Speciation and the evolution of dispersal along environmental gradients

We analyze the joint evolution of an ecological character and of dispersal distance in asexual and sexual populations inhabiting an environmental gradient. Several interesting phenomena resulting from the evolutionary interplay of these characters are revealed. First, asexual and sexual populations exhibit two analogous evolutionary regimes, in which either speciation in the ecological character occurs in conjunction with evolution of short-range dispersal, or dispersal distance remains high and speciation does not occur. Second, transitions between these two regimes qualitatively differ between asexual and sexual populations, with the former showing speciation with long-range dispersal and the latter showing no speciation with short-range dispersal. Third, a phenotypic gradient following the environmental gradient occurs only in the last case, i.e., for non-speciating sexual populations evolving towards short-range dispersal. Fourth, the transition between the evolutionary regimes of long-range dispersal with no speciation and short-range dispersal with speciation is typically abrupt, mediated by a positive feedback between incipient speciation and the evolution of short-range dispersal. Fifth, even though the model of sexual evolution analyzed here does not permit assortative mating preferences, speciation occurs for a surprisingly wide range of conditions. This illustrates that dispersal evolution is a powerful alternative to preference evolution in enabling spatially distributed sexual populations to respond to frequency-dependent disruptive selection.     

Distance-dependent performance of asexual progeny in Allium vineale (Liliaceae)

Local adaptation within and among populations may have an impact on processes ranging from speciation to the evolution of mixed breeding systems and dispersal strategies. It is also one potential factor that could favor the production of asexual over sexual propagules. This field experiment tested whether asexually produced bulbils of Allium vineale demonstrate local adaptation to the parental microsite at the scale of natural dispersal from the parent (5, 25, 50, 100, and 1000 cm). Both "home' and randomly chosen "away' genotypes were planted at each location to determine the relative performance of the "home' genotype. Overall, bulbil performance declined with distance from the parent. In particular, "home' bulbils outperformed "away' bulbils at a distance of 25 cm from the parent, indicating that local adaptation has occurred at the scale of natural dispersal in this species. The variance in propagule performance also increased at farther distances from the parent, indicating that the predictability of offspring performance decreases with distance. Fine-scale local adaptation within the range of seed dispersal in this population may be one factor favoring asexual reproduction in Allium vineale.     

Conditional dispersal, clines, and the evolution of dispersiveness

Conditional dispersal, in which an individual’s decision over whether to disperse is a response to environmental conditions, features prominently in studies of dispersal evolution. Using models of clines, I examine how one widely discussed cost of dispersal, namely, that dispersal impedes local adaptation, changes with conditional dispersal and what this implies for dispersal evolution. I examine the consequences for dispersal evolution of the responsiveness of dispersal to the environment, the accuracy of any proximal cues that individuals rely upon to assess habitat quality, and whether dispersal responds to fitness itself or only to some fitness components (juvenile survivorship). All of the conditional dispersal behaviors that I consider weaken the indirect cost of dispersal inhibiting local adaptation. However, if individuals rely on imprecise cues to assess habitat quality and base dispersal decisions on juvenile survivorship, then conditional dispersal can incur additional costs by exacerbating overcrowding. Conditional dispersal initially leads to steeper clines in traits under direct selection, but when dispersiveness can itself evolve, conditional dispersal allows sigmoidal clines to persist long after those obtained with unconditional movement would become stepped. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Population subdivision in Australian temperate marine invertebrates: Larval connections versus historical factors*

This paper reviews the use of genetic data, in combination with manipulative experimentation, to infer the mode of reproduction and the extent and directionality of dispersal for a range of Australian temperate marine invertebrates. Local populations of obligately sexually reproducing species have been inferred to be strongly interconnected by larval dispersal, over distances of thousands of kilometres. Their larvae may be subject to strong post-settlement selection, but this selection is independent of obvious geographic or intertidal gradients. Within local populations selection may therefore result in apparently chaotic genetic patchiness which is eliminated by the effects of sexual reproduction and the widespread dispersal and mixing of the colonizing larvae of each generation. In partial contrast, local populations of species which rely on asexual reproduction for the maintenance of populations show evidence of similar larval connections, but no recent settlement of their sexually generated larvae has been demonstrated. The apparent connectedness of these populations may reflect either historical events or a more episodic pattern of settlement by sexually generated larvae. Local populations of these species are more highly differentiated as a result of the continued asexual replication of a limited number of genotypes. In one of these species, reciprocal transplantation of the clones within and among populations has revealed that resident clones can be highly locally adapted (as reflected by much higher asexual fecundity), which implies that selection is an important determinant of the composition of local populations. Nevertheless, the failure to detect continuing sexual recruitment into these populations obscures the evolutionary significance of this finding.     

