Unravelling the global invasion routes of a worldwide invader,the red swamp crayfish (Procambarus clarkii)

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Genetic structure of an introduced paper wasp,Polistes chinensis antennalis (Hymenoptera,Vespidae) in New Zealand

Several eusocial wasps are prominent invaders to remote islands. The paper wasp Polistes chinensis antennalis is native to East Asia, was introduced to New Zealand in 1979 and has expanded its distribution there. This provides an excellent opportunity to examine the impacts of an initial bottleneck and subsequent expansion on genetic structure. We analysed and compared the genetic population structures of the native (Japan and South Korea) and invasive New Zealand populations. Although 94% of individuals had shared haplotypes detected across both populations, the remaining 6% had private haplotypes identified in only one of the three countries. The genetic variation at microsatellite loci was lower in New Zealand than in native countries, and the genetic structure in New Zealand was clearly distinct from that in its native range. Higher frequencies of diploid‐male‐ and triploid‐female‐producing colonies were detected in New Zealand than in the native countries, showing the reduction in genetic variation via a genetic bottleneck. At least two independent introductions were suggested, and the putative source regions for New Zealand were assigned as Kanto (central island) and Kyushu (south island) in Japan. Serial founder events following the initial introduction were also indicated. The estimated dispersal distance between mother and daughter in New Zealand was twice that in Japan. Thus, the introduction history of P. chinensis antennalis in New Zealand is probably the result of at least two independent introductions, passing through a bottleneck during introduction, followed by population expansion from the point of introduction.     

The worldwide expansion of the Argentine ant

Aim The aim of this study was to determine the number of successful establishments of the invasive Argentine ant outside native range and to see whether introduced supercolonies have resulted from single or multiple introductions. We also compared the genetic diversity of native versus introduced supercolonies to assess the size of the propagules (i.e. the number of founding individuals) at the origin of the introduced supercolonies. Location Global. Methods We used mitochondrial DNA (mtDNA) markers and microsatellite loci to study 39 supercolonies of the Argentine ant Linepithema humile covering both the native (n = 25) and introduced range (n = 14). Results Data from three mitochondrial genes and 13 nuclear microsatellites suggest that the introduced supercolonies studied originated from at least seven founding events out of the native area in Argentina (primary introductions). The distribution of mtDNA haplotypes also suggests that supercolonies in the introduced range each derive from a single source supercolony and that one of these source supercolonies has been particularly successful, being the basis of many introduced populations spread across the world. Comparison of the genetic diversity of supercolonies based on the five most diverse loci also revealed that native and introduced supercolonies have greatly overlapping ranges of diversity, although the genetic diversity is on average less in introduced than in native supercolonies. Main conclusions Both primary introductions (from the native range) and secondary introductions (from sites with established invasive supercolonies) were important in the global expansion of the Argentine ant. In combination with the similar social organization of colonies in the native and introduced range, this indicates that invasiveness did not evolve recently as a unique and historically contingent event (e.g. reduction of genetic diversity) in this species. Rather, native L. humile supercolonies have characteristics that make them pre‐adapted to invade new – and in particular disturbed – habitats when given the opportunity. These results have important implications with regard to possible strategies to be used to control invasive ants.     

BIODIVERSITY RESEARCH: Genetic diversity in two introduced biofouling amphipods (Ampithoe valida & Jassa marmorata) along the Pacific North American coast: investigation into molecular identification and cryptic diversity

