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Invasions are commonly found to benefit from disturbance events. However, the importance of the relative timing of the invasion and disturbance for invader success and impact on community composition remains uncertain. Here, we experimentally test this by invading a five-species bacterial community on eight separate occasions-four before a disturbance and four after. Invader success and impact on community composition was greatest when the invasion immediately followed the disturbance. However, the subsequent invasions had negligible success or impact. Pre-disturbance, invader success and impact was greatest when the invader was added just before the disturbance. Importantly, however, the first three pre-disturbance invasion events had significantly greater success than the last three post-disturbance invasions. Moreover, these findings were consistent across a range of propagule pressures. Overall, we demonstrate that timing is highly important for both the success and impact on community composition of an invader, with both being lower as time since disturbance progresses.

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Behavioural traits are key to promote invasion success because they are easier to adjust to changing environmental conditions than morphological or life history traits. Often, research has overlooked variance in behavioural traits within populations or has assumed it to be mere noise. However, a recent focus towards individual variation of behaviour of successful invaders has revealed new and more profound insights into the invasion process. Behavioural variation within a population could lead to more successful invasions, as they include individuals with diverse behaviours, which ensures at least some individuals could be able to cope with changing conditions. The aim of this research was to examine if invasive guppies (Poecilia reticulata) present within-population differences in their sociability (time spent associating with a shoal) when interacting with conspecifics or heterospecifics. Guppies presented significant differences in their individual tendencies to associate with conspecific or heterospecific shoals. There were among-individual differences in the time spent shoaling with conspecifics versus heterospecifics, where most individuals did not differ in their sociability with conspecifics or heterospecifics, and only 22% of individuals presented a higher tendency to associate with conspecifics. Our results are the first to show individual differences in fish' tendencies to associate with heterospecifics among individuals of the same population and rearing conditions. Given that associations with heterospecific natives have been found to be as beneficial as associations with conspecifics for invaders, our results contribute to the understanding of mechanisms behind heterospecific sociability between natives and invaders.

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The conservation-invasion paradox (CIP) refers to a long-term phenomenon wherein species threatened in their native range can sustain viable populations when introduced to other regions. Understanding the drivers of CIP is helpful for conserving threatened species and managing invasive species, which is unfortunately still lacking. We compiled a global data set of 1071 introduction events, including 960 CIP events (successful establishment of threatened species outside its native range) and 111 non-CIP events (unsuccessful establishment of threatened species outside its native range after introduction), involving 174 terrestrial vertebrates. We then tested the relative importance of various predictors at the location, event, and species levels with generalized linear mixed models and model averaging. Successful CIP events occurred across taxonomic groups and biogeographic realms, especially for the mammal group in the Palearctic and Australia. Locations of successful CIP events had fewer native threat factors, especially less climate warming in invaded regions. The probability of a successful CIP event was highest when species introduction efforts were great and there were more local congeners and fewer natural enemies. These results can inform threatened species ex situ conservation and non-native invasive species mitigation.

Causantes mundiales de la paradoja conservación­invasión Resumen La paradoja de conservación­invasión (PCI) se refiere al evento a largo plazo en el que las especies amenazadas en su distribución nativa puedan mantener poblaciones viables cuando se les introduce a otras regiones. Es de mucha ayuda para la conservación de especies amenazadas y el manejo de especies invasoras entender las causantes de la PCI, entendimiento que todavía es escaso. Compilamos un conjunto mundial de datos de 174 vertebrados terrestres en 1071 eventos de introducción, incluyendo 960 eventos de PCI (el establecimiento exitoso de especies amenazadas fuera de su distribución nativa) y 111 eventos no PCI (el fracaso en el establecimiento de especies amenazadas fuera de su distribución nativa después de la introducción). Después analizamos con modelos lineales mixtos generalizados y promedio de modelos la importancia relativa de varios pronosticadores en la localidad, en el evento y a nivel de especie. Los eventos exitosos de PCI ocurrieron en todos los grupos taxonómicos y en todos los reinos biogeográficos, especialmente para los mamíferos del Paleártico y Australia. Las localidades de los eventos exitosos de PCI tuvieron menos factores nativos de amenaza, especialmente un menor calentamiento climático en las regiones invadidas. La probabilidad de que un evento de PCI sea exitoso fue mayor cuando los esfuerzos de introducción fueron mayores y hubo más congéneres locales y menos enemigos naturales. Estos resultados pueden orientar la conservación ex situ de especies y la mitigación de especies invasoras no nativas.