Reproductive strategy, clonal structure and genetic diversity in populations of the aquatic macrophyte Sparganium emersum in river systems

Many aquatic and riparian plant species are characterized by the ability to reproduce both sexually and asexually. Yet, little is known about how spatial variation in sexual and asexual reproduction affects the genotypic diversity within populations of aquatic and riparian plants. We used six polymorphic microsatellites to examine the genetic diversity within and differentiation among 17 populations (606 individuals) of Sparganium emersum, in two Dutch-German rivers. Our study revealed a striking difference between rivers in the mode of reproduction (sexual vs. asexual) within S. emersum populations. The mode of reproduction was strongly related to locally reigning hydrodynamic conditions. Sexually reproducing populations exhibited a greater number of multilocus genotypes compared to asexual populations. The regional population structure suggested higher levels of gene flow among sexually reproducing populations compared to clonal populations. Gene flow was mainly mediated via hydrochoric dispersal of generative propagules (seeds), impeding genetic differentiation among populations even over river distances up to 50 km. Although evidence for hydrochoric dispersal of vegetative propagules (clonal plant fragments) was found, this mechanism appeared to be relatively less important. Bayesian-based assignment procedures revealed a number of immigrants, originating from outside our study area, suggesting intercatchment plant dispersal, possibly the result of waterfowl-mediated seed dispersal. This study demonstrates how variation in local environmental conditions in river systems, resulting in shifting balances of sexual vs. asexual reproduction within populations, will affect the genotypic diversity within populations. This study furthermore cautions against generalizations about dispersal of riparian plant species in river systems.     

Speciation through competition: a critical review

We examined causes of speciation in asexual populations in both sympatry and parapatry, providing an alternative explanation for the speciation patterns reported by Dieckmann and Doebeli (1999) and Doebeli and Dieckmann (2003). Both in sympatry and parapatry, they find that speciation occurs relatively easily. We reveal that in the sympatric clonal model, the equilibrium distribution is continuous and the disruptive selection driving evolution of discrete clusters is only transient. Hence, if discrete phenotypes are to remain stable in the sympatric sexual model, there should be some source of nontransient disruptive selection that will drive evolution of assortment. We analyze sexually reproducing populations using the Bulmer's infinitesimal model and show that cost-free assortment alone leads to speciation and disruptive selection only arises when the optimal distribution cannot be matched--in this example, because the phenotypic range is limited. In addition, Doebeli and Dieckmann's analyses assumed a high genetic variance and a high mutation rate. Thus, these theoretical models do not support the conclusion that sympatric speciation is a likely outcome of competition for resources. In their parapatric model (Doebeli and Dieckmann 2003), clustering into distinct phenotypes is driven by edge effects, rather than by frequency-dependent competition.     

Interspecific Competition, Environmental Gradients, Gene Flow, and the Coevolution of Species' Borders

Darwin viewed species range limits as chiefly determined by an interplay between the abiotic environment and interspecific interactions. Haldane argued that species' ranges could be set intraspecifically when gene flow from a species' populous center overwhelms local adaptation at the periphery. Recently, Kirkpatrick and Barton have modeled Haldane's process with a quantitative genetic model that combines density-dependent local population growth with dispersal and gene flow across a linear environmental gradient in optimum phenotype. To address Darwin's ideas, we have extended the Kirkpatrick and Barton model to include interspecific competition and the frequency-dependent selection that it generates, as well as stabilizing selection on a quantitative character. Our model includes local population growth, movements over space, natural selection, and gene flow. It simultaneously addresses the evolution of character displacement and species borders. It reproduces the Kirkpatrick and Barton single-species result that limited ranges can be produced with sufficiently steep environmental gradients and strong dispersal. Further, in the absence of environmental gradients or barriers to dispersal, interspecific competition will not limit species ranges at evolutionary equilibrium. However, interspecific competition can interact with environmental gradients and gene flow to generate limited ranges with much less extreme gradient and dispersal parameters than in the single-species case. Species display character displacement in sympatry, yet the reduction in competition that results from this displacement does not necessarily allow the two species to become sympatric everywhere. When species meet, competition reduces population densities in the region of overlap, which, in turn, intensifies the asymmetry in gene flow from center to margin. This reduces the ability of each species to adapt to local physical conditions at their range limits. If environmental gradients are monotonic but not linear, the transition zone between species at coevolutionary equilibrium occurs where the environmental gradient is steepest. If productivity gradients are also introduced into the model, then patterns similar to Rapoport's rule emerge. Interacting species respond to climate change, as it affects the optimal phenotype over space, by a combination of range shifts and local evolution in mean phenotype, while solitary species respond solely by range shifts. Finally, we compare empirical estimates for intrinsic growth rates and diffusion coefficients for several species to those needed by the single-species model to produce a stable limited range. These empirical values are generally insufficient to produce limited ranges in the model suggesting a role for interspecific interactions.     