Aim We investigated patterns of genetic diversity among invasive populations of Ampithoe valida and Jassa marmorata from the Pacific North American coast to assess the accuracy of morphological identification and determine whether or not cryptic diversity and multiple introductions contribute to the contemporary distribution of these species in the region. Location Native range: Atlantic North American coast; Invaded range: Pacific North American coast. Methods We assessed indices of genetic diversity based on DNA sequence data from the mitochondrial cytochrome c oxidase subunit I (COI) gene, determined the distribution of COI haplotypes among populations in both the invasive and putative native ranges of A. valida and J. marmorata and reconstructed phylogenetic relationships among COI haplotypes using both maximum parsimony and Bayesian approaches. Results Phylogenetic inference indicates that inaccurate species‐level identifications by morphological criteria are common among Jassa specimens. In addition, our data reveal the presence of three well supported but previously unrecognized clades of A. valida among specimens in the north‐eastern Pacific. Different species of Jassa and different genetic lineages of Ampithoe exhibit striking disparity in geographic distribution across the region as well as substantial differences in genetic diversity indices. Main conclusions Molecular genetic methods greatly improve the accuracy and resolution of identifications for invasive benthic marine amphipods at the species level and below. Our data suggest that multiple cryptic introductions of Ampithoe have occurred in the north‐eastern Pacific and highlight uncertainty regarding the origin and invasion histories of both Jassa and Ampithoe species. Additional morphological and genetic analyses are necessary to clarify the taxonomy and native biogeography of both amphipod genera.     

Reconstructing the invasion history of Heracleum persicum (Apiaceae) into Europe

Sparse, incomplete and inappropriate historical records of invasive species often hamper invasive species management interventions. Population genetic analyses of invaders might provide a suitable context for the identification of their source populations and possible introduction routes. Here, we describe the population genetics of Heracleum persicum Desf. ex Fisch and trace its route of introduction into Europe. Microsatellite markers revealed a significantly higher genetic diversity of H. persicum in its native range, and the loss of diversity in the introduced range may be attributed to a recent genetic bottleneck. Bayesian cluster analysis on regional levels identified three and two genetic clusters in the native and the introduced ranges, respectively. A global structure analysis revealed two worldwide distinct genetic groups: one primarily in Iran and Denmark, the other primarily in Norway. There were also varying degrees of admixture in England, Sweden, Finland and Latvia. Approximate Bayesian computation indicated two independent introductions of H. persicum from Iran to Europe: the first one in Denmark and the second one in England. Finland was subsequently colonized by English populations. In contrast to the contemporary hypothesis of English origin of Norwegian populations, we found Finland to be a more likely source for Norwegian populations, a scenario supported by higher estimated histor‐ical migration from Finland to Norway. Genetic diversity per se is not a primary determinant of invasiveness in H. persicum. Our results indicate that, due to either pre‐adaptations or rapid local adaptations, introduced populations may have acqu‐ired invasiveness after subsequent introductions, once a suitable environment was encountered.     

High genetic diversity is not essential for successful introduction

Some introduced populations thrive and evolve despite the presumed loss of diversity at introduction. We aimed to quantify the amount of genetic diversity retained at introduction in species that have shown evidence of adaptation to their introduced environments. Samples were taken from native and introduced ranges of Arctotheca populifolia and Petrorhagia nanteuilii. Using microsatellite data, we identified the source for each introduction, estimated genetic diversity in native and introduced populations, and calculated the amount of diversity retained in introduced populations. These values were compared to those from a literature review of diversity in native, confamilial populations and to estimates of genetic diversity retained at introduction. Gene diversity in the native range of both species was significantly lower than for confamilials. We found that, on average, introduced populations showing evidence of adaptation to their new environments retained 81% of the genetic diversity from the native range. Introduced populations of P. nanteuilii had higher genetic diversity than found in the native source populations, whereas introduced populations of A. populifolia retained only 14% of its native diversity in one introduction and 1% in another. Our literature review has shown that most introductions demonstrating adaptive ability have lost diversity upon introduction. The two species studied here had exceptionally low native range genetic diversity. Further, the two introductions of A. populifolia represent the largest percentage loss of genetic diversity in a species showing evidence of substantial morphological change in the introduced range. While high genetic diversity may increase the likelihood of invasion success, the species examined here adapted to their new environments with very little neutral genetic diversity. This finding suggests that even introductions founded by small numbers of individuals have the potential to become invasive.     