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Anthropogenic activities expose many ecosystems to multiple novel disturbances simultaneously. Despite this, how biodiversity responds to simultaneous disturbances remains unclear, with conflicting empirical results on their interactive effects. Here, we experimentally test how one disturbance (an invasive species) affects the diversity of a community over multiple levels of another disturbance regime (pulse mortality). Specifically, we invade stably coexisting bacterial communities under four different pulse frequencies, and compare their final resident diversity to uninvaded communities under the same pulse mortality regimes. Our experiment shows that the disturbances synergistically interact, such that the invader significantly reduces resident diversity at high pulse frequency, but not at low. This work therefore highlights the need to study simultaneous disturbance effects over multiple disturbance regimes as well as to carefully document unmanipulated disturbances, and may help explain the conflicting results seen in previous multiple-disturbance work.

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Invasive species and chemical pollution both threaten biodiversity. Here, we discuss how pollution, through its impacts on wildlife behaviour, shapes invasion dynamics by altering species interactions. Addressing knowledge gaps will have implications for the management of invasive species and conservation of native ecosystems in an increasingly toxic world.

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Ecosystem functions are heavily dependent on the functional composition of the plant community, i.e., the functional traits of plants forming the community. This, on the one hand, depends on plant occurrence, but on the other hand, depends on the intraspecific variability of functional traits of the species, which are influenced by climate and nutrient availability and affected by plant-plant interactions. To illustrate that, we studied the effects of drought and nitrogen addition (+ N), two important abiotic variables which are changing with ongoing global change, as well as their combined effect on the functional responses of grassland communities in semi-arid environments of Northern Africa comprising of natural and invasive species. We conducted an experiment where we planted three native species and one invasive plant species in artificial communities of five individuals per species per plot. We exposed these communities to four different treatments: a drought treatment, an N-addition treatment, the combination between drought and N-addition, as well as a control. To assess the performance of plants within treatments, we measured selected plant functional traits (plant height, specific leaf area [SLA], leaf dry matter content [LDMC], N content of the leaves [Nmass], specific root length [SRL], and root diameter) for all individuals occurring in our plots, and additionally assessed the above and belowground biomass for each plant individual. We found that the invasive species showed a higher performance (higher biomass accumulation, taller plants, higher SLA, Nmass, SRL, and root diameter as well as lower LDMC) than the native species under drought conditions. The invasive species was especially successful with the combined impact of drought + N, which is a likely scenario in ongoing global change for our research area. Thus, plant functional traits might be a key factor for the invasion success of plant species which will be even more pronounced under ongoing global change.

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PREMISE: Invasive plant species often escape from specialist herbivores and are more likely to be attacked by generalist herbivores in the exotic range. Consequently, the shifting defense hypothesis predicts that invasive plants will produce higher concentrations of qualitative defense compounds to deter dominant generalist herbivores in the exotic range. Here, I additionally propose a reduced chemical diversity hypothesis (RCDH), which predicts that reduced herbivory pressure will select for invasive plant genotypes that produce lower diversities of chemical defense compounds in the exotic range. METHODS: I tested whether (1) invasive Brassica nigra populations express a lower diversity and an overall higher concentration of glucosinolate compounds than native-range B. nigra; (2) Brassica nigra individuals that express high diversities and concentrations of glucosinolate compounds are more attractive to specialist and deterrent to generalist herbivores; and (3) tissues of invasive B. nigra are less palatable than tissues of native-range B. nigra to the generalist herbivores Theba pisana and Helix aspersa. RESULTS: Invasive B. nigra populations produced a significantly lower diversity of glucosinolate compounds, a marginally higher concentration of total glucosinolates, and a significantly higher concentration of sinigrin (the dominant glucosinolate). Leaf tissues of invasive B. nigra were significantly less palatable to T. pisana and marginally less so to H. aspersa. Brassica nigra individuals that expressed high concentrations of total glucosinolate compounds were visited by a low diversity of generalist herbivore species in the field. CONCLUSIONS: In line with the RCDH, the lower diversity of glucosinolate compounds produced by invasive B. nigra populations likely resulted from selection imposed by reduced herbivory pressure in the exotic range.

