Genetic relationships of the marine invasive crab parasite <Emphasis Type="Italic">Loxothylacus panopaei</Emphasis>: an analysis of DNA sequence variation,host specificity,and distributional range

Host specificity is a key variable of the niche breath of parasites that can be an important determinant of a parasite’s ability to invade new areas. There is increasing evidence that many parasite species may comprise a variety of genetically variable lineages, which differ in host specificity and geographic range. In this study, we (1) explored the extent of diversity in the invasive parasitic barnacle Loxothylacus panopaei (Rhizocephala) infecting mud crabs (2) examined the geographic origin for the invasive lineage and (3) assessed if further southward spread of the parasite may be impeded. Along the US Atlantic coast, L. panopaei infects different hosts in its invaded range (Chesapeake Bay to north of Cape Canaveral) compared to one portion of the native range in Southeast Florida. This difference was reflected in genetic lineages on two independent loci, mitochondrial cytochrome oxidase I and nuclear cytochrome c. Both loci were concordant in that they showed one lineage infecting crabs of the genus Panopeus in the native range and one lineage infecting Eurypanopeus depressus and Rhithropanopeus harrisii hosts in the invaded range and in the Gulf of Mexico, thus indicating Gulf of Mexico populations as the most likely source of introduction into Chesapeake Bay. Interestingly, the nuclear marker resolved an additional lineage of parasites infecting panopeid hosts in the native range. All three parasite lineages were well supported, but a decision about species status must await further analyses. Since its introduction in the 1960s, the invasive L. panopaei lineage has expanded its range southward along the US Atlantic coast, now almost reaching the northern limit of native Panopeus-infecting lineages at Cape Canaveral, Florida. We hypothesized that parasite-free E. depressus in Southeast Florida, living in sympatry with infected panopeid populations, might be resistant to infection by the invasive lineage. Our infection experiments rejected this hypothesis, suggesting that any impediment to further southward range expansion might be expected from temperature regimes of the subtropical zoogeographic region south of Cape Canaveral.     

Phylogeography and cryptic introduction of the ragworm Hediste diversicolor (Annelida,Nereididae) in the Northwest Atlantic

Many benthic marine invertebrates show striking range disjunctions across broad spatial scales. Without direct evidence for endemism or introduction, these species remain cryptogenic. The common ragworm Hediste diversicolor plays a pivotal role in sedimentary littoral ecosystems of the North Atlantic as an abundant prey item and ecosystem engineer, but exhibits a restricted dispersal capacity that may limit connectivity at both evolutionary and ecological time scales. In Europe, H. diversicolor is subdivided into cryptic taxa and genetic lineages whose distributions have been modified by recent invasions. Its origin in the northwest Atlantic has not been adequately addressed. To trace the age and origin of North American ragworm populations, we analyzed mtDNA sequence data (COI) from the Gulf of Maine and Bay of Fundy (n=73 individuals) and compared our findings with published data from the northeast Atlantic. Our results together with previous data indicate that two species of the H. diversicolor complex have independently colonized the northwest Atlantic at least three different times, resulting in two distinct conspecific assemblages in the Bay of Fundy and Gulf of Maine (respectively) that are different from the species found in the Gulf of St. Lawrence. North American populations had significantly lower genetic diversity compared with populations in the northeast Atlantic, and based on patterns of shared identity, populations in the Bay of Fundy originated from the Baltic Sea and North Sea. Populations from the Gulf of Maine were phylogenetically distinct and most likely originated from unsampled European populations. Analyses of the North American populations revealed patterns of post‐colonization gene flow among populations within the Gulf of Maine and Bay of Fundy. However, we failed to detect shared haplotypes between the two regions, and this pattern of complete isolation corroborates a strong phylogeographic break observed in other species.     

