Structured populations of the oriental fruit moth in an agricultural ecosystem

Intercontinental trade has led to multiple introductions of invasive pest species at a global scale. Molecular analyses of the structure of populations support the understanding of ecological strategies and evolutionary patterns that promote successful biological invasions. The oriental fruit moth, Grapholita (=Cydia) molesta, is a cosmopolitan and economically destructive pest of stone and pome fruits, expanding its distribution range concomitantly with global climate warming. We used ten newly developed polymorphic microsatellite markers to examine the genetic structure of G. molesta populations in an agricultural ecosystem in the Emilia‐Romagna region of northern Italy. Larvae collected in eight sampling sites were assigned to a mosaic of five populations with significant intra‐regional structure. Inferred measures of gene flow within populations implicated both active dispersal, and passive dispersal associated with accidental anthropogenic displacements. Small effective population sizes, coupled with high inbreeding levels, highlighted the effect of orchard management practices on the observed patterns of genetic variation within the sampling sites. Isolation by distance did not appear to play a major role at the spatial scale considered. Our results provide new insights into the population genetics and dynamics of an invasive pest species at a regional scale.     

Population genetic structure of the round goby in Lake Michigan: implications for dispersal of invasive species

Understanding subsequent dispersal of non-native species following introduction is important for predicting the extent and speed of range expansion and is critical for effective management and risk assessment. Post-introduction dispersal may occur naturally or via human transport, but assessing the relative contribution of each is difficult for many organisms. Here, we use data from seven microsatellite markers to study patterns of dispersal and gene flow among 12 pierhead populations of the round goby (Neogobius melanostomus) in Lake Michigan. We find significant population structure among sampling sites within this single Great Lake: (1) numerous populations exhibited significant pairwise F _ST and (2) a Bayesian assignment analysis revealed three distinct genetic clusters, corresponding to different pierhead locations, and genetic admixture between these clusters in the remaining populations. Genetic differentiation (F _ST) is generally related to geographic distance (i.e., isolation by distance), but is periodically interrupted at the scale of Lake Michigan due to gene flow among geographically distant sites. Moreover, average genetic differentiation among populations exhibit a significant, negative correlation with the amount of shipping cargo at ports. Our results, therefore, provide evidence that genetic structure of the round goby in Lake Michigan results from limited natural dispersal with frequent long-distance dispersal through anthropogenic activities such as commercial shipping. Our study suggests that while round gobies can undoubtedly disperse and found new populations through natural dispersal mechanisms, their spread within and among the Great Lakes is likely aided by transport via ships. We, therefore, recommend that ballast-water treatment and management may limit the spread of non-native species within the Great Lakes after the initial introduction in addition to preventing the introduction of non-native species to the Great Lakes.     

Disentangling the dynamics of invasive fireweed (<Emphasis Type="Italic">Senecio madagascariensis</Emphasis> Poir. species complex) in the Hawaiian Islands

Studies investigating the genetic variation of invasive species render opportunities to better understand the dynamics of biological invasions from an ecological and evolutionary perspective. In this study, we investigate fine-scale population genetic structure of invasive Senecio madagascariensis (fireweed) using microsatellite markers to determine levels of genetic diversity and how it pertains to introduction history of this species within and among the Hawaiian Islands. Dispersal patterns were interpreted and, together with a habitat suitability analysis, we aim to describe the potential range expansion of S. madgascariensis within the islands. Bayesian and frequency-based analyses revealed genetic structure with two major genetic demes corresponding to the two fireweed-infested islands of Maui and Hawaii. Both these demes showed further genetic sub-structure, each consisting of three genetically distinct subgroups. Overall, fireweed showed significant levels of inbreeding. Major genetic demes (Maui and Hawaii) differed in observed heterozygosities, inbreeding and genetic structure, each harbouring a large proportion of private alleles. In contrast to the current understanding of fireweed’s introduction history between the Hawaiian Islands, fine-scale population genetic parameters suggest that this species has been introduced at least twice, possibly even more, to the archipelago. Spatial analyses also revealed high correlation between genetic similarity and geographical proximity (>2 km apart) followed by a sharp decline. In addition, a single population was identified that likely resulted from a rare human- or animal-mediated extreme long-distance dispersal event from Maui to Hawaii. Bayesian and likelihood estimates of ‘first generation migrants’ also concurred that contemporary dispersal occurs more frequently over smaller spatial scales than larger scales. These findings indicate that spread in this species occurs primarily via a stratified strategy. Predictions from habitat suitability models indicate all Hawaiian Islands as highly suitable for fireweed invasion and the movement of propagules to currently uninfested islands and outlying suitable habitats should be avoided to circumvent further expansions of the invasion.     