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PREMISE: The enemy release hypothesis predicts that release from natural enemies, including soil-borne pathogens, liberates invasive plants from a negative regulating force. Nevertheless, invasive plants may acquire novel enemies and mutualists in the introduced range, which may cause variable effects on invader growth. However, how soil microorganisms may influence competitive ability of invasive plants along invasion chronosequences has been little explored. METHODS: Using the invasive plant Solidago canadensis, we tested whether longer residence times are associated with stronger negative plant-soil feedbacks and thus weaker competitive abilities at the individual level. We grew S. canadensis individuals from 36 populations with different residence times across southeastern China in competition versus no competition and in three different types of soils: (1) conspecific rhizospheric soils; (2) soils from uninvaded patches; and (3) sterilized soil. For our competitor treatments, we constructed synthetic communities of four native species (Bidens parviflora, Solanum nigrum, Kalimeris indica, and Mosla scabra), which naturally co-occur with Solidago canadensis in the field. RESULTS: Solidago canadensis populations with longer residence times experienced stronger positive plant-soil feedbacks and had greater competitive responses (i.e., produced greater above-ground biomass and grew taller) in conspecific rhizospheric soils than in sterilized or uninvaded soils. Moreover, S. canadensis from older populations significantly suppressed above-ground biomass of the native communities in rhizospheric and uninvaded soils but not in sterilized soil. CONCLUSIONS: The present results suggest that older populations of S. canadensis experience stronger positive plant-soil feedbacks, which may enhance their competitive ability against native plant communities.

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Invasive species Symphyotrichum lanceolatum (Willd.) G. L. Nesom is spreading uncontrollably along wet habitats as well as in disturbed ecosystems. All those habitats function as corridors that facilitate seed dispersal. One way to prevent the spread of invasive species is to know their reproductive ecology. The present study evaluates the potential for generative reproduction of S. lanceolatum and determines how different temperatures, amounts of nutrients, and light regimes, affect seed germination. Seeds collected from 13 natural populations were germinated at four fluctuating temperature regimes (15/6, 20/10, 30/15, and 35/20 °C). To test the influence of nitrate on seed germination, two KNO3 concentrations were used (5 mM and 50 mM solution). For each treatment, three replicates of 30 seeds were placed in complete darkness or a 14 h photoperiod. The results showed that the germination increased with increasing temperature. The optimal temperature regimes were 30/15 °C and 35/20 °C with approximately 88% germination. The overall effect of KNO3 on germination was positive. The concentration of 50 mM KNO3 had a less stimulating effect compared to 5 mM KNO3. Seeds showed sensitivity to lack of light during germination but were able to germinate in a significant percentage in such conditions. Considering that S. lanceolatum often occurs in disturbed sites, these results suggest that seed reaction to alternating temperature, nutrients concentration, and light can be determining factors that affect seed germination of this species and, thus, its spread.

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Disturbances can facilitate biological invasions, with the associated increase in resource availability being a proposed cause. Here, we experimentally tested the interactive effects of disturbance regime (different frequencies of biomass removal at equal intensities) and resource abundance on invasion success using a factorial design containing five disturbance frequencies and three resource levels. We invaded populations of the bacterium Pseudomonas fluorescens with two ecologically different invader morphotypes: a fast-growing "colonizer" type and a slower growing "competitor" type. As resident populations were altered by the treatments, we additionally tested their effect on invader success. Disturbance frequency and resource abundance interacted to affect the success of both invaders, but this interaction differed between the invader types. The success of the colonizer type was positively affected by disturbance under high resources but negatively under low. However, disturbance negatively affected the success of the competitor type under high resource abundance but not under low or medium. Resident population changes did not alter invader success beyond direct treatment effects. We therefore demonstrate that the same disturbance regime can either be beneficial or detrimental for an invader depending on both community resource abundance and its life history. These results may help to explain some of the inconsistencies found in the disturbance-invasion literature.

