GENETIC DATA HINT AT A COMMON DONOR REGION FOR INVASIVE ATLANTIC AND PACIFIC POPULATIONS OF GRACILARIA VERMICULOPHYLLA (GRACILARIALES,RHODOPHYTA)1

Gracilaria vermiculophylla (Ohmi) Papenf., an agar‐producing red alga introduced from northeast Asia to Europe and North America, is often highly abundant in invaded areas. To assay its genetic diversity and identify the putative source of invasive populations, we analyzed the mitochondrial cytochrome c oxidase subunit I (cox1) gene from 312 individuals of G. vermiculophylla collected in 37 native and 32 introduced locations. A total of 19 haplotypes were detected: 17 in northeast Asia and three in Europe and eastern and western North America, with only one shared among all regions. The shared haplotype was present in all introduced populations and in ～99% of individuals in the introduced areas. This haplotype was also found at three native locations in east Korea, west Japan, and eastern Russia. Both haplotype and nucleotide diversities were extremely low in Europe and North America compared to northeast Asia. Our study indicates that the East Sea/Sea of Japan is a likely donor region of the invasive populations of G. vermiculophylla in the east and west Atlantic and the east Pacific.     

Flora and fauna associated with the introduced red alga Gracilaria vermiculophylla

This paper presents the first detailed study of the spread of the introduced marine red alga Gracilaria vermiculophylla on the west coast of Sweden, and of the fauna and flora associated with this alga in Scandinavia and the western mid-Atlantic. G. vermiculophylla was discovered in the archipelago of Göteborg, Sweden, in the summer of 2003, and in 2005 its distribution range covered at least 150 km. The species is typically found as loose-lying thalli or attached to small stones and mollusc shells within low-energy bays and estuaries. Both gametophytic and tetrasporophytic specimens were found, as well as specimens with mixed reproductive stages. In order to assess the importance of this introduced alga as a habitat for native benthic organisms, attached and loose-lying individuals of G. vermiculophylla were sampled from invaded locations in Sweden, Denmark and Virginia (United States). In total we found 92 taxa associated with G. vermiculophylla. The dominant classes were Malacostraca, Gastropoda and Florideophyceae. The diversity of the associated taxa was not affected by attachment status, or G. vermiculophylla biomass. In Virginia and Sweden animal abundances were positively correlated with the biomass of algae and plants associated with G. vermiculophylla. If G. vermiculophylla primarily invades non-vegetated soft-sediment estuaries, the invasion may lead to an increase in abundances of small native invertebrates (e.g. gastropods and crustaceans) and epiphytic algae, with likely cascading effects on higher trophic levels.     

Distribution and ecological role of the non-native macroalga Gracilaria vermiculophylla in Virginia salt marshes

Intertidal salt marshes are considered harsh habitats where relatively few stress-resistant species survive. Most studies on non-native species in marshes describe terrestrial angiosperms. We document that a non-native marine macroalga, Gracilaria vermiculophylla, is abundant throughout Virginia’s Atlantic coastline. We sampled eight marshes, characterized by low slopes and by the presence of the tube-building polychaete Diopatra cuprea on adjacent mudflats, which have been shown previously to be associated with G. vermiculophylla. G. vermiculophylla was found in 71% of the sampled quadrats on the border between the mudflat and tall Spartina alterniflora, 51% within the tall S. alterniflora zone, and 12% further inland. We also tagged G. vermiculophylla from two habitats: (1) unattached G. vermiculophylla within marshes and (2) G. vermiculophylla ‘incorporated’ onto D. cuprea tubes on the adjacent mudflats. Of the incorporated thalli, 3–9% ended up in the marsh, demonstrating connectivity between habitats. In addition, 21% of unattached thalli remained for 2 weeks within the marsh, suggesting that entanglement around marsh plants reduces tidal drift. Growth experiments in mesh bags indicate that most of the G. vermiculophylla transferred from the lagoon to the marsh decomposed there, potentially enhancing local nutrient levels. Finally, we document that G. vermiculophylla in marshes had a reduced associated flora and fauna compared to G. vermiculophylla on the adjacent Diopatra mudflats. In conclusion, unattached G. vermiculophylla is abundant along marsh borders in the tall S. alterniflora zone in Virginia, and we hypothesize that this non-native species has significant impacts in terms of marsh habitat complexity, species abundance and diversity, nutrient dynamics, productivity, and trophic interactions.     