Mixed‐stock analysis of wintertime aggregations of striped bass along the Mid‐Atlantic coast

Most larger individuals of migratory striped bass Morone saxatilis from the two major Atlantic coast stocks, the Chesapeake Bay and Hudson River, appear to winter in mid‐Atlantic coastal waters. But it is not known whether they exhibit differential wintertime distributions in accordance with the latitudinal differences in locations of these two estuaries. Mixed‐stock analyses were conducted based on mitochondrial DNA and nuclear DNA genotypic frequencies on wintertime collections of striped bass from coastal waters. No significant differences (P > 0.05) were seen in the proportions of striped bass from the two stocks between collections made from the Delaware Bay mouth and Cape Hatteras in 1997. However, there was a substantially higher Hudson contribution to a 1995 collection from coastal New Jersey (0.349, SD = 0.136) than to the combined 1997 Delaware Bay mouth and Cape Hatteras collection (0.157, SD = 0.072), suggesting this question deserves further study. Additionally, use of the original four reference samples from Chesapeake Bay tributaries (Choptank, Potomac, Rappahannock, Upper Bay) proved adequate alone in characterizing the Chesapeake Bay stock in simulations in which additional tributary collections (Nanticoke, Patuxent, Pocomoke) were added.     

Genetic and historical evidence disagree on likely sources of the Atlantic amethyst gem clam Gemma gemma (Totten, 1834) in California

Aim Historical information about source populations of invasive species is often limited; therefore, genetic analyses are used. We compared inference about source populations from historical and genetic data for the oyster‐associated clam, Gemma gemma that invaded California from the USA Atlantic coast. Location Mid‐Atlantic (North Carolina, Maryland), Northeastern (New Jersey, New York, Massachusetts) and the California coasts (Elkhorn Slough, San Francisco Bay, Bolinas Lagoon, Tomales Bay, Bodega Harbor). Methods The documented history of transplantation of Eastern oysters to California was reviewed. Cytochrome c oxidase subunit I (COI) sequences from recent and archived clams were examined in a haplotype network. We used AMOVA to detect geographic genetic structure and a permutation test for significant reductions in diversity. Results Chesapeake Bay oysters were transplanted to New York prior to shipment to San Francisco Bay and from there to peripheral bays. Gemma in the Northeastern and Mid‐Atlantic regions were genetically differentiated. In California, populations in Bodega Harbor and Tomales Bay were genetically similar to those in the Mid‐Atlantic area while clams in San Francisco Bay, Elkhorn Slough and Bolinas Lagoon resembled populations in the Northeastern region. In California, genetic variation was not highest in San Francisco Bay despite greater magnitude of oyster plantings. Haplotypes varied over time in native and introduced populations. Main Conclusions Historical records and inferences from genetics agree that both Northeastern and Mid‐Atlantic regions were sources for Gemma in California. Only complex genetic hypotheses reconcile the strong segregation of haplotypes in California to the historical evidence of mixing in their proximate source (New York). These hypotheses include sorting of mixtures of haplotypes or selection in non‐native areas. Haplotype turnover in San Francisco and Massachusetts samples over time suggests that the sorting hypothesis is plausible. We suggest, however, that Gemma was introduced independently and recently to Tomales Bay and Bodega Harbor.     

Geographic structure, genetic diversity and source tracking of Spartina alterniflora