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Successful control and prevention of biological invasions depend on identifying traits of non-native species that promote fitness advantages in competition with native species. Here, we show that, among 76 native and non-native woody plants of deciduous forests of North America, invaders express a unique functional syndrome that combines high metabolic rate with robust leaves of longer lifespan and a greater duration of annual carbon gain, behaviours enabled by seasonally plastic xylem structure and rapid production of thin roots. This trait combination was absent in all native species examined and suggests the success of forest invaders is driven by a novel resource-use strategy. Furthermore, two traits alone-annual leaf duration and nuclear DNA content-separated native and invasive species with 93% accuracy, supporting the use of functional traits in invader risk assessments. A trait syndrome reflecting both fast growth capacity and understorey persistence may be a key driver of forest invasions.

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Understanding drivers of success for alien species can inform on potential future invasions. Recent conceptual advances highlight that species may achieve invasiveness via performance along at least three distinct dimensions: 1) local abundance, 2) geographic range size, and 3) habitat breadth in naturalized distributions. Associations among these dimensions and the factors that determine success in each have yet to be assessed at large geographic scales. Here, we combine data from over one million vegetation plots covering the extent of Europe and its habitat diversity with databases on species' distributions, traits, and historical origins to provide a comprehensive assessment of invasiveness dimensions for the European alien seed plant flora. Invasiveness dimensions are linked in alien distributions, leading to a continuum from overall poor invaders to super invaders-abundant, widespread aliens that invade diverse habitats. This pattern echoes relationships among analogous dimensions measured for native European species. Success along invasiveness dimensions was associated with details of alien species' introduction histories: earlier introduction dates were positively associated with all three dimensions, and consistent with theory-based expectations, species originating from other continents, particularly acquisitive growth strategists, were among the most successful invaders in Europe. Despite general correlations among invasiveness dimensions, we identified habitats and traits associated with atypical patterns of success in only one or two dimensions-for example, the role of disturbed habitats in facilitating widespread specialists. We conclude that considering invasiveness within a multidimensional framework can provide insights into invasion processes while also informing general understanding of the dynamics of species distributions.

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BACKGROUND AND AIMS: Non-native plant species are not restricted to lowlands, but increasingly are invading high elevations. While for both native and non-native species we expected variability of plant functional traits due to the changing environmental conditions along elevational gradients, we additionally assumed that non-native species are characterized by a more acquisitive growth strategy, as traits reflecting such a strategy have been found to correlate with invasion success. Furthermore, the typical lowland introduction of non-native species coming from multiple origins should lead to higher trait variability within populations of non-native species specifically at low elevations, and they might therefore occupy a larger total trait space. METHODS: Along an elevational gradient ranging from 55 to 1925 m a.s.l. on Tenerife, we collected leaves from eight replicate individuals in eight evenly distributed populations of five native and six non-native forb species. In each population, we measured ten eco-morphological and leaf biochemical traits and calculated trait variability within each population and the total trait space occupied by native and non-native species. KEY RESULTS: We found both positive (e.g. leaf dry matter content) and negative (e.g. leaf N) correlations with elevation for native species, but only few responses for non-native species. For non-native species, within-population variability of leaf dry matter content and specific leaf area decreased with elevation, but increased for native species. The total trait space occupied by all non-native species was smaller than and a subset of that of native species. CONCLUSIONS: We found little evidence that intraspecific trait variability is associated with the success of non-native species to spread towards higher elevations. Instead, for non-native species, our results indicate that intermediate trait values that meet the requirements of various conditions are favourable across the changing environmental conditions along elevational gradients. As a consequence, this might prevent non-native species from overcoming abruptly changing environmental conditions, such as when crossing the treeline.

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Anthropogenic changes are often studied in isolation but may interact to affect biodiversity. For example, climate change could exacerbate the impacts of biological invasions if climate change differentially affects invasive and native species. Behavioural plasticity may mitigate some of the impacts of climate change, but species vary in their degree of behavioural plasticity. In particular, invasive species may have greater behavioural plasticity than native species since plasticity helps invasive species establish and spread in new environments. This plasticity could make invasives better able to cope with climate change. Here our goal was to examine whether reproductive behaviours and behavioural plasticity vary between an introduced and a native Onthophagus dung beetle species in response to warming temperatures and how differences in behaviour influence offspring survival. Using a repeated measures design, we exposed small colonies of introduced O. taurus and native O. hecate to three temperature treatments, including a control, low warming and high warming treatment, and then measured reproductive behaviours, including the number, size and burial depth of brood balls. We reared offspring in their brood balls in developmental temperatures that matched those of the brood ball burial depth to quantify survival. We found that the introduced O. taurus produced more brood balls and larger brood balls, and buried brood balls deeper than the native O. hecate in all treatments. However, the two species did not vary in the degree of behavioural plasticity in response to warming. Differences in reproductive behaviours did affect survival such that warming temperatures had a greater effect on survival of offspring of native O. hecate compared to introduced O. taurus. Overall, our results suggest that differences in behaviour between native and introduced species are one mechanism through which climate change may exacerbate negative impacts of biological invasions.