GRACILARIA VERMICULOPHYLLA (RHODOPHYTA,GRACILARIALES) IN THE VIRGINIA COASTAL BAYS,USA: COX1 ANALYSIS REVEALS HIGH GENETIC RICHNESS OF AN INTRODUCED MACROALGA

Gracilaria vermiculophylla (Ohmi) Papenfuss is an invasive alga that is native to Southeast Asia and has invaded many estuaries in North America and Europe. It is difficult to differentiate G. vermiculophylla from native forms using morphology and therefore molecular techniques are needed. In this study, we used three molecular markers (rbcL, cox2‐cox3 spacer, cox1) to identify G. vermiculophylla at several locations in the western Atlantic. RbcL and cox2‐cox3 spacer markers confirmed the presence of G. vermiculophylla on the east coast of the USA from Massachusetts to South Carolina. We used a 507 base pair region of cox1 mtDNA to (i) verify the widespread distribution of G. vermiculophylla in the Virginia (VA) coastal bays and (ii) determine the intraspecific diversity of these algae. Cox1 haplotype richness in the VA coastal bays was much higher than that previously found in other invaded locations, as well as some native locations. This difference is likely attributed to the more intensive sampling design used in this study, which was able to detect richness created by multiple, diverse introductions. On the basis of our results, we recommend that future studies take differences in sampling design into account when comparing haplotype richness and diversity between native and non‐native studies in the literature.     

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

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The effect of temporal variability in salinity on the invasive red alga Gracilaria vermiculophylla

Non-native, invasive species are often characterized by being tolerant to environmental stressors, leaving them more fit relative to native species. The red alga Gracilaria vermiculophylla originates from the NW Pacific but has recently spread along the coastlines of Western Europe, where it has become abundant in many shallow, soft-bottom estuaries. Salinity is important for the local and regional distribution of algae. The distribution of G. vermiculophylla in Europe suggests that it thrives well in hyposaline environments and that it may be more fit than some native algae under such conditions. Little, however, is known about the ecophysiology of G. vermiculophylla and it is therefore difficult to predict its spread and future distribution. Laboratory experiments with G. vermiculophylla showed that steady-state salinity above 15?psu was optimal for growth and that the growth rate was reduced at salinities below 15?psu. Variable salinity reduced the growth rate and larger oscillations were more stressful than small ones. Exposure to very low salinity (0–5?psu) was stressful for the alga and algae exposed to these low levels for 2–4 days were unable to recover fully. Gracilaria vermiculophylla did not seem to perform better in hyposaline conditions than many native, estuarine species. The present distribution of G. vermiculophylla in Scandinavia can be explained well by its response to salinity, but this may not explain its present success relative to many naturally occurring algal species.     

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.     

Phenology and thallus size in a non-native population of Gracilaria vermiculophylla

Phenology, or seasonal variation in life cycle events, is poorly described for many macroalgal species. We describe the phenology of a non-native population of Gracilaria vermiculophylla whose thalli are free-living or anchored by decorating polychaetes to tube caps. At a site in South Carolina, USA, we sampled 100 thalli approximately every month from January 2014 to January 2015. We assessed the reproductive state and measured thallus size based on wet weight, thallus length, and thallus surface area from herbarium mounts. Because life cycle stage cannot be assigned using morphology, we implemented a PCR assay to determine the life cycle stage—tetrasporophyte, female gametophyte, or male gametophyte—of each thallus. Tetrasporophytes dominated throughout the year, making up 81%–100% of thalli sampled per month. Reproductive tetrasporophytes varied between 0% and 65% of monthly samples and were most common in warm summer months (July through September) when thalli also tended to be larger. The vast majority of the reproductive thalli were worm-anchored and not fixed to hard substratum via a holdfast. Thus, free-living thalli can be reproductive and potentially seed new non-native populations. Given G. vermiculophylla reproduction seems tied closely to temperature, our work suggests phenology may change with climate-related changes in seawater temperatures. We also highlight the importance of understanding the natural history of macroalgae to better understand the consequence of range expansions on population dynamics.     