Genetic reconstruction of a bullfrog invasion to elucidate vectors of introduction and secondary spread

Reconstructing historical colonization pathways of an invasive species is critical for uncovering factors that determine invasion success and for designing management strategies. The American bullfrog (Lithobates catesbeianus) is endemic to eastern North America, but now has a global distribution and is considered to be one of the worst invaders in the world. In Montana, several introduced populations have been reported, but little is known of their sources and vectors of introduction and secondary spread. We evaluated the genetic composition of introduced populations at local (Yellowstone River floodplain) and regional (Montana and Wyoming) scales in contrast to native range populations. Our objectives were to (1) estimate the number of introductions, (2) identify probable native sources, (3) evaluate genetic variation relative to sources, and (4) characterize properties of local‐ and regional‐scale spread. We sequenced 937 bp of the mitochondrial cytochrome b locus in 395 tadpoles collected along 100 km of the Yellowstone River, from three additional sites in MT and a proximate site in WY. Pairwise Φ_ST revealed high divergence among nonnative populations, suggesting at least four independent introductions into MT from diverse sources. Three cyt b haplotypes were identical to native haplotypes distributed across the Midwest and Great Lakes regions, and AMOVA confirmed the western native region as a likely source. While haplotype (H_d = 0.69) and nucleotide diversity (π = 0.005) were low in introduced bullfrogs, the levels of diversity did not differ significantly from source populations. In the Yellowstone, two identified haplotypes implied few introduction vectors and a significant relationship between genetic and river distance was found. Evidence for multiple invasions and lack of subsequent regional spread emphasizes the importance of enforcing legislation prohibiting bullfrog importation and the need for continuing public education to prevent transport of bullfrogs in MT. More broadly, this study demonstrates how genetic approaches can reveal key properties of a biological invasion to inform management strategies.     

Preadaptation and post‐introduction evolution facilitate the invasion of Phragmites australis in North America

Compared with non‐invasive species, invasive plant species may benefit from certain advantageous traits, for example, higher photosynthesis capacity and resource/energy‐use efficiency. These traits can be preadapted prior to introduction, but can also be acquired through evolution following introduction to the new range. Disentangling the origins of these advantageous traits is a fundamental and emerging question in invasion ecology. We conducted a multiple comparative experiment under identical environmental condition with the invasive haplotype M lineage of the wetland grass Phragmites australis and compared the ecophysiological traits of this invasive haplotype M in North America with those of the European ancestor and the conspecific North American native haplotype E lineage, P. australis ssp. americanus. The invasive haplotype M differed significantly from the native North American conspecific haplotype E in several ecophysiological and morphological traits, and the European haplotype M had a more efficient photosynthetic apparatus than the native North American P. australis ssp. americanus. Within the haplotype M lineage, the introduced North American P. australis exhibited different biomass allocation patterns and resource/energy‐use strategies compared to its European ancestor group. A discriminant analysis of principal components separated the haplotype M and the haplotype E lineages completely along the first canonical axis, highly related to photosynthetic gas‐exchange parameters, photosynthetic energy‐use efficiency and payback time. The second canonical axis, highly related to photosynthetic nitrogen use efficiency and construction costs, significantly separated the introduced P. australis in North America from its European ancestor. Synthesis. We conclude that the European P. australis lineage was preadapted to be invasive prior to its introduction, and that the invasion in North America is further stimulated by rapid post‐introduction evolution in several advantageous traits. The multicomparison approach used in this study could be an effective approach for distinguishing preadaptation and post‐introduction evolution of invasive species. Further research is needed to link the observed changes in invasive traits to the genetic variation and the interaction with the environment.     