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Human introductions of species beyond their natural ranges and their subsequent establishment are defining features of global environmental change. However, naturalized plants are not uniformly distributed across phylogenetic lineages, with some families contributing disproportionately more to the global alien species pool than others. Additionally, lineages differ in diversification rates, and high diversification rates have been associated with characteristics that increase species naturalization success. Here, we investigate the role of diversification rates in explaining the naturalization success of angiosperm plant families. We use five global data sets that include native and alien plant species distribution, horticultural use of plants, and a time-calibrated angiosperm phylogeny. Using phylogenetic generalized linear mixed models, we analysed the effect of diversification rate, different geographical range measures, and horticultural use on the naturalization success of plant families. We show that a family's naturalization success is positively associated with its evolutionary history, native range size, and economic use. Investigating interactive effects of these predictors shows that native range size and geographic distribution additionally affect naturalization success. High diversification rates and large ranges increase naturalization success, especially of temperate families. We suggest this may result from lower ecological specialization in temperate families with large ranges, compared with tropical families with smaller ranges.

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Invasive alien species are driving global biodiversity loss, compromising ecosystem function and service provision, and human, animal and plant health. Habitat characteristics and geographical origin may predict invasion success, and in aquatic environments could be mediated principally by salinity tolerance. Crustacean invaders are causing global problems and we urgently require better predictive power of their invasiveness. Here, we compiled global aquatic gammarid (Crustacea: Amphipoda: Gammaroidea) diversity and examined their salinity tolerances and regions of origin to test whether these factors predict invasion success. Across 918 aquatic species within this superfamily, relatively few gammarids (n = 27, 3%) were reported as aliens, despite extensive invasion opportunities and high numbers of published studies on amphipod invasions. However, reported alien species were disproportionately salt-tolerant (i.e. 32% of brackish-water species), with significantly lower proportions of aliens originating from freshwater and marine environments (both 1%). Alien gammarids also significantly disproportionally originated from the Ponto-Caspian (20% of these taxa) when compared with all 'other' grouped regions (1%), and principally invaded Eurasian waters, with translocations of salt-tolerant taxa to freshwaters being pervasive. This suggests habitat characteristics, alongside regional contexts, help predict invasibility. In particular, broad environmental tolerances to harsh environments and associated evolutionary history probably promote success of aliens globally.

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As a successful invasive cyanobacterial species, Raphidiopsis raciborskii is distributed globally and shows a tendency to replace Microcystis aeruginosa in many subtropical and temperate waters, but the ecological traits that contribute to its invasiveness are still unclear. In this study, we found that R. raciborskii occurred in 149 sites in 42 lakes in eastern and central China and coexisted with M. aeruginosa at most sites. Based on field results, a combination of invasion and competition experiments using a biomass gradient to evaluate the invasiveness and competitiveness of R. raciborskii compared with M. aeruginosa was conducted. In invasive groups, both R. raciborskii and M. aeruginosa were shown to have positive specific growth rates, indicating that R. raciborskii could coexist with M. aeruginosa. Furthermore, R. raciborskii was shown to grow faster from invasion while M. aeruginosa reduced growth for invasion. In competitive groups, R. raciborskii reached a higher maximum biomass and grew longer than M. aeruginosa. The specific growth rate of R. raciborskii was not inhibited by M. aeruginosa biomass, whereas the growth of M. aeruginosa was inhibited by R. raciborskii biomass. It was shown during the whole experiment that R. raciborskii tended to replace M. aeruginosa to become dominant owing to its faster growth rate and the eventual decline in growth of M. aeruginosa. With an increase in biomass of M. aeruginosa, the vegetative cell size and filament length of R. raciborskii gradually increased. This study has demonstrated that the inherent invasive traits of R. raciborskii, size differences, niche differences, and relative fitness differences between R. raciborskii and M. aeruginosa are crucial reasons for the invasive success of R. raciborskii. Our results revealed the invasiveness and domination of R. raciborskii from a new perspective.