Range expansions of sexual versus asexual organisms: Effects of reproductive assurance and migration load

It is generally considered that sexual organisms show faster evolutionary adaptation than asexual organisms because sexuals can accumulate adaptive mutations through recombination. Yet, empirical evidence often shows that the geographic range size of sexual species is narrower than that of closely related asexual species, which may seem as if asexuals can adapt to more varied environments. Two potential explanations for this apparent contradiction considered by the existing theory are reproduction assurance and migration load. Here, we consider both reproductive assurance and migration load within a single model to comparatively examine their effects on range expansions of sexuals and asexuals across an environmental gradient. The model shows that higher dispersal propensity decreases sexuals' disadvantage in reproductive assurance while increasing their disadvantage in migration load. Moreover, lower mutation rate constrains adaptation more strongly in asexuals than in sexuals. Thus, high dispersal propensity and high mutation rates promote that asexuals have wider range sizes than sexuals. Intriguingly, our model reveals that sexuals can have wider geographic range sizes than asexuals under low dispersal propensity and low mutation rates, a pattern consistent with a few exceptional empirical cases. Combining reproductive assurance and migration load provides a useful perspective to better understand the relationships between species' mating systems and their geographic ranges.     

Density‐dependent dispersal and the formation of range borders.

Knowledge about the mechanisms of range formation is crucial for scientifically based species conservation strategies in the face of ongoing global climate change. In recent years an increasing amount of studies have focused on the influences of density‐dependent dispersal on demographic and biogeographical patterns. However, it still remains unclear, to what extent and in what ways this strategy would affect the range formation of species. In order to fill this gap, we present a study using individual‐based simulations of a species with discrete generations living along a dispersal mortality gradient. We compare the evolution of range sizes for species following density‐dependent and density‐independent emigration. Furthermore we assess the influence of environmental stochasticity and Allee effects on range formation, as both processes are known to play an important role for dispersal evolution. We find that density‐dependent dispersal always results in much wider ranges than unconditional dispersal. Increasing environmental stochasticity, a predicted consequence of climate change, can remarkably expand the ranges of species living in such connectivity gradients if dispersal decisions are based on local population density. A strong Allee effect causes range contraction for both strategies, but the effect is considerably less dramatic under density‐dependent compared to density‐independent emigration. We strongly recommend accounting for these findings in future attempts to model species’ range shifts due to climate change.     

Variations in sexual and asexual reproduction of Scirpus mariqueter along an elevational gradient

In order to assess the importance of sexual and asexual reproduction during the life history of Scirpus mariqueter, its reproductive and growth characters were concurrently examined along an elevational gradient (from low elevation to high elevation). The proportions of flowering shoot and inflorescence mass, seed : flower ratio and seed weight were used to quantify the investment in sexual reproduction. The proportions of current-year shoot and rhizome mass were used to quantify the investment in asexual reproduction, and the proportion of corm mass was used for growth, respectively. It was found that vegetative propagation predominated at low elevation, whereas sexual reproduction predominated at high elevation; and that sexual reproduction increased with declining asexual reproduction along the gradient. The results suggest that asexual reproduction is relatively favored in the early life stage, whereas sexual reproduction is favored when the population becomes mature and aged, probably because of the functional differentiation between the two reproductive types. Sexual productive characters (i.e. the proportions of flowering shoot and inflorescence mass) were negatively correlated to both growth and asexual reproductive characters along the gradient, indicating there might exist some trade-offs among growth, sexual and asexual reproduction during the life history. However, no obvious pattern was found between asexual reproductive characters and growth characters along the elevational gradient, possibly because of the varied relationships between them at different life stages. The variations in sexual and asexual reproduction in the species and the relationship between them are thought to be of great significance for local population growth, species persistence and evolution.     