Comparative phylogeography of invasive Rattus rattus and Rattus norvegicus in the U.S. reveals distinct colonization histories and dispersal

Invasive Rattus are arguably the most costly and destructive invasive species on the planet, but little is known concerning their invasion history and population structure in the U.S. We utilized both nuclear microsatellites and mitochondrial DNA sequences (mtDNA) to compare the colonization history, patterns of gene flow, and levels of genetic diversity of Rattus rattus and R. norvegicus in the U.S. Analyses of mtDNA suggest R. rattus is characterized by a single rapid expansion into the U.S. from one or two very closely related mtDNA lineages or geographic sources. For R. norvegicus, mtDNA analyses suggest at least four invasions distinct in space and/or time have occurred to establish its distribution in the U.S. Microsatellite analyses suggest for R. rattus that dispersal is characterized by an isolation-by-distance pattern, suggesting a relatively low frequency of long distance dispersal, and low levels of establishment for novel propagules. In contrast, microsatellite analyses of R. norvegicus suggest high frequencies of long distance dispersal and essentially panmixia among nearly all sampled populations, as well as a high frequency of novel propagules entering at the east and west coasts and assimilating into established populations. We discuss these results in the context of invasive Rattus management in the U.S. and their implications for invasive species in general, as well as the implications for managing the spread of rat-borne pathogens.     

Population genomic analysis suggests strong influence of river network on spatial distribution of genetic variation in invasive saltcedar across the southwestern United States

Understanding the complex influences of landscape and anthropogenic elements that shape the population genetic structure of invasive species provides insight into patterns of colonization and spread. The application of landscape genomics techniques to these questions may offer detailed, previously undocumented insights into factors influencing species invasions. We investigated the spatial pattern of genetic variation and the influences of landscape factors on population similarity in an invasive riparian shrub, saltcedar (Tamarix L.) by analysing 1,997 genomewide SNP markers for 259 individuals from 25 populations collected throughout the southwestern United States. Our results revealed a broad‐scale spatial genetic differentiation of saltcedar populations between the Colorado and Rio Grande river basins and identified potential barriers to population similarity along both river systems. River pathways most strongly contributed to population similarity. In contrast, low temperature and dams likely served as barriers to population similarity. We hypothesize that large‐scale geographic patterns in genetic diversity resulted from a combination of early introductions from distinct populations, the subsequent influence of natural selection, dispersal barriers and founder effects during range expansion.     

Propagule size is not a good predictor for regional population subdivision or fine-scale spatial structure in lichenized fungi

Propagule size has important consequences on the genetic structure of wind-dispersed species, as species with small propagules have higher capability of long-distance dispersal. Here, we studied reproductive modes and compared local and regional population structures in three Macaronesian lichenized fungi differing in propagule size. First, we quantified size distribution of propagules in each species. Second, genotype simulations based on microsatellite data were used to infer the reproductive mode. Third, using spatial analysis and population genetic approaches, we quantified the local and regional scale genetic structures of the fungal species. The three species differed in size distributions of propagules. The majority of populations exhibited clonal reproductive mode. Identical reproductive modes occurred often across species in the same sites, implying a possible relationship between reproductive mode and local site conditions. Contrary to expectation, at the local scale, the species exhibited similar patterns of spatial autocorrelation in genotypes. However, in agreement with the expectation based on propagule size, the species with highest frequency of small vegetative propagules (L. pulmonaria) exhibited lowest regional genetic differentiation. Nevertheless, altogether, our results show that propagule size is not a good predictor of population subdivision in lichenized fungi, neither at local nor regional spatial scale.     