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.     

Invasion of novel habitats uncouples haplo‐diplontic life cycles

Baker's Law predicts uniparental reproduction will facilitate colonization success in novel habitats. While evidence supports this prediction among colonizing plants and animals, few studies have investigated shifts in reproductive mode in haplo‐diplontic species in which both prolonged haploid and diploid stages separate meiosis and fertilization in time and space. Due to this separation, asexual reproduction can yield the dominance of one of the ploidy stages in colonizing populations. We tested for shifts in ploidy and reproductive mode across native and introduced populations of the red seaweed Gracilaria vermiculophylla. Native populations in the northwest Pacific Ocean were nearly always attached by holdfasts to hard substrata and, as is characteristic of the genus, haploid–diploid ratios were slightly diploid‐biased. In contrast, along North American and European coastlines, introduced populations nearly always floated atop soft‐sediment mudflats and were overwhelmingly dominated by diploid thalli without holdfasts. Introduced populations exhibited population genetic signals consistent with extensive vegetative fragmentation, while native populations did not. Thus, the ecological shift from attached to unattached thalli, ostensibly necessitated by the invasion of soft‐sediment habitats, correlated with shifts from sexual to asexual reproduction and slight to strong diploid bias. We extend Baker's Law by predicting other colonizing haplo‐diplontic species will show similar increases in asexuality that correlate with the dominance of one ploidy stage. Labile mating systems likely facilitate colonization success and subsequent range expansion, but for haplo‐diplontic species, the long‐term eco‐evolutionary impacts will depend on which ploidy stage is lost and the degree to which asexual reproduction is canalized.     

Genetic structure in chloroplast and nuclear microsatellites in Rosa rugosa around sea straits in northern Japan

Sea‐strait geological history and seawater dispersal patterns can affect the genetic structure of coastal plants. We investigated microsatellite (simple sequence repeat, SSR) variations in chloroplast (cp) and nuclear (n) DNA of the coastal shrub R osa rugosa, native to northeastern Asia, and three species closely related to it. In northern Japan, we sampled R. rugosa along coasts around the Soya and Tsugaru straits with different histories of the Quaternary land formation as a result of different water depths. Both cpSSR and nSSR variations suggest rare hybridization between R . rugosa and the closely related species. Variations in one of two cpSSRs showed genetic differentiation between eastern and western Hokkaido. This genetic structure may result from introgression from R. davurica in eastern Hokkaido and/or isolation between the eastern and western coastlines of Hokkaido as a result of the Quaternary land formation in the Soya Strait. On the other hand, variations in 10 nSSRs were geographically homogeneous with weak isolation by distance along coastlines. These results suggest that extensive gene flow has been homogenizing the genetic structure of R . rugosa in northern Japan.     

Genetic patterns across multiple introductions of the globally invasive crab genus Carcinus

The European green crab Carcinus maenas is one of the world's most successful aquatic invaders, having established populations on every continent with temperate shores. Here we describe patterns of genetic diversity across both the native and introduced ranges of C. maenas and its sister species, C. aestuarii, including all known non‐native populations. The global data set includes sequences from the mitochondrial cytochrome c oxidase subunit I gene, as well as multilocus genotype data from nine polymorphic nuclear microsatellite loci. Combined phylogeographic and population genetic analyses clarify the global colonization history of C. maenas, providing evidence of multiple invasions to Atlantic North America and South Africa, secondary invasions to the northeastern Pacific, Tasmania, and Argentina, and a strong likelihood of C. maenas × C. aestuarii hybrids in South Africa and Japan. Successful C. maenas invasions vary broadly in the degree to which they retain genetic diversity, although populations with the least variation typically derive from secondary invasions or from introductions that occurred more than 100 years ago.     