Genetic diversity and colony breeding structure in native and introduced ranges of the Formosan subterranean termite, Coptotermes formosanus

The Formosan subterranean termite, Coptotermes formosanus is recognized as one of the most important invasive pest species. Originating from China, C. formosanus has spread to many tropical and subtropical regions around the globe in the last 400 years, including Japan, Hawaii and the continental USA. Although the current distribution is well documented, information about the patterns of invasion and effects of introduction on the population genetics of this species is largely lacking. We analyzed the genetic structure of populations from two native populations (Guangdong and Hunan provinces, China) and two introduced populations (Maui and Kauai, Hawaii) using microsatellite genotyping. We also reanalyzed published data of additional populations from China, Japan, Hawaii, and the continental USA. The population from Hunan, the earliest introduction outside of China (Japan) and the first introduction to the continental USA (South Carolina) showed little genetic similarity with any of the native or introduced populations investigated. However, populations from Oahu (HI), New Orleans (LA) and Rutherford County (NC) showed close similarity. In general, genetic patterns suggest multiple introductions to the USA, with, for example, two separate introductions to the island of Maui. Bottleneck effects were detected in almost all recent introductions (after 1940). All populations in the introduced range showed lower genetic diversity than those in the native range. However, this low genetic diversity did not result in the formation of polygynous supercolonies as has been described for other invasive termite and ant species.     

The lack of genetic bottleneck in invasive Tansy ragwort populations suggests multiple source populations.

Jacobaea vulgaris (Asteraceae) is a species of Eurasian origin that has become a serious non-indigenous weed in Australia, New Zealand, and North America. We used neutral molecular markers to (1) test for genetic bottlenecks in invasive populations and (2) to investigate the invasion pathways. It is for the first time that molecular markers were used to unravel the process of introduction in this species.The genetic variation of 15 native populations from Europe and 16 invasive populations from Australia, New Zealand and North America were compared using the amplified fragment length polymorphisms (AFLP's). An analysis of molecular variance showed that a significant part (10%) of the total genetic variations between all individuals could be explained by native or invasive origin.Significant among-population differentiation was detected only in the native range, whereas populations from the invasive areas did not significantly differ from each other; nor did the Australian, New Zealand and North American regions differ within the invasive range. The result that native populations differed significantly from each other and that the amount of genetic variation, measured as the number of polymorphic bands, did not differ between the native and invasive area, strongly suggests that introductions from multiple source populations have occurred. The lack of differentiation between invasive regions suggests that either introductions may have occurred from the same native sources in all invasive regions or subsequent introductions took place from one into another invasive region and the same mix of genotypes was subsequently introduced into all invasive regions.An assignment test showed that European populations from Ireland, the Netherlands and the United Kingdom most resembled the invasive populations.     

Getting here from there: testing the genetic paradigm underpinning introduction histories and invasion success

Aim To explore the potential of genetic processes and mating systems to influence successful plant invasions, we compared genetic diversity of the highly invasive tropical treelet, Miconia calvescens, in nine invasive populations and three native range populations. Specifically, we tested how genetic diversity is partitioned in native and invaded regions, which have different invasion histories (multiple vs. single introductions). Lastly, we infer how levels of inbreeding in different regions impact invasion success. Location Invaded ranges in the Pacific (Hawaii, Tahiti, New Caledonia) and Australia and native range in Costa Rica. Methods Genetic diversity was inferred by analysing variation at nine microsatellite loci in 273 individuals from 13 populations of M. calvescens. Genetic structure was assessed using amova , isolation by distance (IBD) within regions, a Bayesian clustering approach, and principal coordinates analysis. Results Microsatellite analysis revealed that invaded regions exhibit low levels of allelic richness and genetic diversity with few private alleles. To the contrary, in the native range, we observed high levels of allelic richness, high heterozygosity and 78% of all private alleles. Surprisingly, despite evident genetic bottlenecks in all invasive regions, similarly high levels of inbreeding were detected in both invasive and native ranges (F_IS: 0.345 and 0.399, respectively). Bayesian clustering analysis showed a lack of geographical structure in the Pacific and evidence of differing invasion histories between the Pacific and Australia. While Pacific populations are derived from a single introduction to the region, multiple introductions have taken place in Australia from different source regions. Main conclusions Multiple introductions have not resulted in increased genetic diversity for M. calvescens invasions. Moreover, similar inbreeding levels between native and invaded ranges suggests that there is no correlation between levels of inbreeding and levels of standing genetic diversity for M. calvescens. Overall, our results show that neither inbreeding nor low genetic diversity is an impediment to invasion success.     