Aim To examine the distribution and structure of genetic variation among native Spartina alterniflora and to characterize the evolutionary mechanisms underlying the success of non‐native S. alterniflora. Location Intertidal marshes along the Atlantic, Gulf and Pacific coasts of North America. Methods amova , parsimony analysis, haplotype networks of chloroplast DNA (cpDNA) sequences, neighbour‐joining analysis, Bayesian analysis of population structure, and individual assignment testing were used. Results Low levels of gene flow and geographic patterns of genetic variation were found among native S. alterniflora from the Atlantic and Gulf coasts of North America. The distribution of cpDNA haplotypes indicates that Atlantic coast S. alterniflora are subdivided into ‘northern’ and ‘southern’ groups. Variation observed at microsatellite loci further suggests that mid‐Atlantic S. alterniflora are differentiated from S. alterniflora found in southern Atlantic and New England coastal marshes. Comparisons between native populations on the Atlantic and Gulf coasts and non‐native Pacific coast populations substantiate prior studies demonstrating reciprocal interspecific hybridization in San Francisco Bay. Our results corroborate historical evidence that S. alterniflora was introduced into Willapa Bay from multiple source populations. However, we found that some Willapa Bay S. alterniflora are genetically divergent from putative sources, probably as a result of admixture following secondary contact among previously allopatric native populations. We further recovered evidence in support of models suggesting that S. alterniflora has secondarily spread within Washington State, from Willapa Bay to Grays Harbor. Main conclusions Underlying genetic structure has often been cited as a factor contributing to ecological variation of native S. alterniflora. Patterns of genetic structure within native S. alterniflora may be the result of environmental differences among biogeographical provinces, of migration barriers, or of responses to historical conditions. Interactions among these factors, rather than one single factor, may best explain the distribution of genetic variation among native S. alterniflora. Comprehensive genetic comparisons of native and introduced populations can illustrate how biological invasions may result from dramatically different underlying factors – some of which might otherwise go unrecognized. Demonstrating that invasions can result from several independent or interacting mechanisms is important for improving risk assessment and future forecasting. Further research on S. alterniflora not only may clarify what forces structure native populations, but also may improve the management of non‐native populations by enabling post‐introduction genetic changes and the rapid evolution of life‐history traits to be more successfully exploited.     

Global distribution of cryptic native,introduced and hybrid lineages in the widespread estuarine amphipod Ampithoe valida

Biological invasions can pose a severe threat to coastal ecosystems, but are difficult to track due to inaccurate species identifications and cryptic diversity. Here, we clarified the cryptic diversity and introduction history of the marine amphipod Ampithoe valida by sequencing a mtDNA locus from 683 individuals and genotyping 10,295 single-nucleotide polymorphisms (SNPs) for 349 individuals from Japan, North America and Argentina. The species complex consists of three cryptic lineages: two native Pacific and one native Atlantic mitochondrial lineage. It is likely that the complex originated in the North Pacific and dispersed to the north Atlantic via a trans-arctic exchange approximately 3 MYA. Non-native A. valida in Argentina have both Atlantic mitochondrial and nuclear genotypes, strongly indicating an introduction from eastern North America. In two eastern Pacific estuaries, San Francisco Bay and Humboldt Bay, California, genetic data indicate human-mediated hybridization of Atlantic and Pacific sources, and possible adaptive introgression of mitochondrial loci, nuclear loci, or both. The San Francisco Bay hybrid population periodically undergoes population outbreaks and profoundly damages eelgrass Zostera marina thalli via direct consumption, and these ecological impacts have not been documented elsewhere. We speculate that novel combinations of Atlantic and Pacific lineages could play a role in A. valida’s unique ecology in San Francisco Bay. Our results reinforce the notion that we can over-estimate the number of non-native invasions when there is cryptic native structure. Moreover, inference of demographic and evolutionary history from mitochondrial loci may be misleading without simultaneous survey of the nuclear genome.

     

Diversity and zoogeography of marine Tubificidae (Annelida,Oligochaeta) with notes on variation in widespread species

The specific and generic diversities of the marine Tubificidae (Annelida, Oligochaeta) of the NE Pacific are compared to those of the NE and NW Atlantic as well as to those of Heron Island, Australia. Diversity in the NE Pacific is relatively high when compared to that of the NE Atlantic. The Tubificidae of the NW Atlantic (limited to the eastern coast of the USA) show two distinct zoogeographic regions: Florida to Cape Hatteras; Cape Hatteras to Massachusetts. Diversity, both in terms of the number of species and number of genera, is approximately the same in these two regions, and is similar to that of both the NE Pacific and Heron Island. Evidence suggests that the widespread marine species, in particular Tubificoides pseudogaster, have a range of morphotypes across their distributions. The apparent wide distributions of these species may be due to a taxonomy unable to resolve the differences between the morphotypes. The tubificid oligochaete fauna of the NE Atlantic appears impoverished compared to the other regions examined. The NE Pacific, NW Atlantic, and Heron Island regions are not dominated by one group of species while the NE Atlantic fauna is dominated by Tubificoides benedeni and Clitellio arenarius.     