Home and Away: Comparisons of Resource Utilization by a Marine Species in Native and Invaded Habitats

The exotic Asian shore crab, Hemigrapsus sanguineus, was recently introduced to the northeastern coast of North America and during the 1990's breeding populations were established throughout southern New England. In 1997–1998, ecological studies of several co-occurring brachyuran crabs were conducted and in native (Tanabe Bay, Japan) and invaded (Long Island Sound, USA) habitats of H. sanguineus. Standardized comparisons of H. sanguineus were made between the 2 habitats using data on crab sizes, utilization of space, and food habits. Results revealed that (1) the resource use of H. sanguineus was quite different from that of other resident species in its invaded habitat, and (2) there were no substantial changes in resource utilization by H. sanguineus after it became established in the invaded habitat (relative to native Tanabe Bay). Differing patterns of resource use by H. sanguineus and other crabs in the invaded habitat, the lack of restriction in resource use by H. sanguineus following its introduction, and the climatological and physical similarities between native and invaded regions likely contributed to the successful invasion of H. sanguineus into rocky intertidal habitats in southern New England.     

The desire for variety: Italian wall lizard (Podarcis siculus) populations introduced to the United States via the pet trade are derived from multiple native-range sources

Tests of invasion success often require comparisons between introduced and native populations, but determining the native-range sources for introduced populations can be difficult. Molecular markers can help clarify the geographic extent of native-range sources, helping to identify which populations are appropriate for comparative studies. The Italian Wall Lizard (Podarcis siculus) was introduced multiple times to the United States with extant populations in California, Kansas, New Jersey, and New York. We used phylogeographic analysis of mtDNA sequences (cytb gene) for individuals sampled from these introduced populations and across the native range to identify the number of independent introductions and the location of the source populations. Haplotypes sampled from introduced populations were nested within three geographically distinct, well-supported clades that together encompassed a large portion of the native range. Combining these phylogeographic results with documentation of the introductions revealed putative sources: California individuals are derived from Sicily; Kansas and New York populations are from Tuscany near Florence; and the New Jersey population is likely from the Adriatic coastal region, but a more specific locality is not possible. The pet trade dominates the invasion pathway for P. siculus introductions to the US. The genetically and geographically diverse sampling of its native range may be driven by the desire for phenotypic variety in the pet trade, a hypothesis that needs future testing.     

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.     

Gracilaria vermiculophylla: A western Pacific species of Gracilariaceae (Rhodophyta) first recorded from the eastern Pacific

We report Gracilaria vermiculophylla (Ohmi) Papenfuss from the Pacific coast of North America based on a morphoanatomical revision and comparison of sequences of the nuclear gene coding for the small subunit of ribosomal RNA and the internal transcribed spacers of populations from Baja California, Mexico and Hiroshima, Japan. It is the first convincing report of this species out of its center of distribution in western north Pacific, where it has been considered as a synonym of former ‘G. verrucosa’ records. G. vermiculophylla also occurs in central California as indicated by internal transcribed spacer sequences of a previously unknown Gracilaria Greville material. In the northeastern Pacific G. vermiculophylla is characterized by a robust somewhat vermiform, well‐branched cylindrical thallus, with gradual cell size transition from cortex to medulla, deep spermatangial conceptacles, regular chains of carposporangia and carposporangial initials, downwardly oriented tubular nutritive cells, but rarely with upwardly oriented tubular nutritive cells. In some features of its cystocarp anatomy G. vermiculophylla is related to Gracilariopsis E. Y. Dawson or Hydropuntia Montagne and it is a relevant species for discussions about Gracilariaceae genera.     