Clonal diversity and spatial genetic structure of Potamogeton pectinatus in managed pond and river populations

Higher levels of genetic diversity of river macrophytes are expected in downstream parts because of potential accumulation of various genotypes from upstream sites. We assessed the clonal diversity and spatial genetic structure of fennel pondweed (Potamogeton pectinatus or Stuckenia pectinata) populations with emphasis on the estimation of dispersal via clonal propagules along a river in connection to upstream ponds. We analysed genetic diversity of 354 plant shoots sampled in 2005 and 2006 at three pond and five river sites in the Woluwe river catchment (Belgium). Nine microsatellite DNA markers revealed 88 genets of which 89% occurred in only one site. Clonal propagule dispersal was detected up to 10 km along the river. Few multilocus genotypes were repeatedly present along a major part of the river indicating vegetative spread. Populations of ponds contained a higher amount of clonal diversity, indicating the importance of local seed recruitment. A fine-scaled spatial genetic structure indicated that most seedling recruitment occurred at a distance <5 m in pond populations whereas clones in river sites were unrelated and showed no spatial autocorrelation. The clonal diversity decreased along the river from upstream to downstream due to establishment of few large clones.     

CONTRIBUTIONS OF SEXUAL AND ASEXUAL REPRODUCTION TO POPULATION STRUCTURE IN THE CLONAL SOFT CORAL,ALCYONIUM RUDYI

Numerous studies of population structure in sessile clonal marine invertebrates have demonstrated low genotypic diversity and nonequilibrium genotype frequencies within local populations that are monopolized by relatively few, highly replicated genets. All of the species studied to date produce planktonic sexual propagules capable of dispersing long distances; despite local genotypic disequilibria, populations are often panmictic over large geographic areas. The population structure paradigm these species represent may not be typical of the majority of clonal invertebrate groups, however, which are believed to produce highly philopatric sexual propagules. I used allozyme variation to examine the population structure of the temperate soft coral, Alcyonium rudyi, a typical clonal species whose sexually produced larvae and asexually produced ramets both have very low dispersal capabilities. Like other clonal plants and invertebrates, the local population dynamics of A. rudyi are dominated by asexual reproduction, and recruitment of new sexually produced genets occurs infrequently. As expected from its philopatric larval stage, estimates of genetic differentiation among populations of A. rudyi were highly significant at all spatial scales examined (mean θ = 0.300 among 20 populations spanning a 1100-km range), suggesting that genetic exchange seldom occurs among populations separated by as little as a few hundred meters. Mapping of multilocus allozyme genotypes within a dense aggregation of A. rudyi ramets confirmed that dispersal of asexual propagules is also very limited: members of the same genet usually remain within < 50 cm of one another on the same rock surface. Unlike most previously studied clonal invertebrates, populations of A. rudyi do not appear to be dominated by a few widespread genets: estimates of genotypic diversity (G_o) within 20 geographically distinct populations did not differ from expectations for outcrossing, sexual populations. Despite theoretical suggestions that philopatric dispersal combined with typically small effective population sizes should promote inbreeding in clonal species, inbreeding does not appear to contribute significantly to the population structure of A. rudyi. Genet genotype frequencies conformed to Hardy-Weinberg expectations in all populations, and inbreeding coefficients (f) were close to zero. In general, the population structure of A. rudyi did not differ significantly from that observed among outcrossing sexual species with philopatric larval dispersal. Age estimates suggest, however, that genets of A. rudyi live for many decades. Genet longevity may promote high genotypic diversity within A. rudyi populations and may be the most important evolutionary consequence of clonal reproduction in this species and the many others that share its dispersal characteristics.     

Looking at both sides of the invasion: patterns of colonization in the violet tunicate Botrylloides violaceus

Understanding the ecological and evolutionary forces that shape the genetic structure of invasive populations and facilitate their expansion across a large spectrum of environments is critical for the prediction of spread and management of ongoing invasions. Here, we study the dynamics of postestablishment colonization in the colonial ascidian Botrylloides violaceus, a notorious marine invader. After its initial introduction from the Northwest Pacific, B. violaceus spread rapidly along the Pacific and Atlantic coasts of North America, impacting both aquaculture facilities and natural ecosystems. We compare genetic diversity and patterns of gene flow among 25 populations (N=679) from the West and East coasts, and evaluate the contribution of sexual vs. asexual reproduction to this species' invasion success using data from the mitochondrial cytochrome c oxidase subunit I (COI) gene and 13 nuclear polymorphic microsatellite loci. Our results reveal contrasting patterns of spread in the coastal waters of North America. While the West coast was colonized by noncontiguous (long-distance) dispersal, the East coast invasion appears to have occurred through contiguous (stepping-stone) spread. Molecular data further indicate that although dispersal in colonial ascidians is predominantly achieved through sexually produced propagules, aquaculture practices such as high-pressure washing can facilitate fragmentation and potentially exacerbate infestations and spread via asexual propagules. The results presented here suggest that caution should be used against the general assumption that all invasions, even within a single species, exhibit similar patterns of colonization, as highly contrasting dynamics may transpire in different invaded ranges.     

