Interspecific hybridization between a native and an invading plant species, or two invading species, sometimes results in a new, sexually reproducing taxon. Several examples of such taxa have been confirmed by recent molecular and isozyme analyses. Further study of these new taxa, when recognized soon after their origin, should aim to elucidate the factors that influence their subsequent establishment and spread, thus leading to a better understanding of the processes that lead to successful speciation. Plant hybrids formed following a plant invasion provide great potential for the study of 'evolution in action'.

Diversity in the tRNALEU1 intron of the chloroplast genome of Spartina was used to study hybridization of native California cordgrass, Spartina foliosa, with S. alterniflora, introduced to San Francisco Bay approximately 25 years ago. We sequenced 544 bases of the tRNALEU1 intron and found three polymorphic sites, a pyrimidine transition at site 126 and transversions at sites 382 and 430. Spartina from outside of San Francisco Bay, where hybridization between these species is impossible, gave cpDNA genotypes of the parental species. S. foliosa had a single chloroplast haplotype, CCT, and this was unique to California cordgrass. S. alterniflora from the native range along the Atlantic coast of North America had three chloroplast haplotypes, CAT, TAA, and TAT. Hybrids were discriminated by random amplified polymorphic DNA (RAPD) phenotypes developed in a previous study. We found one hybrid that contained a cpDNA haplotype unknown in either parental species (TCT). The most significant finding was that hybridization proceeds in both directions, assuming maternal inheritance of cpDNA; 26 of the 36 hybrid Spartina plants from San Francisco Bay contained the S. foliosa haplotype, nine contained haplotypes of the invading S. alterniflora, and one had the cpDNA of unknown origin. Furthermore, cpDNA of both parental species was distributed throughout the broad range of RAPD phenotypes, suggesting ongoing contributions to the hybrid swarm from both. The preponderance of S. foliosa cpDNA has entered the hybrid swarm indirectly, we propose, from F1s that backcross to S. foliosa. Flowering of the native precedes by several weeks that of the invading species, with little overlap between the two. Thus, F1 hybrids would be rare and sired by the last S. foliosa pollen upon the first S. alterniflora stigmas. The native species produces little pollen and this has low viability. An intermediate flowering time of hybrids as well as pollen that is more vigourous and abundant than that of the native species would predispose F1s to high fitness in a vast sea of native ovules. Thus, spread of hybrids to other S. foliosa marshes could be an even greater threat to the native species than introductions of alien S. alterniflora.

Populations of exotic earthworms responded positively to the presence of the nitrogen-fixing tree, Myrica faya, which is currently invading early succesional habitats in Hawaii Volcanoes National Park. Earthworm biomass in one high-density stand of Myrica was over three times the levels in nearby submontane forest and rainforest. Comparisons of earthworm populations under pairs of Myrica and the dominant native tree, Metrosideros polymorpha, showed biomass levels to be elevated from over two- to almost eightfold under the exotic tree. The increased rate of burial of nitrogenrich litter by earthworms can alter the rate of nitrogen accretion and cycling in these ecosystems.

The relationship between plant invasions and global change is complex. Whereas some components of global change, such as rising CO2, usually promote invasion, other components, such as changing temperature and precipitation, can help or hinder plant invasion. Additionally, experimental studies and models suggest that invasive plants often respond unpredictably to multiple components of global change acting in concert. Such variability adds uncertainty to existing risk assessments and other predictive tools. Here, we review current knowledge about relationships between plant invasion and global change, and highlight research needed to improve forecasts of invasion risk. Managers should be prepared for both expansion and contraction of invasive plants due to global change, leading to increased risk or unprecedented opportunities for restoration.

Invasive plants are an economic problem and a threat to the conservation of natural systems. Escape from natural enemies might contribute to successful invasion, with most work emphasizing the role of insect herbivores; however, microbial pathogens are attracting increased attention. Soil biota in some invaded ecosystems may promote 'exotic' invasion, and plant-soil feedback processes are also important. Thus, relatively rare species native to North America consistently demonstrate negative feedbacks with soil microbes that promote biological diversity, whereas abundant exotic and native species demonstrate positive feedbacks that reduce biological diversity. Here we report that soil microbes from the home range of the invasive exotic plant Centaurea maculosa L. have stronger inhibitory effects on its growth than soil microbes from where the weed has invaded in North America. Centaurea and soil microbes participate in different plant-soil feedback processes at home compared with outside Centaurea's home range. In native European soils, Centaurea cultivates soil biota with increasingly negative effects on the weed's growth, possibly leading to its control. But in soils from North America, Centaurea cultivates soil biota with increasingly positive effects on itself, which may contribute to the success of this exotic species in North America.