Molecular evidence for limited dispersal of vegetative propagules in the epiphytic lichen Lobaria pulmonaria

Propagation, whether sexual or asexual, is a fundamental step in the life cycle of every organism. In lichenized fungi, a great variety of vegetative propagules have evolved in order for the symbiotic partners to disperse simultaneously. For lichens with the ability of sexual and asexual reproduction, the relative contribution of vegetative dispersal is unknown but could, nonetheless, be inferred by studying genotype distribution. The genetic structure of three Lobaria pulmonaria (Lobariaceae) populations from Switzerland was investigated based on the observed variation at six microsatellite loci. All three populations had a clustered distribution of identical genotypes at small spatial scales. The maximum distance between identical genotypes was 230 m. At a distance of 350 m from a source tree, seemingly suitable habitat patches were too far apart to be colonized. Some multilocus genotypes were frequent within local populations but no genotypes were shared among populations. The restricted occurrences of common genotypes as well as the clustered distributions are evidence for a limited dispersal of vegetative propagules in L. pulmonaria. Gene flow among isolated populations will ultimately depend on the capacity of long-distance dispersal and thus probably depend on sexual reproduction.     

Effects of dispersal plasticity on population divergence and speciation

Phenotypic plasticity is thought to have a role in driving population establishment, local adaptation and speciation. However, dispersal plasticity has been underappreciated in this literature. Plasticity in the decision to disperse is taxonomically widespread and I provide examples for insects, molluscs, polychaetes, vertebrates and flowering plants. Theoretical work is limited but indicates an interaction between dispersal distance and plasticity in the decision to disperse. When dispersal is confined to adjacent patches, dispersal plasticity may enhance local adaptation over unconditional (non-plastic) dispersal. However, when dispersal distances are greater, plasticity in dispersal decisions strongly reduces the potential for local adaptation and population divergence. Upon dispersal, settlement may be random, biased but genetically determined, or biased but plastically determined. Theory shows that biased settlement of either type increases population divergence over random settlement. One model suggests that plasticity further enhances chances of speciation. However, there are many strategies for deciding on where to settle such as a best-of-N strategy, sequential sampling with a threshold for acceptance or matching with natal habitat. To date, these strategies do not seem to have been compared within a single model. Although we are just beginning to explore evolutionary effects of dispersal plasticity, it clearly has the potential to enhance as well as inhibit population divergence. Additional work should pay particular attention to dispersal distance and the strategy used to decide on where to settle.     

Evolution of sexual systems, dispersal strategies and habitat selection in the liverwort genus Radula

? Shifts in sexual systems are among the most common and important transitions in plants and are correlated with a suite of life-history traits. The evolution of sexual systems and their relationships to gametophyte size, sexual and asexual reproduction, and epiphytism are examined here in the liverwort genus Radula. ? The sequence of trait acquisition and the phylogenetic correlations between those traits was investigated using comparative methods. ? Shifts in sexual systems recurrently occurred from dioecy to monoecy within facultative epiphyte lineages. Production of specialized asexual gemmae was correlated to neither dioecy nor strict epiphytism. ? The significant correlations among life-history traits related to sexual systems and habitat conditions suggest the existence of evolutionary trade-offs. Obligate epiphytes do not produce gemmae more frequently than facultative epiphytes and disperse by whole gametophyte fragments, presumably to avoid the sensitive protonemal stage in a habitat prone to rapid changes in moisture availability. As dispersal ranges correlate with diaspore size, this reinforces the notion that epiphytes experience strong dispersal limitations. Our results thus provide the evolutionary complement to metapopulation, metacommunity and experimental studies demonstrating trade-offs between dispersal distance, establishment ability, and life-history strategy, which may be central to the evolution of reproductive strategies in bryophytes.     