Low abundance distribution of Pfiesteria piscicida in Pacific and Western Atlantic as detected by mtDNA-18S rDNA real-time polymerase chain reaction

Information on the abundance of Pfiesteria piscicida in thenatural environment is needed for understanding the ecologicalroles of this dinoflagellate. In this study, a real-time polymerasechain reaction (PCR) assay was developed using mitochondrialcytochrome b upstream region and 18S rDNA (PpmtDNA and Pp18S),and the geographic and temporal distribution of P. piscicidawas investigated in several locations. Both PpmtDNA and Pp18Sgenerally gave similar results, indicating that P. piscicidawas present at all studied regions along the American coast(from Maine to North Carolina along the US Atlantic coast andLos Lagos along the Chilean Pacific coast). Despite its widespreaddistribution, P. piscicida was only detected in 36% of the 431water samples analyzed, and its abundance was generally low(<1.0–1.5 cells mL^–1). Populations detected atthe five stations in the Neuse River (North Carolina) and twostations in Chesapeake Bay (Maryland) were genetically homogeneous,whereas those from other locations appeared to be geneticallydiverse. It can be concluded that (i) the PpmtDNA–Pp18Sreal-time PCR assay is sensitive and specific for detectingand quantifying P. piscicida in the natural environment and(ii) P. piscicida is widespread along the American coasts, butnormally only as a minor component of plankton even in the high-riskestuaries (Neuse River, Chesapeake Bay).     

NORTH CAROLINA MARINE ALGAE. IX. ONSLOWIA ENDOPHYTICA GEN. ET SP. NOV. (PHAEOPHYTA,SPHACELARIALES) AND NOTES ON OTHER NEW RECORDS FOR NORTH CAROLINA1

Four benthic algae are reported here for the first time in the North Carolina flora. The new brown algal genus and species, Onslowia endophytica Searles, is described as an endophyte of Halymenia floridana from the North Carolina continental shelf. New records of Boodleopsis pusilla and Naccaria corymbosa from North Carolina constitute range extensions of these tropical species on the American coast north from Florida. Blastophysa rhizopus, an endophyte and epiphyte known from the North Atlantic coast of Europe and America as well as the Caribbean is reported from North Carolina for the first time and in a new host, Predaea feldmannii.     

A new skink species (Oligosoma taumakae sp. nov.; Reptilia: Scincidae) from the Open Bay Islands,New Zealand

Abstract

We describe a new skink species (Oligosoma taumakae sp. nov.) from the Open Bay Islands, New Zealand. This species is diagnosed on the basis of several morphological characteristics, and its specific status is supported by mitochondrial sequence data (ND2, ND4). The new species appears to be most closely related to O. acrinasum, O. infra‐punctatum, O. otagense and O. waimatense. The new taxon appears to be rare and endemic to the island of Taumaka in the Open Bay Islands (off the west coast of the South Island). Predation by a flightless rail (weka, Gallirallus australis), native to New Zealand but introduced to the Open Bay Islands, is a major conservation concern.     

Is Cyclocardia (Conrad) a wastebasket taxon? Exploring the phylogeny of the most diverse genus of the Carditidae (Archiheterodonta,Bivalvia)