As increasing attention has been paid to below-ground ecosystems in recent years, the impact of exotic plant invasions on soil ecosystems has emerged as a central issue in invasion ecology. In this paper, effects of exotic plant invasions on soil biota and soil nutrient cycling processes are reviewed, and the mechanisms that underlie these impacts are discussed. The available literature suggests that no consistent pattern exists on the impacts of exotic plant invasions on soil microbes, soil animals, soil carbon cycling and nitrogen cycling. The mechanisms underlying the impacts of exotic plant invasions are also complex and diverse. The discrepancies most likely arise from the differences between invasive and native plants in a variety of physiological and ecological traits, such as litter quality and quantity, root distribution, and phenology. Future studies are recommended to (a) compare the impacts across multiple scales and multiple ecosystems; (b) intensify mechanical studies of the impacts; (c) link the impacts on biodiversity and ecosystem processes; and (d) explore the mutualistic interactions between soil ecosystems and exotic plants.

Various biotic and abiotic components of soil ecology differed significantly across an area where Halogeton glomeratus is invading a native winterfat, [ Krascheninnikovia (= Ceratoides) lanata] community. Nutrient levels were significantly different among the native, ecotone, and exotic-derived soils. NO3, P, K, and Na all increased as the cover of halogeton increased. Only Ca was highest in the winterfat area. A principal components analysis, conducted separately for water-soluble and exchangeable cations, revealed clear separation between halogeton- and winterfat-derived soils. The diversity of soil bacteria was highest in the exotic, intermediate in the ecotone, and lowest in the native community. Although further studies are necessary, our results offer evidence that invasion by halogeton alters soil chemistry and soil ecology, possibly creating conditions that favor halogeton over native plants.]]>

Plants can change soil biology, chemistry and structure in ways that alter subsequent plant growth. This process, referred to as plant-soil feedback (PSF), has been suggested to provide mechanisms for plant diversity, succession and invasion. Here we use three meta-analytical models: a mixed model and two Bayes models, one correcting for sampling dependence and one correcting for sampling and hierarchical dependence (delta-splitting model) to test these hypotheses. All three models showed that PSFs have medium to large negative effects on plant growth, and especially grass growth, the life form for which we had the most data. This supports the hypothesis that PSFs, through negative frequency dependence, maintain plant diversity, especially in grasslands. PSFs were also large and negative for annuals and natives, but the delta-splitting model indicated that more studies are needed for these results to be conclusive. Our results support the hypotheses that PSFs encourage successional replacements and plant invasions. Most studies were performed using monocultures of grassland species in greenhouse conditions. Future research should examine PSFs in plant communities, non-grassland systems and field conditions.

We investigated the influence of female age on five reproductive traits and on the offspring size-number trade-off from an extensive data set spanning 20 years of study on free-ranging Alpine marmots. Offspring mass increased with female age, whereas litter size and reproductive allocation remained constant in females up to 10 years of age and declined thereafter. Although reproductive allocation declined, post-weaning juvenile survival and the size-number trade-off did not change markedly throughout a female's lifetime. Senescence of annual reproductive success (i.e., the number of offspring surviving their first hibernation within a given litter) only resulted from senescence of litter size. The data were insufficient to determine whether the decrease in litter size with age was caused by declining litter size at birth, offspring pre-weaning survival, or both. Regardless, our findings demonstrate that marmot females display a size-number trade-off invariant with age, and that their reproductive tactic involves increasing offspring size at the cost of decreasing litter size with increasing age. As a result, reproductive performance remains constant throughout a female's lifetime, despite the deleterious effects of senescence in litter size.

Both ecological and evolutionary processes can influence community assembly and stability, and native community members may respond both ecologically and evolutionarily as additional species enter established communities. Biological invasions provide a unique opportunity to examine these responses of native community members to novel species additions. Here, I use reciprocal transplant experiments among naturally invaded and uninvaded environments, along with experimental removals of exotic species, to determine whether exotic plant competitors and exotic insect herbivores evoke evolutionary changes in native plants. Specifically, I address whether the common native plant species Lotus wrangelianus has responded evolutionarily to a series of biological invasions by adapting to the presence of the exotic plant Medicago polymorpha and the exotic insect herbivore Hypera brunneipennis. Despite differences in selection regimes between invaded and uninvaded environments and the presence of genetic variation for traits relevant to the novel competitive and plant-herbivore interactions, these experiments failed to reveal evidence that Lotus has responded evolutionarily to the double invasion of Medicago followed by H. brunneipennis. However, when herbivory from H. brunneipennis was experimentally reduced, Lotus plants from source populations invaded by Medicago outperformed plants from uninvaded source populations when transplanted into heavily invaded destination environments. Therefore, Lotus showed evidence of adaptation to Medicago invasion but not to the newer invasion of an exotic shared herbivore. The presence of this exotic insect herbivore alters the outcome of evolutionary responses in this system and counteracts adaptation by the native Lotus to invasion by the exotic plant Medicago. This result has broad implications for the conservation of native communities. While native species may be able to adapt to the presence of one or a few exotics, a multitude of invasions may limit the ability of natives to respond evolutionarily to the novel and frequently changing selection pressures that arise with subsequent invasions.