基于个体的空间显性模型和遥感技术模拟入侵植物扩张机制

基于个体的空间显性模型和遥感技术,以互花米草为例,模拟了自1997到2010年的种群扩张动态,揭示了土地利用变化与潮间带高程的影响;并通过全局敏感性分析揭示了种子扩散、成体存活率、有性和无性繁殖等种群统计学特征对互花米草种群扩张的相对重要性。研究结果发现:1)有性繁殖与无性繁殖共同决定互花米草种群快速扩张;2)潮间带高程和土地利用变化显著影响模型预测的精度,对互花米草种群扩张有非常重要的影响;3)成体存活率与种子长距离扩散是影响互花米草种群扩张速度最重要的因素;无性繁殖比有性繁殖对种群扩张的影响更大;种子长距离扩散比本地扩散更为重要,同时,小概率的种子长距离扩散事件对种群扩张有非常重要的影响。为了经济有效地控制外来入侵植物的扩张,应该抑制种子的长距离扩散和移除种子长距离扩散形成的位于入侵前沿的小斑块。     

EVOLUTION OF TRADE-OFFS BETWEEN SEXUAL AND ASEXUAL PHASES AND THE ROLE OF REPRODUCTIVE PLASTICITY IN THE GENETIC ARCHITECTURE OF APHID LIFE HISTORIES

Life‐history theory postulates that evolution is constrained by trade‐offs (i.e., negative genetic correlations) among traits that contribute to fitness. However, in organisms with complex life cycles, trade‐offs may drastically differ between phases, putatively leading to different evolutionary trajectories. Here, we tested this possibility by examining changes in life‐history traits in an aphid species that alternates asexual and sexual reproduction in its life cycle. The quantitative genetics of reproductive and dispersal traits was studied in 23 lineages (genotypes) of the bird cherry‐oat aphid Rhopalosiphum padi, during both the sexual and asexual phases, which were induced experimentally under specific environmental conditions. We found large and significant heritabilities (broad‐sense) for all traits and several negative genetic correlations between traits (trade‐offs), which are related to reproduction (i.e., numbers of the various sexual or asexual morphs) or dispersal (i.e., numbers of winged or wingless morphs). These results suggest that R. padi exhibits lineage specialization both in reproductive and dispersal strategies. In addition, we found important differences in the structure of genetic variance–covariance matrices ( G ) between phases. These differences were due to two large, negative genetic correlations detected during the asexual phase only: (1) between fecundity and age at maturity and (2) between the production of wingless and winged parthenogenetic females. We propose that this differential expression in genetic architecture results from a reallocation scheme during the asexual phase, when sexual morphs are not produced. We also found significant G × E interaction and nonsignificant genetic correlations across phases, indicating that genotypes could respond independently to selection in each phase. Our results reveal a rather unique situation in which the same population and even the same genotypes express different genetic (co)variation under different environmental conditions, driven by optimal resource allocation criteria.     

Dispersal and life history strategies in epiphyte metacommunities: alternative solutions to survival in patchy,dynamic landscapes

Host trees for obligate epiphytes are dynamic patches that emerge, grow and fall, and metacommunity diversity critically depends on efficient dispersal. Even though species that disperse by large asexual diaspores are strongly dispersal limited, asexual dispersal is common. The stronger dispersal limitation of asexually reproducing species compared to species reproducing sexually via small spores may be compensated by higher growth rates, lower sensitivity to habitat conditions, higher competitive ability or younger reproductive age. We compared growth and reproduction of different groups of epiphytic bryophytes with contrasting dispersal (asexual vs. sexual) and life history strategies (colonists, short- and long-lived shuttle species, perennial stayers) in an old-growth forest stand in the boreo-nemoral region in eastern Sweden. No differences were seen in relative growth rates between asexual and sexual species. Long-lived shuttles had lower growth rates than colonists and perennial stayers. Most groups grew best at intermediate bark pH. Interactions with other epiphytes had a small, often positive effect on growth. Neither differences in sensitivity of growth to habitat conditions nor differences in competitive abilities among species groups were found. Habitat conditions, however, influenced the production of sporophytes, but not of asexual diaspores. Presence of sporophytes negatively affected growth, whereas presence of asexual diaspores did not. Sexual species had to reach a certain colony size before starting to reproduce, whereas no such threshold existed for asexual reproduction. The results indicate that the epiphyte metacommunity is structured by two main trade-offs: dispersal distance vs. reproductive age, and dispersal distance vs. sensitivity to habitat quality. There seems to be a trade-off between growth and sexual reproduction, but not asexual. Trade-offs in species traits may be shaped by conflicting selection pressures imposed by habitat turnover and connectivity rather than by species interactions. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