Geographic patterns of genetic diversity from the native range of <Emphasis Type="Italic">Cactoblastis cactorum</Emphasis> (Berg) support the documented history of invasion and multiple introductions for invasive populations

Spread of the invasive cactus-feeding moth Cactoblastis cactorum has been well documented since its export from Argentina to Australia as a biocontrol agent, and records suggest that all non-native populations are derived from a single collection in the moth’s native range. The subsequent global spread of the moth has been complex, and previous research has suggested multiple introductions into North America. There exists the possibility of additional emigrations from the native range in nursery stock during the late twentieth century. Here, we present mitochondrial gene sequence data (COI) from South America (native range) and North America (invasive range) to test the hypothesis that the rapid invasive spread in North America is enhanced by unique genetic combinations from isolated portions of the native range. We found that haplotype richness in the native range of C. cactorum is high and that there was 90% lower richness in Florida than in Argentina. All Florida C. cactorum haplotypes are represented in a single, well-defined clade, which includes collections from the reported region of original export from Argentina. Thus, our data are consistent with the documented history suggesting a single exportation of C. cactorum from the eastern region of the native range. Additionally, the presence of geographic structure in three distinct haplotypes within the same clade across Florida supports the hypothesis of multiple introductions into Florida from a location outside the native range. Because the common haplotypes in Florida are also known to occur in the neighboring Caribbean Islands, the islands are a likely source for independent North American colonization events. Our data show that rapid and successful invasion within North America cannot be attributed to unique genetic combinations. This suggests that successful invasion of the southeastern US is more likely the product of a fortuitous introduction into favorable abiotic conditions and/or defense responses of specific Opuntia hosts, rapid adaptation, or a release from native enemies.     

Contrasting mitochondrial diversity of European starlings (Sturnus vulgaris) across three invasive continental distributions

European starlings (Sturnus vulgaris) represent one of the most widespread and problematic avian invasive species in the world. Understanding their unique population history and current population dynamics can contribute to conservation efforts and clarify evolutionary processes over short timescales. European starlings were introduced to Central Park, New York in 1890, and from a founding group of about 100 birds, they have expanded across North America with a current population of approximately 200 million. There were also multiple introductions in Australia in the mid‐19th century and at least one introduction in South Africa in the late 19th century. Independent introductions on these three continents provide a robust system to investigate invasion genetics. In this study, we compare mitochondrial diversity in European starlings from North America, Australia, and South Africa, and a portion of the native range in the United Kingdom. Of the three invasive ranges, the North American population shows the highest haplotype diversity and evidence of both sudden demographic and spatial expansion. Comparatively, the Australian population shows the lowest haplotype diversity, but also shows evidence for sudden demographic and spatial expansion. South Africa is intermediate to the other invasive populations in genetic diversity but does not show evidence of demographic expansion. In previous studies, population genetic structure was found in Australia, but not in South Africa. Here we find no evidence of population structure in North America. Although all invasive populations share haplotypes with the native range, only one haplotype is shared between invasive populations. This suggests these three invasive populations represent independent subsamples of the native range. The structure of the haplotype network implies that the native‐range sampling does not comprehensively characterize the genetic diversity there. This study represents the most geographically widespread analysis of European starling population genetics to date.     

New insights on the geographical origins of the Caribbean raccoons

The introduction of species outside their natural range is one of the major threats to biodiversity and has often been identified as a menace to agricultural production and human health. The raccoon is recognized as a globally invasive species. However, several populations in the Caribbean were long considered native and endemic species. Although previous genetic studies have shown that raccoons from the islands of the West Indies belong to the northern raccoon Procyon lotor, the history and origin of these introductions remain poorly known. In this study, we investigated the geographical origin of Caribbean raccoon populations using newly available molecular genetic data. We used haplotype network analyses of two mitochondrial markers, Cytochrome b and Control Region, with new sequences and those from GenBank. We also specifically investigated the origin of the endangered endemic Cozumel raccoon, Procyon pygmaeus, by re-analyzing data. Our results confirmed that all Caribbean raccoon populations belong to the northern raccoon. Bahamian populations originated from two different sources in Florida, and the Lesser Antilles raccoons seem to originate from northern regions of the native range. In addition, our results question the taxonomic status of the Cozumel raccoon, as currently available genetic data support a conspecific status with the northern raccoon. These results have important implications in the context of conservation and ecosystem management. Identifying origins of introduced populations and understanding the history of their introductions will facilitate studies on the impact of the raccoon on insular ecosystems.     