Although the impacts of exotic plant invasions on community structure and ecosystem processes are well appreciated, the pathways or mechanisms that underlie these impacts are poorly understood. Better exploration of these processes is essential to understanding why exotic plants impact only certain systems, and why only some invaders have large impacts. Here, we review over 150 studies to evaluate the mechanisms underlying the impacts of exotic plant invasions on plant and animal community structure, nutrient cycling, hydrology and fire regimes. We find that, while numerous studies have examined the impacts of invasions on plant diversity and composition, less than 5% test whether these effects arise through competition, allelopathy, alteration of ecosystem variables or other processes. Nonetheless, competition was often hypothesized, and nearly all studies competing native and alien plants against each other found strong competitive effects of exotic species. In contrast to studies of the impacts on plant community structure and higher trophic levels, research examining impacts on nitrogen cycling, hydrology and fire regimes is generally highly mechanistic, often motivated by specific invader traits. We encourage future studies that link impacts on community structure to ecosystem processes, and relate the controls over invasibility to the controls over impact.

Plant invaders have been suggested to change soil microbial communities and biogeochemical cycling in ways that can feedback to benefit themselves. In this paper, we ask when do these feedbacks influence the spread of exotic plants. Because answering this question is empirically challenging, we show how ecological theory on 'pushed' and 'pulled' invasions can be used to examine the problem. We incorporate soil feedbacks into annual plant invasion models, derive the conditions under which such feedbacks affect spread, and support our approach with simulations. We show that in homogeneous landscapes, strong positive feedbacks can influence spreading velocity for annual invaders, but that empirically documented feedbacks are not strong enough to do so. Moreover, to influence spread, invaders must modify the soil environment over a spatial scale larger than is biologically realistic. Though unimportant for annual invader spread in our models, feedbacks do affect invader density and potential impact. We discuss how future research might consider the way landscape structure, dispersal patterns, and the time scales over which plant-soil feedbacks develop regulate the effects of such feedbacks on invader spread.

Plant invasions are unintended consequences of globalisation , which facilitates the transglobal movement of plant species across all geographical and physical boundaries with the vastly increasingmovement of people and commercial goods. As an internationalised city that has various channels through which plant s′movements occur , Shanghai has been subject to a heavy invasion of plants. Thispaper firstly deals with some characteristics of plant invasions in Shanghai. It is concluded that : 1) alien plants play an important role in the flora of Shanghai , accounting for 57. 4 %of the total flora ; 2) annual and biennial plant s are the major component s of Shanghai′s alien flora , representing 69.1 %; 3) most alien plants belong to a relatively few families (e. g. Asteraceae and Poaceae) ; 4) monocotylous families tend to have a higher proportion of alien plants with respect to their world′s number of species than do dicotylous families ; and 5) plant invasions will continue to sweep across literally every part of Shanghai as the consequence of it s further urbanisation and increasing global trade (c. f . China′s ent ry into WTO by November 2001) . These patterns of plant invasions reflect the role of humans as global plant dispersers and of human disturbance in plant invasions. In the rest of the paper , we briefly review other issues in the field of plant invasions , including att ributes of invasive species , habitat invisibility , environmental and economic costs of plant invasions , and future research directions.