Genetic patterns across an invasion's history: a test of change versus stasis for the Eurasian round goby in North America

Biological invasions comprise accidental evolutionary experiments, whose genetic compositions underlie relative success, spread and persistence in new habitats. However, little is known about whether, or how, their population genetic patterns change temporally and/or spatially across the invasion's history. Theory predicts that most would undergo founder effect, exhibit low genetic divergence across the new range and gain variation over time via new arriving propagules. To test these predictions, we analyse population genetic diversity and divergence patterns of the Eurasian round goby Neogobius melanostomus across the two decades of its North American invasion in the Laurentian Great Lakes, comparing results from 13 nuclear DNA microsatellite loci and mitochondrial DNA cytochrome b sequences. We test whether ‘genetic stasis’, ‘genetic replacement’ and/or ‘genetic supplement’ scenarios have occurred at the invasion's core and expansion sites, in comparison with its primary native source population in the Dnieper River, Black Sea. Results reveal pronounced genetic divergence across the exotic range, with population areas remaining genetically distinct and statistically consistent across two decades, supporting ‘genetic stasis’ and ‘founder takes most’. The original genotypes continue to predominate, whose high population growth likely outpaced the relative success of later arrivals. The original invasion core has stayed the most similar to the native source. Secondary expansion sites indicate slight allelic composition convergence towards the core population over time, attributable to some early ‘genetic supplementation’. The geographic and temporal coverage of this investigation offers a rare opportunity to discern population dynamics over time and space in context of invasion genetic theory vs. reality.     

Multiple dispersal vectors drive range expansion in an invasive marine species

The establishment and subsequent spread of invasive species is widely recognized as one of the most threatening processes contributing to global biodiversity loss. This is especially true for marine and estuarine ecosystems, which have experienced significant increases in the number of invasive species with the increase in global maritime trade. Understanding the rate and mechanisms of range expansion is therefore of significant interest to ecologists and conservation managers alike. Using a combination of population genetic surveys, environmental DNA (eDNA) plankton sampling and hydrodynamic modelling, we examined the patterns of introduction of the predatory Northern Pacific seastar (Asterias amurensis) and pathways of secondary spread within southeast Australia. Genetic surveys across the invasive range reveal some genetic divergence between the two main invasive regions and no evidence of ongoing gene flow, a pattern that is consistent with the establishment of the second invasive region via a human‐mediated translocation event. In contrast, hydrodynamic modelling combined with eDNA plankton sampling demonstrated that the establishment of range expansion populations within a region is consistent with natural larval dispersal and recruitment. Our results suggest that both anthropogenic and natural dispersal vectors have played an important role in the range expansion of this species in Australia. The multiple modes of spread combined with high levels of fecundity and a long larval duration in A. amurensis suggests it is likely to continue its range expansion and significantly impact Australian marine ecosystems.     

Reconstruction of a windborne insect invasion using a particle dispersal model,historical wind data,and Bayesian analysis of genetic data

Understanding how invasive species establish and spread is vital for developing effective management strategies for invaded areas and identifying new areas where the risk of invasion is highest. We investigated the explanatory power of dispersal histories reconstructed based on local‐scale wind data and a regional‐scale wind‐dispersed particle trajectory model for the invasive seed chalcid wasp Megastigmus schimitscheki (Hymenoptera: Torymidae) in France. The explanatory power was tested by: (1) survival analysis of empirical data on M. schimitscheki presence, absence and year of arrival at 52 stands of the wasp's obligate hosts, Cedrus (true cedar trees); and (2) Approximate Bayesian analysis of M. schimitscheki genetic data using a coalescence model. The Bayesian demographic modeling and traditional population genetic analysis suggested that initial invasion across the range was the result of long‐distance dispersal from the longest established sites. The survival analyses of the windborne expansion patterns derived from a particle dispersal model indicated that there was an informative correlation between the M. schimitscheki presence/absence data from the annual surveys and the scenarios based on regional‐scale wind data. These three very different analyses produced highly congruent results supporting our proposal that wind is the most probable vector for passive long‐distance dispersal of this invasive seed wasp. This result confirms that long‐distance dispersal from introduction areas is a likely driver of secondary expansion of alien invasive species. Based on our results, management programs for this and other windborne invasive species may consider (1) focusing effort at the longest established sites and (2) monitoring outlying populations remains critically important due to their influence on rates of spread. We also suggest that there is a distinct need for new analysis methods that have the capacity to combine empirical spatiotemporal field data, genetic data, and environmental data to investigate dispersal and invasion.     