Sequence data reflect the introduction pathways of the Sirex woodwasp parasitoid,Ibalia leucospoides (Ibaliidae,Hymenoptera)

1. The parasitoid wasp Ibalia leucospoides is native to the northern hemisphere and has been introduced to the southern hemisphere as a biological control agent for the invasive woodwasp Sirex noctilio. Two subspecies of the parasitoid, Ibalia leucospoides leucospoides (Palearctic distribution) and Ibalia leucospoides ensiger (Nearctic distribution), were introduced and are reported to have hybridized.
2. Despite extensive records of the numbers and origins of the wasps imported into the southern hemisphere, nothing is known regarding their current population diversity. We investigated the genetic variation of I. leucospoides in its native and introduced ranges using mitochondrial (COI) and nuclear (ITS) markers.
3. Mitochondrial DNA diversity in the introduced range was limited, with only five haplotypes, although sequence divergence between these haplotypes was high. Similarly, the ITS rDNA sequences revealed multiple clades present in the introduced range.
4. These results reflect introductions from a wide geographical range but where genetic bottlenecks have possibly reduced the genetic diversity. The data further reflect the origin of the I. leucospoides populations in South America and South Africa from New Zealand or Australia. We found no evidence of hybridization between the two subspecies of the parasitoid in its introduced range, and no evidence that I. leucospoides ensiger has established outside its native range.

     

Invasion of the Hawaiian Islands by a parasite infecting imperiled stream fishes

Points of origin and pathways of spread are often poorly understood for introduced parasites that drive disease emergence in imperiled native species. Co‐introduction of parasites with non‐native hosts is of particular concern in remote areas like the Hawaiian Islands, where the introduced nematode Camallanus cotti has become the most prevalent parasite of at‐risk native stream fishes. In this study, we evaluated the prevailing hypothesis that C. cotti entered the Hawaiian Islands with poeciliid fishes from the Americas, and spread by translocation of poeciliid hosts across the archipelago for mosquito control. We also considered the alternative hypothesis of multiple independent co‐introductions with host fishes originating from Asia. We inferred conduits of introduction and spread of C. cotti across the archipelago from geographic patterns of mtDNA sequence variation and allelic variation across 11 newly developed microsatellite markers. The distribution of haplotypes suggests that C. cotti spread across the archipelago following an initial introduction on O'ahu. Approximate Bayesian Computation modeling and allelic variation also indicate that O'ahu is the most likely location of introduction, from which C. cotti dispersed to Maui followed by spread to the other islands in the archipelago. Evidence of significant genetic structure across islands indicates that contemporary dispersal is limited. Our findings parallel historical records of non‐native poeciliid introductions and suggest that remediating invasion hotspots could reduce the risk of infection in native stream fishes, which illustrates how inferences on parasite co‐introductions can improve conservation efforts by guiding responses to emerging infectious disease in species of concern.     