The carditid genus Cyclocardia is currently the most diverse genus of the family, including nearly 180 nominal species encompassing wide stratigraphical (Cretaceous–Recent) and geographical (Antarctica, South and North America, Europe, Africa, Alaska, Russia, Japan and New Zealand) ranges. Due to the lack of autapomorphies in the diagnosis of the genus and its large account of species, we re-evaluate the systematic and phylogenetic status of Cyclocardia. We applied three approaches: bibliographic revision, phylogenetic analysis and an exploration of morphological disparity. We used a shell–character matrix comprising 65 taxa (2 outgroups, 29 non-Cyclocardia carditids and 28 species of Cyclocardia) for phylogenetic and disparity analyses. Maximum Observable Rescaled Distances was used to construct a distance matrix to compare Cyclocardia species and other carditid groups. According to our results, Cyclocardia represents a non-monophyletic taxon and is thus a ‘wastebasket taxon’, chiefly because its diagnosis was based mainly on plesiomorphic characters. The European species C. kickxi and C. chameformis are placed within Scalaricardita, and the previously proposed genus Crassicardia is monophyletic (including C. crassidens, C. crebricostata, C. isaotakii and C. rjabininae). Three new genera are proposed for new groups identified by the phylogenetic analysis: South American Oesterheldia gen. nov. (including O. cannada and O. dalek), western North American Coanicardita gen. nov. (including C. ventricosa and C. occidentalis), and North Pacific Hippocampocardia gen. nov. (including H. barbarensis, H. hamiltonensis and H. yakatagensis). The newly defined monophyletic Cyclocardia is restricted to the Atlantic Ocean species C. borealis, C. novangliae and C. compressa.     

88 Preliminary observations on the identity of the red algal genus Caloglossa (delesseriaceae,ceramiales) in the northwestern gulf of mexico

Phylogenetic analyses inferred from rbcL sequences indicates that specimens of Caloglossa (Harvey) G. Martens collected from Vermilion Bay, Louisiana and Galveston, Texas, reported as C. leprieurii (Montagne) G. Martens (1869) constitutes a separate taxon. Studies are presently being conducted to compare this northwestern Gulf of Mexico taxon with material of C. intermedia Kamiya et West in Kamiya et al. (2000, J. Phycol. 36:411‐420) from the Western Atlantic coast and with C. apomeiotica West et Zuccarello in West et al. (1994, Bot. Mar. 37:381–390) from Baja California, Mexico.     

Invasion by a Japanese marine microorganism in western North America

The earliest record in western North America of Trochammina hadai Uchio, a benthic foraminifer common in Japanese estuaries, is from sediment collected in Puget Sound in 1971. It was first found in San Francisco Bay in sediment samples taken in 1983, and since 1986 has been collected at 91% of the sampled sites in the Bay, constituting up to 93% of the foraminiferal assemblage at individual sites. The species is also present in recent sediment samples from 12 other sites along the west coast of North America. The evidence indicates that T. hadai is a recent introduction to San Francisco Bay, and is probably also not native to the other North American sites. Trochammina hadai was probably transported from Japan in ships' ballast tanks, in mud associated with anchors, or in sediments associated with oysters imported for mariculture. Its remarkable invasion of San Francisco Bay suggests the potential for massive, rapid invasions by other marine microorganisms.     

The North-American amphipods, Melita nitida Smith, 1873 and Incisocalliope aestuarius (Watling and Maurer, 1973) (Crustacea: Amphipoda: Gammaridea), introduced to the Western Scheldt estuary (The Netherlands)

The American amphipod species Melita nitida andIncisocalliope aestuarius have been found in the WesternScheldt estuary (the Netherlands). This is the first record of these species inthe north-east Atlantic. Shipping is the most likely vector of introduction.Thedistribution of both species is investigated and compared with the distributionand the microhabitat of co-occurring amphipod species. Melitanitida is known from both the east and west coast of North Americaand I. aestuarius originates from the east coast of NorthAmerica. Until now neither has been reported from other parts of the world. Inthe Netherlands both species are restricted to the mesohaline part of theWestern Scheldt. Melita nitida occurs predominantly underPacific oysters at the underside of boulders, mainly sublittorally.Incisocalliope aestuarius is associated to hydrozoans.Bothmicrohabitats are hardly utilized by other amphipod species. Therefore, thetheory that the existence of many empty niches in north-western Europeanbrackish waters make this environment particularly susceptible to invasions ofalien species is corroborated. The application of hard substrates in a regionoriginally predominated by soft bottoms moreover facilitates the introductionofexotic species. The species community on hard substrates in the mesohaline partof the Western Scheldt contains a high proportion of introduced species:approximately one third of the macrofauna species is of allochthonous origin.     