High levels of diversity and population structure in the potato late blight pathogen at the Mexico centre of origin

Globally destructive crop pathogens often emerge by migrating out of their native ranges. These pathogens are often diverse at their centre of origin and may exhibit adaptive variation in the invaded range via multiple introductions from different source populations. However, source populations are generally unidentified or poorly studied compared to invasive populations. Phytophthora infestans, the causal agent of late blight, is one of the most costly pathogens of potato and tomato worldwide. Mexico is the centre of origin and diversity of P. infestans and migration events out of Mexico have enormously impacted disease dynamics in North America and Europe. The debate over the origin of the pathogen, and population studies of P. infestans in Mexico, has focused on the Toluca Valley, whereas neighbouring regions have been little studied. We examined the population structure of P. infestans across central Mexico, including samples from Michoacán, Tlaxcala and Toluca. We found high levels of diversity consistent with sexual reproduction in Michoacán and Tlaxcala and population subdivision that was strongly associated with geographic region. We determined that population structure in central Mexico has contributed to diversity in introduced populations based on relatedness of U.S. clonal lineages to Mexican isolates from different regions. Our results suggest that P. infestans exists as a metapopulation in central Mexico, and this population structure could be contributing to the repeated re‐emergence of P. infestans in the United States and elsewhere.     

The invasion of <Emphasis Type="Italic">Senecio pterophorus</Emphasis> across continents: multiple,independent introductions,admixture and hybridization

Senecio pterophorus (Compositae) is a perennial shrub native to eastern South Africa that was introduced into the Western Cape in South Africa and Australia approximately 100 years ago and into Europe (Italy and Spain) more than 25–30 years ago. In this study, the aims were to unravel the putative sources of the introduced populations and identify the changes in genetic diversity after invasion using molecular markers and phylogeographic and population genetic analyses. We sampled the entire area of distribution for S. pterophorus extensively. Based on the results, three lineages were established along a latitudinal and climatic gradient in the native range (south, central, central/north) with high levels of admixture. Multiple, independent introductions occurred in the four invaded ranges. The central/northern lineage (humid climate) was the primary source for all of the invaded regions (with drier climates), although a secondary role was revealed for the southern lineage in the Western Cape and the central/northern lineage in Australia and Spain. The genetic diversity was slightly lower in the Spanish and Australian populations than that in the native populations. A variety of demographic and genetic processes affected the amount and structure of genetic diversity in the invaded areas, including multiple introductions and admixture (Western Cape, Australia and Spain) as well as pre-invasive hybridization (Italy). The patterns of dispersion support a hypothesis of rapid evolution of S. pterophorus after invasion in response to novel climatic conditions.     

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.     

Alien parasitic copepods in mussels and oysters of the Wadden Sea

Molluscan intestinal parasites of the genus Mytilicola, specifically M. intestinalis, were initially introduced into bivalves in the North Sea in the 1930s. It was presumably introduced from the Mediterranean with ship-fouling mussels, then attained epidemic proportions in Mytilus edulis in the 1950s and is now widely established in the North Sea region. Mytilicola orientalis was co-introduced with Pacific oysters to France in the 1970s and in the southern North Sea in the early 1990s. Its main host Crassostrea gigas has massively invaded the Wadden Sea with a concomitant decline in mussels. To explore whether introduced mytilicolid parasites could play a role in the shifting dominance from native mussels to invasive oysters, we analysed 390 mussels and 174 oysters collected around the island of Sylt in the northern Wadden Sea. We show that M. intestinalis has a prevalence >90% and a mean intensity of 4 adult copepods in individual mussels with >50 mm shell length at all sheltered sites. By contrast, none were found in the oysters. However, at one site, we found M. orientalis in C. gigas with a prevalence of 10% and an intensity of 2 per host individual (August 2008). This constitutes the most northern record in Europe for this Pacific parasite until now. Alignments of partial sequences of the mitochondrial cytochrome oxidase I (COI) gene and the nuclear internal transcribed spacers (ITS) and 18S rDNA sequences each show a distinct difference between the two species, which confirms our morphological identification. We suggest that the high parasite load in mussels compared to oysters may benefit the continued expansion of C. gigas in the Wadden Sea.