Pathways of introduction of the invasive aquatic plant Cabomba caroliniana

The pathway and frequency of species' introductions can affect the extent, impact, and management of biological invasions. Here, we examine the pathway of introduction of the aquatic plant Cabomba caroliniana (fanwort) into Canada and the northern United States using plastid DNA sequence (intergenic spacers atpF‐atpH, trnH‐psbA, and trnL‐trnF) and DNA content analyses. We test the hypothesis that the spread of fanwort is a result of commercial trade by comparing a Canadian population (Kasshabog Lake, ON) to native populations from southern U.S., introduced populations in northern U.S., and plants from commercial retailers. Thirteen plastid haplotypes were identified throughout North America, including one dominant haplotype, which was present in all C. caroliniana populations. Several rare haplotypes were used to infer shared colonization history. In particular, the Canadian population shared two rare alleles with a population from Massachusetts, suggesting range expansion of C. caroliniana from the northern U.S. However, the possibility of a commercial introduction cannot be excluded, as common alleles were shared between the Canadian population and both commercial and southern U.S. sources. Variation in C. caroliniana genome size was bimodal and populations were classified into “high” and “low” categories. The Canadian population had DNA contents similar to several northern U.S. populations (low DNA content). This may provide additional support for range expansion from these introduced populations rather than from commercial sources or populations in the southern U.S., which had high DNA content.     

Genetic Bottlenecks in Time and Space: Reconstructing Invasions from Contemporary and Historical Collections

Herbarium accession data offer a useful historical botanical perspective and have been used to track the spread of plant invasions through time and space. Nevertheless, few studies have utilised this resource for genetic analysis to reconstruct a more complete picture of historical invasion dynamics, including the occurrence of separate introduction events. In this study, we combined nuclear and chloroplast microsatellite analyses of contemporary and historical collections of Senecio madagascariensis, a globally invasive weed first introduced to Australia c. 1918 from its native South Africa. Analysis of nuclear microsatellites, together with temporal spread data and simulations of herbarium voucher sampling, revealed distinct introductions to south-eastern Australia and mid-eastern Australia. Genetic diversity of the south-eastern invasive population was lower than in the native range, but higher than in the mid-eastern invasion. In the invasive range, despite its low resolution, our chloroplast microsatellite data revealed the occurrence of new haplotypes over time, probably as the result of subsequent introduction(s) to Australia from the native range during the latter half of the 20^th century. Our work demonstrates how molecular studies of contemporary and historical field collections can be combined to reconstruct a more complete picture of the invasion history of introduced taxa. Further, our study indicates that a survey of contemporary samples only (as undertaken for the majority of invasive species studies) would be insufficient to identify potential source populations and occurrence of multiple introductions.     

Introduced or glacial relict? Phylogeography of the cryptogenic tunicate Molgula manhattensis (Ascidiacea,Pleurogona)

Aim The tunicate Molgula manhattensis has a disjunct amphi‐Atlantic distribution and a recent history of world‐wide introductions. Its distribution could be the result of regional extinctions followed by post‐glacial recolonization, or anthropogenic dispersal. To determine whether the North Atlantic distribution of M. manhattensis is natural or human‐mediated, we analysed mtDNA cytochrome c oxidase subunit I (COI) sequence variation in individuals from cryptogenic and introduced ranges. Location North Atlantic Europe and America; Black Sea; San Francisco Bay; Osaka Bay. Methods Nuclear 18S rDNA sequences were used to resolve phylogenetic relationships and mtDNA COI sequences for phylogeographic analyses. Results Phylogenetic analyses confirmed that M. manhattensis and M. socialis, which are frequently confused, are distinct species. MtDNA haplotype diversity was nearly three times higher with deeper relationships among haplotypes on the North‐east American coast than in Europe. Diversity declined from south to north in America but not in Europe. In areas of known introductions (Black Sea, Japan, San Francisco Bay), M. manhattensis showed variable levels of haplotype diversity. Medium‐to‐high‐frequency haplotypes originating from the North‐west Atlantic were present in two locations of known introductions, but not in Europe. Private haplotypes were found on both sides of the Atlantic and in introduced populations. The mismatch distribution for the North‐east Atlantic coast indicates a recent expansion. Main conclusions Molgula manhattensis is native in North‐east America. However, whether it was introduced or is native to Europe remains equivocal. Additional sampling might or might not reveal the presence of putative private European haplotypes in America. The low European diversity may be explained by low effective population size and a recent expansion, or by low propagule pressure of anthropogenic introduction. Absence of medium‐to‐high‐frequency American haplotypes in Europe may be the result of exclusive transport from southern ports, or long‐term residence. These arguments are ambiguous, and M. manhattensis remains cryptogenic in Europe.