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Knowing the causes of biological invasion success can be relevant to combat future invasive processes. The recent invasion of the horseshoe whip snake Hemorrhois hippocrepis on the island of Ibiza provides the opportunity to compare natural history traits between invasive and source populations, and to unravel what makes this snake a successful invader that is threatening the only endemic vertebrate of the island, Podarcis pityusensis. This study compares the basic reproductive traits of mainland native and invasive populations of the snake. Our results revealed that invasive populations were characterized by female maturity at a smaller size, extended reproductive period, and much lower reproduction frequency compared to the native population. In contrast, some major reproductive traits-the abdominal fat body cycle, clutch size, hatchling body size, and hatchling body condition, did not differ between the two populations. Some of these results must reflect the environmental differences in the recently invaded island with respect to the source area, and overall plasticity of reproductive traits. Plasticity is evolutionarily interesting, and may aid the successful growth of this species in their invasiveness of Mediterranean islands like Ibiza. The most significant finding is that this expression of phenotypic plasticity occurred rapidly in this invasive population, within a period of 14 years maximum. Our results on the reproduction ecology of the invasive population were not conclusive regarding the factors determining the invasiveness of the snake and pointed to alternative causes.

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Invasive species often cause serious economic and ecological damage. Despite decades of extensive impacts of invasives on bio-diversity and agroforestry, the mechanisms underlying the genetic adaptation and rapid evolution of invading populations remain poorly understood. The black locust gall midge, Obolodiplosis robiniae, a highly invasive species that originated in North America, spread widely throughout Asia and Europe in the past decade. Here, we used 11 microsatellite DNA markers to analyze the genetic variation of 22 O. robiniae populations in China (the introduced region) and two additional US populations (the native region). A relatively high level of genetic diversity was detected among the introduced populations, even though they exhibited lower diversity than the native US populations. Evidence for genetic differentiation among the introduced Chinese populations was also found based on the high Fst value compared to the relatively low among the native US populations. Phylogenetic trees, structure graphical output, and principal coordinate analysis plots suggested that the Chinese O. robiniae populations (separated by up to 2,540 km) cluster into two main groups independent of geographical distance. Genetic variation has been observed to increase rapidly during adaptation to a new environment, possibly contributing to population establishment and spread. Our results provide insights into the genetic mechanisms underlying successful invasion, and identify factors that have contributed to colonization by an economically important pest species in China. In addition, the findings improve our understanding of the role that genetic structure plays during invasion by O. robiniae.

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It is important to illuminate the effects of litter decomposition of invasive alien species on soil N-fixing bacterial communities (SoNiBa), especially under heavy metal pollution to better outline the mechanisms for invasion success of invasive alien species. This study attempts to identify the effects of litter decomposition of Solidago canadensis L. on SoNiBa under cadmium (Cd) pollution with different concentrations (i.e., low concentration, 7.5 mg/kg soil; high concentration, 15 mg/kg soil) via a polyethylene litterbags-experiment. Electrical conductivity and total N of soil were the most important environmental factors for determining the variations of SoNiBa composition. S. canadensis did not significantly affect the alpha diversity of SoNiBa but significantly affect the beta diversity of SoNiBa and SoNiBa composition. Thus, SoNiBa composition, rather than alpha diversity of SoNiBa, was the most important determinant of the invasion success of S. canadensis. Cd with 15 mg/kg soil did not address distinct effects on alpha diversity of SoNiBa, but Cd with 7.5 mg/kg soil noticeably raised the number of species and species richness of SoNiBa mainly due to the hormonal effects. The combined S. canadensis and Cd with 15 mg/kg soil obviously decreased cumulative mass losses and the rate of litter decomposition (k) of S. canadensis, but the combined S. canadensis and Cd with 7.5 mg/kg soil evidently accelerated cumulative mass losses and k of S. canadensis. Thus, Cd with 7.5 mg/kg soil can accelerate litter decomposition of S. canadensis, but Cd with 15 mg/kg soil can decline litter decomposition of S. canadensis.

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