Seasonal Abundance and Occurrence of the Asian Isopod Synidotea laevidorsalis in Delaware Bay, USA

In 1999 the marine isopod Synidotea laevidorsalis (Miers 1881), indigenous to the northwest Pacific, was first documented in Delaware Bay, USA. We monitored weekly recruitment of this isopod and several other motile species in the Maurice River, a tributary of Delaware Bay. A spatial survey was also conducted. Abundance of S. laevidorsalis varied seasonally but overwhelmingly dominated other co-occurring species by an order of magnitude or more throughout most of the year. Isopod abundance increased through the summer of 2004 and peaked in September, coincident with the passing of Hurricane Ivan. Field observations documented large populations, frequently associated with pilings and buoy lines, throughout Delaware Bay in salinities of 4 through 22 ppt. The dramatic abundance of this isopod indicates that there is considerable potential for altering community structure. This isopod has yet to be observed along the Atlantic Coast of New Jersey or in Chesapeake Bay, but it has been reported near Charleston, SC.     

Predation by blue crabs, Callinectes sapidus, on rapa whelks, Rapana venosa: possible natural controls for an invasive species?

Blue crabs Callinectes sapidus are voracious predators in Chesapeake Bay and other estuarine habitats. The rapa whelk Rapana venosa is native to Asian waters but was discovered in Chesapeake Bay in 1998. This predatory gastropod grows to large terminal sizes (in excess of 150 mm shell length (SL)) and has a thick shell that may contribute to an ontogenetic predation refuge. However, juvenile rapa whelks in Chesapeake Bay may be vulnerable to predation by the blue crab given probable habitat overlap, relative lack of whelk shell architectural defenses, and the relatively large size of potential crab predators. Feeding experiments using three size classes of blue crab predators in relation to a size range of rapa whelks of two different ages (Age 1 and Age 2) were conducted. Blue crabs of all sizes tested consumed Age 1 rapa whelks; 58% of all Age 1 whelks offered were eaten. Age 2 rapa whelks were consumed by medium (67% of whelks offered were eaten) and large (70% of whelks offered were eaten) blue crabs but not by small crabs. The attack methods of medium and large crabs changed with whelk age and related shell weight. Age 1 whelks were typically crushed by blue crabs while Age 2 whelk shells were chipped or left intact by predators removing prey. Rapa whelks less than approximately 35 mm SL are vulnerable to predation by all sizes of blue crabs tested. Rapa whelk critical size may be greater than 55 mm SL in the presence of large blue crabs indicating that a size refugia from crab predation may not be achieved by rapa whelks in Chesapeake Bay until at least Age 2 or Age 3. Predation by blue crabs on young rapa whelks may offer a natural control strategy for rapa whelks in Chesapeake Bay and other estuarine habitats along the North American Atlantic coast.     

Evolution of the Neotropical ant genus Linepithema

Abstract A comprehensive species‐level phylogeny of the ant genus Linepithema Mayr, a Neotropical group best known for the invasive Argentine ant L. humile (Mayr), is inferred for the first time using fragments from three nuclear loci [wingless (WG), long‐wavelength rhodopsin (LWR) and internal transcribed spacer (ITS‐2)] and the mitochondrial cytochrome oxidase subunit I (COI) gene. Monophyly of the genus is strongly supported in parsimony, likelihood and Bayesian analyses of the concatenated data, as is the monophyly of four species groups defined previously on the basis of morphology. An Andean species, L. oblongum (Santschi), is the sister taxon of the Argentine ant. Eight of the 11 species whose monophyly was testable in the analysis were inferred to be monophyletic. Several instances of species paraphyly and one case of mitochondrial introgression suggest that complex population genetic processes underpin the patterns of diversity in Linepithema, and that simple genetic approaches to taxonomy such as DNA barcoding should be treated with caution. A maximum likelihood reconstruction of ancestral distributions suggests that Linepithema is of southern South American origin and that populations in the Greater Antilles are the result of four independent colonization events.     

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.