The community context of species' borders: ecological and evolutionary perspectives

Species distributional limits may coincide with hard dispersal barriers or physiological thresholds along environmental gradients, but they may also be influenced by species interactions. We explore a number of models of interspecific interactions that lead to (sometimes abrupt) distribution limits in the presence and absence of environmental gradients. We find that gradients in competitive ability can lead to spatial segregation of competitors into distinct ranges, but that spatial movement tends to broaden the region of sympatry between the two species, and that Allee effects tend to sharpen these boundaries. We generalize these simple models to include metapopulation dynamics and other types of interactions including predator–prey and host–parasite interactions. We derive conditions for range limits in each case. We also consider models that include coevolution and gene flow and find that character displacement along environmental gradients can lead to stable parapatric distributions. We conclude that it is essential to consider coevolved species interactions as a potential mechanism limiting species distributions, particularly when barriers to dispersal are weak and environmental gradients are gradual.     

Asexual and sexual reproductive strategies in clonal plants

Most plants can reproduce both sexually and asexually (or vegetatively),and the balance between the two reproductive modes may vary widely between and within species.Extensive clonal growth may affect the evolution of life history traits in many ways.First,in some clonal species,sexual reproduction and sex ratio vary largely among populations.Variation in sexual reproduction may strongly affect plant's adaptation to local environments and the evolution of the geographic range.Second,clonal growth can increase floral display,and thus pollinator attraction,while it may impose serious constraints and evolutionary challenges on plants through geitonogamy that may strongly influence pollen dispersal.Geitonogamous pollination can bring a cost to plant fitness through both female and male functions.Some co-evolutionary interactions,therefore,may exist between the spatial structure and the mating behavior of clonal plants.Finally,a trade-off may exist between sexual reproduction and clonal growth.Resource allocation to the two reproductive modes may depend on environmental conditions,competitive dominance,life span,and genetic factors.If different reproductive modes represent adaptive strategies for plants in different environments,we expect that most of the resources should be allocated to sexual reproduction in habitats with fluctuating environmental conditions and strong competition,while clonal growth should be dominant in stable habitats.Yet we know little about the consequence of natural selection on the two reproductive modes and factors which control the balance of the two reproductive modes.Future studies should investigate the reproductive strategies of clonal plants simultaneously from both sexual and asexual perspectives.     

Evolution of conditional dispersal: evolutionarily stable strategies in spatial models

We consider a two-species competition model in which the species have the same population dynamics but different dispersal strategies. Both species disperse by a combination of random diffusion and advection along environmental gradients, with the same random dispersal rates but different advection coefficients. Regarding these advection coefficients as movement strategies of the species, we investigate their course of evolution. By applying invasion analysis we find that if the spatial environmental variation is less than a critical value, there is a unique evolutionarily singular strategy, which is also evolutionarily stable. If the spatial environmental variation exceeds the critical value, there can be three or more evolutionarily singular strategies, one of which is not evolutionarily stable. Our results suggest that the evolution of conditional dispersal of organisms depends upon the spatial heterogeneity of the environment in a subtle way.     

The relative importance of sexual and asexual reproduction in the spread of Spartina alterniflora using a spatially explicit individual-based model

This study investigates a spatially explicit, individual-based model for simulating the spread of invasive smooth cordgrass (Spartina alterniflora) in Yancheng coastal wetlands from 1995 to 2010. The model, which considers the landscape heterogeneity and changes detected by remote sensing, also reveals the relative importance of sexual and asexual reproduction in the spread by global sensitivity analysis. The model was verified as suitable for simulating the range expansion of S. alterniflora. The results show that: (1) although seedling recruitment is low, it significantly contributes to the range expansion of S. alterniflora. Removing sexual propagation greatly reduces the expansion rate. Rapid expansion requires both sexual and asexual reproduction; (2) in the global sensitivity analysis, the most significant affecters of S. alterniflora invasion were seed dispersal distance, adult survival rate and asexual recruitment survival rate. Sexual propagation contributes much more significantly to quick range expansion than asexual reproduction, but asexual reproduction is the main source of recruitment. Invasion control strategies should target a single reproduction mode. Here, limiting the germination and dispersal of seeds is suggested as a realistic strategy for controlling and managing invasion by this species.