Contributions from the field of population biology hold promise for understanding and managing invasiveness; invasive species also offer excellent opportunities to study basic processes in population biology. Life history studies and demographic models may be valuable for examining the introduction of invasive species and identifying life history stages where management will be most effective. Evolutionary processes may be key features in determining whether invasive species establish and spread. Studies of genetic diversity and evolutionary changes should be useful for 0066-4162/01/1215-0305$14.00 305 Annu. Rev. Ecol. Syst. 2001.32:305-332. Downloaded from www.annualreviews.org by NORTH CAROLINA STATE UNIVERSITY on 09/26/12. For personal use only. 306 SAKAI ET AL.understanding the potential for colonization and establishment, geographic patterns of invasion and range expansion, lag times, and the potential for evolutionary responses to novel environments, including management practices. The consequences of biological invasions permit study of basic evolutionary processes, as invaders often evolve rapidly in response to novel abiotic and biotic conditions, and native species evolve in response to the invasion.

Despite the increasing biological and economic impacts of invasive species, little is known about the evolutionary mechanisms that favor geographic range expansion and evolution of invasiveness in introduced species. Here, we focus on the invasive wetland grass Phalaris arundinacea L. and document the evolutionary consequences that resulted from multiple and uncontrolled introductions into North America of genetic material native to different European regions. Continental-scale genetic variation occurring in reed canarygrass' European range has been reshuffled and recombined within North American introduced populations, giving rise to a number of novel genotypes. This process alleviated genetic bottlenecks throughout reed canarygrass' introduced range, including in peripheral populations, where depletion of genetic diversity is expected and is observed in the native range. Moreover, reed canarygrass had higher genetic diversity and heritable phenotypic variation in its invasive range relative to its native range. The resulting high evolutionary potential of invasive populations allowed for rapid selection of genotypes with higher vegetative colonization ability and phenotypic plasticity. Our results show that repeated introductions of a single species may inadvertently create harmful invaders with high adaptive potential. Such invasive species may be able to evolve in response to changing climate, allowing them to have increasing impact on native communities and ecosystems in the future. More generally, multiple immigration events may thus trigger future adaptation and geographic spread of a species population by preventing genetic bottlenecks and generating genetic novelties through recombination.biological invasion ͉ genotypic diversity ͉ multiple immigration ͉ range expansion ͉ phenotypic plasticity

Plant traits-the morphological, anatomical, physiological, biochemical and phenological characteristics of plants-determine how plants respond to environmental factors, affect other trophic levels, and influence ecosystem properties and their benefits and detriments to people. Plant trait data thus represent the basis for a vast area of research spanning from evolutionary biology, community and functional ecology, to biodiversity conservation, ecosystem and landscape management, restoration, biogeography and earth system modelling. Since its foundation in 2007, the TRY database of plant traits has grown continuously. It now provides unprecedented data coverage under an open access data policy and is the main plant trait database used by the research community worldwide. Increasingly, the TRY database also supports new frontiers of trait-based plant research, including the identification of data gaps and the subsequent mobilization or measurement of new data. To support this development, in this article we evaluate the extent of the trait data compiled in TRY and analyse emerging patterns of data coverage and representativeness. Best species coverage is achieved for categorical traits-almost complete coverage for 'plant growth form'. However, most traits relevant for ecology and vegetation modelling are characterized by continuous intraspecific variation and trait-environmental relationships. These traits have to be measured on individual plants in their respective environment. Despite unprecedented data coverage, we observe a humbling lack of completeness and representativeness of these continuous traits in many aspects.We, therefore, conclude that reducing data gaps and biases in the TRY database remains a key challenge and requires a coordinated approach to data mobilization and trait measurements. This can only be achieved in collaboration with other initiatives. Geosphere-Biosphere Program (IGBP) and DIVERSITAS, the TRY database (TRY-not an acronym, rather a statement of sentiment; https ://www.try-db.org; Kattge et al., 2011) was proposed with the explicit assignment to improve the availability and accessibility of plant trait data for ecology and earth system sciences. The Max Planck Institute for Biogeochemistry (MPI-BGC) offered to host the database and the different groups joined forces for this community-driven program. Two factors were key to the success of TRY: the support and trust of leaders in the field of functional plant ecology submitting large databases and the long-term funding by the Max Planck Society, the MPI-BGC and the German Centre for Integrative Biodiversity Research (iDiv) Halle-Jena-Leipzig, which has enabled the continuous development of the TRY database.