Spatial extent of analysis influences observed patterns of population genetic structure in a widespread darter species (Percidae)

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Joint Dispersal Does Not Imply Maintenance of Partnerships in Lichen Symbioses

Dispersal of symbiotic partners by joint propagules is considered as an efficient strategy to maintain successful associations and to circumvent low symbiont availability. Joint dispersal is widespread in diverse symbioses and a particularly common reproductive mode in lichens. We were interested in the implications of joint symbiont dispersal on population genetic structure and investigated patterns of symbiont association in populations of two closely related lichen species in the genus Physconia, with similar range of compatible algal partners. One of the lichen species is characterized by joint dispersal of both symbionts, whereas the other species propagates by meiotic fungal spores alone. The latter species must re-establish the symbiotic stage with appropriate algae sampled from the environment. Both fungal species have specialized on photobionts representing a monophyletic lineage of the algal genus Trebouxia. The results indicate no correlated association of symbiont genotypes in the species with joint symbiont dispersal. We rather show that algal gene diversity in populations of lichenized fungi with different propagation strategies is not necessarily different. The association with algae that differ from the co-dispersed genotypes during the vegetative development of the thalli is the most likely explanation for the observed pattern. Maintenance of symbiotic associations is an option but not a strict consequence of joint symbiont dispersal in lichens.     

Bayesian inference of a complex invasion history revealed by nuclear and chloroplast genetic diversity in the colonizing plant,Silene latifolia

Species invading new ranges are subject to a series of demographic events that can strongly shape genetic diversity. Describing this demographic history is important for understanding where invasive species come from and how they spread, and is critical to testing hypotheses of postinvasion adaptation. Here, we analyse nuclear and chloroplast genetic diversity to study the invasion history of the widespread colonizing weed, Silene latifolia (Caryophyllaceae). Bayesian clustering and PCA revealed strong population structure in the native range of Europe, and although genotypes from multiple native sources were present in the introduced range of North America, the spatial distribution of genetic variance was dramatically reorganized. Using approximate Bayesian computation (ABC), we compared support for different invasion scenarios, including the number and size of independent introduction events and the amount of admixture occurring between sources of introduced genotypes. Our results supported independent introductions into eastern and western North America, with the latter forming a bridgehead for a secondary invasion into the Great Lakes region of central North America. Despite small estimated founder population sizes, the duration of the demographic bottleneck after the initial introduction appeared extremely short‐lived. This pattern of repeated colonization and rapid expansion has effectively eroded the strong population structure and cytonuclear associations present in Europe, but has retained overall high genetic diversity since invasion. Our results highlight the flexibility of the ABC approach for constructing a narrative of the demographic history of species invasions and provide baseline for future studies of evolutionary changes in introduced S. latifolia populations.     

Retention of gene diversity during the spread of a non‐native plant species

Spatial expansion, which is a crucial stage in the process to successful biological invasion, is anticipated to profoundly affect the magnitude and spatial distribution of genetic diversity in novel colonized areas. Here, we show that, contrasting common expectations, Pyrenean rocket (Sisymbrium austriacum), retained SNP diversity as this introduced plant species descended in the Meuse River Basin. Allele frequencies did not mirror between‐population distances along the predominant expansion axis. Reconstruction of invasion history based on the genotypes of historical herbarium specimens indicated no influence of additional introductions or multiple points of entry on this nongradual pattern. Assignment analysis suggested the admixture of distant upstream sources in recently founded downstream populations. River dynamics seem to have facilitated occasional long‐distance dispersal which brought diversity to the expansion front and so maintained evolutionary potential. Our findings highlight the merit of a historical framework in interpreting extant patterns of genetic diversity in introduced species and underscore the need to integrate long‐distance dispersal events in theoretical work on the genetic consequences of range expansion.     

