by European genotypes. Our results indicate that Solidago experiences strong negative soil feedbacks in native soil and that introduced genotypes of Solidago have not lost resistance to these negative effects of soil biota. Both genotypic and landscape-level effects can be important sources of variation in the strength of plant-soil feedbacks.

Conyza canadensis suppresses plant diversity in its nonnative ranges but not at home: a transcontinental comparison.

The impact of invasive species across their native and nonnative ranges is poorly quantified and this impedes a complete understanding of biological invasions. We compared the impact of the native North American plant, Conyza canadensis, which is invasive to Eurasia, on species richness at home and in a number of introduced regions through well replicated transcontinental field studies, glasshouse experiments and individual-based models. Our results demonstrated mostly negative relationships between C. canadensis abundance and native species richness in nonnative ranges, but either positive or no relationships in its native North American range. In glasshouse experiments, the total biomass of Conyza was suppressed more by species from its native range than by species from regions where it is nonnative, but the effects of Conyza on other species did not show a consistent biogeographical pattern. Finally, individual-based models led to the exclusion of Conyza from North American scenarios but to high abundances in scenarios with species from the nonnative ranges of Conyza. We illustrate biogeographical differences in the impact of an invader across regional scales and suggest that inherent differences in one specific aspect of competitive ability, tolerance to the effects of other species, may play some role in these differences.

Acquisition and evolution of enhanced mutualism-an underappreciated mechanism for invasive success?

Soil biota can determine plant invasiveness, yet biogeographical comparisons of microbial community composition and function across ranges are rare. We compared interactions between Conyza canadensis, a global plant invader, and arbuscular mycorrhizal (AM) fungi in 17 plant populations in each native and non-native range spanning similar climate and soil fertility gradients. We then grew seedlings in the greenhouse inoculated with AM fungi from the native range. In the field, Conyza plants were larger, more fecund, and associated with a richer community of more closely related AM fungal taxa in the non-native range. Fungal taxa that were more abundant in the non-native range also correlated positively with plant biomass, whereas taxa that were more abundant in the native range appeared parasitic. These patterns persisted when populations from both ranges were grown together in a greenhouse; non-native populations cultured a richer and more diverse AM fungal community and selected AM fungi that appeared to be more mutualistic. Our results provide experimental support for evolution toward enhanced mutualism in non-native ranges. Such novel relationships and the rapid evolution of mutualisms may contribute to the disproportionate abundance and impact of some non-native plant species.

What happens in Europe stays in Europe: apparent evolution by an invader does not help at home.

Some invasive plant species rapidly evolve greater size and/or competitive ability in their nonnative ranges. However, it is not well known whether these traits transfer back to the native range, or instead represent genotype-by-environment interactions where traits are context specific to communities in the new range where the evolution occurred. Insight into transferability vs. context specificity can be tested using experiments performed with individuals from populations from the native and nonnative ranges of exotic invasive species. Using a widespread invasive plant species in Europe, Solidago gigantea, we established reciprocal common garden experiments in the native range (Montana, North America; n = 4) and the nonnative range (Hungary, Europe; n = 4) to assess differences in size, vegetative shoot number, and herbivory between populations from the native and nonnative ranges. In a greenhouse experiment, we also tested whether the inherent competitive ability of genotypes from 15 native and 15 invasive populations differed when pitted against 11 common native North American competitors. In common gardens, plants from both ranges considered together produced five times more biomass, grew four times taller, and developed five times more rhizomes in the nonnative range garden compared to the native range garden. The interaction between plant origin and the common garden location was highly significant, with plants from Hungary performing better than plants from Montana when grown in Hungary, and plants from Montana performing better than plants from Hungary when grown in Montana. In the greenhouse, there were no differences in the competitive effects and responses of S. gigantea plants from the two ranges when grown with North American natives. Our results suggest that S. gigantea might have undergone rapid evolution for greater performance abroad, but if so, this response does not translate to greater performance at home.

Escape from natural enemies depends on the enemies, the invader, and competition.

The enemy release hypothesis (ERH) attributes the success of some exotic plant species to reduced top-down effects of natural enemies in the non-native range relative to the native range. Many studies have tested this idea, but very few have considered the simultaneous effects of multiple kinds of enemies on more than one invasive species in both the native and non-native ranges. Here, we examined the effects of two important groups of natural enemies-insect herbivores and soil biota-on the performance of Tanacetum vulgare (native to Europe but invasive in the USA) and Solidago canadensis (native to the USA but invasive in Europe) in their native and non-native ranges, and in the presence and absence of competition.In the field, we replicated full-factorial experiments that crossed insecticide, T. vulgare-S. canadensis competition, and biogeographic range (Europe vs. USA) treatments. In greenhouses, we replicated full-factorial experiments that crossed soil sterilization, plant-soil feedback, and biogeographic range treatments. We evaluated the effects of experimental treatments on T. vulgare and S. canadensis biomass.The effects of natural enemies were idiosyncratic. In the non-native range and relative to populations in the native range, T. vulgare escaped the negative effects of insect herbivores but not soil biota, depending upon the presence of S. canadensis; and S. canadensis escaped the negative effects of soil biota but not insect herbivores, regardless of competition. Thus, biogeographic escape from natural enemies depended upon the enemies, the invader, and competition. Synthesis: By explicitly testing the ERH in terms of more than one kind of enemy, more than one invader, and more than one continent, this study enhances our nuanced perspective of how natural enemies can influence the performance of invasive species in their native and non-native ranges.