Invasions of exotic plant species are among the most pervasive and important threats to natural ecosystems. However, the effects of plant invasions on soil processes and soil biota have not been adequately investigated. Changes were studied in soil microbial communities where Mikania micrantha was invading a native forest community in Neilingding Island, Shenzhen, China. The soil microbial community structure (assessed by phospholipid fatty acid [PLFA] profiles) and function (assessed by enzyme activities), as well as soil chemical properties were measured. The results showed that the invasion of M. micrantha into the evergreen broadleaved forests in South China changed most of the characteristics in studied soils. Microbial community structure and function differed significantly among the native, two ecotones, and exotic-derived soils. For PLFA profiles, we observed a significant increase in aerobic bacteria but a decrease in anaerobic bacteria in the M. micrantha monoculture as compared to the native and ecotones. The ratio of cy19:0 to18:1ω7 gradually declined but mono/sat PLFAs increased as M. micrantha became more dominant. Both ratios were significantly related to pH according to regression analysis, therefore, pH was a sensitive indicator reflecting the invaded soil subsystem succession. The microbial community composition clearly separated the native soil from the invaded soils by principal component analysis (PCA) and discriminant analysis (DA). For enzyme activities, 7 of 9 enzymes (β-glucosidase, invertase, protease, urease, acid phosphatase, alkaline phosphatase, and phenol oxidase) showed the similar trend that the activities were highest in the exotic, intermediate in the two ecotones, and lowest in the native community. In most cases, enzyme activities were influenced by soil chemical properties, especially by pH value and soil organic matter. Differences in the structural variables were well correlated to differences in the functional variables as demonstrated by canonical correlation analysis (CCA). It was concluded that M. micrantha invasion had profound effects on the soil subsystem, which must be taken into account when we try to control its invasions.]]>

Eupatorium adenophorum is one of the most noxious invasive weeds in China. To explore the effects of this weed invasion on soil fertility, the activities of six enzymes and 12 physical and chemical factors were compared among the soil within 0~30 cm depth under a more than 10 years old invaded grassland by E. adenophorum, a secondary grassland (abandoned maize field in November of 2001), a primary grassland of Themeda yunnanensis and a two years old artificial grassland of Setaria sphacelata. The results showed that each variable measured in this study changed significantly among communities and soil layers according to a two-way ANOVA. The activities of polyphenol oxidase, alkaline phosphatase and urase, the contents of organic matter, total N, total P, total Ca, hydrolytic N, active P, active K, and pH value decreased with the increase of soil depth. In general, the activities of alkaline phosphatase and urase, the contents of organic matter, total N, total P, total Ca, hydrolytic N, active P, and pH value were higher in the soil under E. adenophorum community compared to the soils under other communities. Total soil K content was lower in the soil under E. adenophorum than in the soils of other communities, while the active K content was similar among the four communities. These results indicate that E. adenophorum invasion for more than 10 years can increase soil fertility, create favorable soil conditions for itself.]]>

How introduced plants, which may be locally adapted to specific climatic conditions in their native range, cope with the new abiotic conditions that they encounter as exotics is not well understood. In particular, it is unclear what role plasticity versus adaptive evolution plays in enabling exotics to persist under new environmental circumstances in the introduced range. We determined the extent to which native and introduced populations of St. John's Wort (Hypericum perforatum) are genetically differentiated with respect to leaf-level morphological and physiological traits that allow plants to tolerate different climatic conditions. In common gardens in Washington and Spain, and in a greenhouse, we examined clinal variation in percent leaf nitrogen and carbon, leaf delta(13)C values (as an integrative measure of water use efficiency), specific leaf area (SLA), root and shoot biomass, root/shoot ratio, total leaf area, and leaf area ratio (LAR). As well, we determined whether native European H. perforatum experienced directional selection on leaf-level traits in the introduced range and we compared, across gardens, levels of plasticity in these traits. In field gardens in both Washington and Spain, native populations formed latitudinal clines in percent leaf N. In the greenhouse, native populations formed latitudinal clines in root and shoot biomass and total leaf area, and in the Washington garden only, native populations also exhibited latitudinal clines in percent leaf C and leaf delta(13)C. Traits that failed to show consistent latitudinal clines instead exhibited significant phenotypic plasticity. Introduced St. John's Wort populations also formed significant or marginally significant latitudinal clines in percent leaf N in Washington and Spain, percent leaf C in Washington, and in root biomass and total leaf area in the greenhouse. In the Washington common garden, there was strong directional selection among European populations for higher percent leaf N and leaf delta(13)C, but no selection on any other measured trait. The presence of convergent, genetically based latitudinal clines between native and introduced H. perforatum, together with previously published molecular data, suggest that native and exotic genotypes have independently adapted to a broad-scale variation in climate that varies with latitude.

Understanding the evolution and demography of invasive populations may be key for successful management. In this study, we test whether or not populations of the non-native, hybrid-derived California wild radish have regionally adapted to divergent climates over their 150-year history in California and determine if population demographic dynamics might warrant different region-specific strategies for control. Using a reciprocal transplant approach, we found evidence for genetically based differences both between and among northern, coastal and southern, inland populations of wild radish. Individual fitness was analyzed using a relatively new statistical method called 'aster modeling' which integrates temporally sequential fitness measurements. In their respective home environments, fitness differences strongly favored southern populations and only slightly favored northern populations. Demographic rates of transition and sensitivities also differed between regions of origin, suggesting that the most effective approach for reducing overall population growth rate would be to target different life-history stages in each region.