Investigating past range dynamics for a weed of cultivation,Silene vulgaris

Since the last glacial maximum (LGM), many plant and animal taxa have expanded their ranges by migration from glacial refugia. Weeds of cultivation may have followed this trend or spread globally following the expansion of agriculture or ruderal habitats associated with human‐mediated disturbance. We tested whether the range expansion of the weed Silene vulgaris across Europe fit the classical model of postglacial expansion from southern refugia, or followed known routes of the expansion of human agricultural practices. We used species distribution modeling to predict spatial patterns of postglacial expansion and contrasted these with the patterns of human agricultural expansion. A population genetic analysis using microsatellite loci was then used to test which scenario was better supported by spatial patterns of genetic diversity and structure. Genetic diversity was highest in southern Europe and declined with increasing latitude. Locations of ancestral demes from genetic cluster analysis were consistent with areas of predicted refugia. Species distribution models showed the most suitable habitat in the LGM on the southern coasts of Europe. These results support the typical postglacial northward colonization from southern refugia while refuting the east‐to‐west agricultural spread as the main mode of expansion for S. vulgaris. We know that S. vulgaris has recently colonized many regions (including North America and other continents) through human‐mediated dispersal, but there is no evidence for a direct link between the Neolithic expansion of agriculture and current patterns of genetic diversity of S. vulgaris in Europe. Therefore, the history of range expansion of S. vulgaris likely began with postglacial expansion after the LGM, followed by more recent global dispersal by humans.     

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.     

Evidence for non-uniform dispersal in the biological invasions of two naturalized North American bird species

Most previous attempts to model the geographical range expansion of an invading species assume random dispersal of organisms through a homogeneous environment. These models result in a series of uniformly increasing circles radiating out from the centre of origin over time. Although these models often give reasonable fits to available data, they do not typically include mechanisms of dispersal. Alternatively, models that include assumptions of non‐random dispersal and a heterogeneous environment inevitably result in an anisotropic or jagged invasion front. This front will include propagules of pioneer individuals for the expanding species. Existing data from biological invasions reveal that the spatial structure of an invading species usually exhibits these propagules. Using population data gathered from the past century, we investigated the propagules of two North American invading bird species: the European starling (Sturnus vulgaris Linnaeus), and the house finch (Carpodacus mexicanus Müller), and found a correlation between propagule location and habitat quality. These results suggest that dispersing individuals seek out favourable habitat and remain there, thus introducing a possible mechanism for explaining non‐uniform dispersal during invasions. When combined with results from other studies, our results suggest that propagules provide starting points for future population expansion of an invading species.     

Invasion genetics of the Ciona intestinalis species complex: from regional endemism to global homogeneity

Determining the degree of population connectivity and investigating factors driving genetic exchange at various geographical scales are essential to understanding population dynamics and spread potential of invasive species. Here, we explore these issues in the highly invasive vase tunicate, Ciona intestinalis, a species whose invasion history has been obscured by its poorly understood taxonomy and population genetics. Recent phylogenetic and comparative genomic studies suggest that C. intestinalis is a cryptic species complex consisting of at least three species. We reconstructed phylogenies based on both mitochondrial (cytochrome c oxidase subunit 3—NADH dehydrogenase subunit 1 region and NADH dehydrogenase subunit 4 gene) and nuclear (internal transcribed spacer 1) sequences, results of which support four major phylogroups corresponding to the previously reported spA, spB and Ciona spp. (spC) as well as an undescribed cryptic species (spD). While spC and spD remain restricted to their native ranges in the Mediterranean Sea and Black Sea, respectively, the highly invasive species (spA and spB) have disjunct global distributions. Despite extensive interspecific divergences, we identified low phylogeographical structure within these two invasive species. Haplotype network analyses revealed comparatively limited mutation steps among haplotypes within each species. Population genetic analyses based on two mtDNA fragments and eight unlinked microsatellites illustrated relatively low population differentiation and high population connectivity at both regional and continental scales in the two invasive species. Human‐mediated dispersal coupled with a high potential for natural dispersal is probably responsible for the observed genetic homogeneity.