The presence of leaf litter of different depths within a tropical forest creates many different microsites for plant establishment. The amount and distribution of leaf litter within a forest can influence patterns of plant establishment. In this study, we determined the spatial variability in leaf litter in the forest understory, and investigated how different litter depths (bare, 1, 6, and 12 cm) affected the establishment of several tropical tree species in both growth house (sun and shade) and field (gap and understory) experiments in the semideciduous tropical forest of Barro Colorado Island, Panama. The tree species used in this study (Aspidospermum cruenta, Ceiba pentandra, Cordia alliodora, Gustavia Superba, Luehea semmannii, Ochroma pyrimidale) were chosen to represent a range of seed masses and a gradient in the light requirement for establishment of the species. The spatial distribution of leaf litter was not correlated between adjacent sampling points within the forest understory, suggesting that the establishment environment for seedlings, with respect to litter, is highly variable at scales of 1—20 m. The presence of litter affected five of the six species, but the nature and the magnitude of the effect were species specific. The smaller seeded shade—intolerant species had fewer seedlings establishing under leaf litter than on bare ground. The species ranged from strongly nagatively affected (Luehae) to moderately negatively affected (Cordia, Ochroma) to affected only by extreme amounts of litter (Ceiba). The presence of litter influenced Gustavia, one of the larger seeded shade—tolerant species, but did not affect Aspidospermum, the other large seeded species. The effect of litter on Gustavia depended on the light environment. Gustavia had more seedlings establishing under litter in the sun, but the presence of litter had no effect in the shade. Differences among the smaller seeded shade—intolerant species in the amount they were negatively influenced by litter were not correlated with seed mass. Data from our field study were consistent with our growth house results for the shade—intolerant species. Additional data from the field study indicated that these species with similar habitat requirements differed in the developmental stage at which they were affected by the presence of litter. Luehea had fewer seeds germinating under litter while the other two species, Ochroma and Cordia, were affected only after germination. Interspecific comparisons done for each light level and litter depth indicated that the presence of litter caused reversals in the relative ranking of species success. For example, Gustavia preferentially established under relatively deep litter depths in the sun where Luehea could not establish. In conclusion, the presence of litter can potentially increase seedling diversity within the forest by creating heterogeneity in the establishment environment and by causing reversals in species' rankings.

Aim A major challenge for invasion ecology is to identify high-impact invaders to guide prioritization of management interventions. We argue that species with the potential to cause regime shifts (altered states of ecosystem structure and function that are difficult or impossible to reverse) should be prioritized. These are species that modify ecosystems in ways that enhance their own persistence and suppress that of native species through reinforcing feedback processes.Methods Using both systems analysis and meta-analysis approaches, we synthesized changes to ecosystems caused by 173 invasive plant species. For the systems analysis, we examined published studies of impacts of invasive plants to determine which presented evidence consistent with a reinforcement of feedback processes. For the meta-analysis, we calculated the effect size ratio between standardized changes in recipient ecosystem and in the status of introduced species as an indication of a reinforcing feedback in particular speciesenvironment combinations. The systems analysis approach allowed us to conceptualize regime shifts in invader-dominated landscapes and to estimate the likelihood of such changes occurring. The meta-analysis allowed us to quantitatively verify the conceptual model and the key invader-context feedbacks and to detect the strength and direction of feedbacks.Results Most reinforcing feedbacks involve impacts on soil-nutrient cycling by shrub and tree invaders in forests and herbaceous invaders in wetlands. Feedbacks resulting in regime shifts were most likely related to processes associated with seed banks, fire and nutrient cycling. Results were used to derive a key for identifying high-impact invaders.Main conclusions Identifying combinations of plant life-forms and ecosystems most likely to result in regime shifts is a robust approach for predicting highimpact invasions and therefore for prioritizing management interventions. The meta-analysis revealed the need for more quantitative studies, including manipulative experiments, on ecosystem feedbacks.

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The search for traits associated with plant invasiveness has yielded contradictory results, in part because most previous studies have failed to recognize that different traits are important at different stages along the introduction–naturalization–invasion continuum. Here we show that across six different habitat types in temperate Central Europe, naturalized non-invasive species are functionally similar to native species occurring in the same habitat type, but invasive species are different as they occupy the edge of the plant functional trait space represented in each habitat. This pattern was driven mainly by the greater average height of invasive species. These results suggest that the primary determinant of successful establishment of alien species in resident plant communities is environmental filtering, which is expressed in similar trait distributions. However, to become invasive, established alien species need to be different enough to occupy novel niche space, i.e. the edge of trait space.

Adaptation to heterogeneous environments can occur via phenotypic plasticity, but how often this occurs is unknown. Reciprocal transplant studies provide a rich dataset to address this issue in plant populations because they allow for a determination of the prevalence of plastic versus canalized responses. From 31 reciprocal transplant studies, we quantified the frequency of five possible evolutionary patterns: (1) canalized response–no differentiation: no plasticity, the mean phenotypes of the populations are not different; (2) canalized response–population differentiation: no plasticity, the mean phenotypes of the populations are different; (3) perfect adaptive plasticity: plastic responses with similar reaction norms between populations; (4) adaptive plasticity: plastic responses with parallel, but not congruent reaction norms between populations; and (5) nonadaptive plasticity: plastic responses with differences in the slope of the reaction norms. The analysis included 362 records: 50.8% life-history traits, 43.6% morphological traits, and 5.5% physiological traits. Across all traits, 52% of the trait records were not plastic, and either showed no difference in means across sites (17%) or differed among sites (83%). Among the 48% of trait records that showed some sort of plasticity, 49.4% showed perfect adaptive plasticity, 19.5% adaptive plasticity, and 31% nonadaptive plasticity. These results suggest that canalized responses are more common than adaptive plasticity as an evolutionary response to environmental heterogeneity.