First record of Cynoscion regalis (Pisces,Sciaenidae) in Portuguese continental waters

The occurrence of Cynoscion regalis in Portuguese continental waters is reported for the first time, with six specimens collected in 2015 from three areas: Tagus Estuary, Sado Estuary and Praia da Vieira (central‐west coast). Analyses of morphometric and meristic characteristics confirmed all six specimens as C. regalis; further validation was obtained by sequencing a 675 bp region of the mitochondrial cytochrome c oxidase I gene. These records constitute a range extension of C. regalis into the southern north‐east Atlantic Ocean.     

First records of the North American catfish Ameiurus melas in Portugal

The North American catfish Ameiurus melas is recorded for the first time in Portugal. Its presence in the Tagus River Basin is probably due to natural spread of individuals from Spanish populations, while episodic translocation could explain its occurence in the previously non-invaded Guadiana and Sado River basins.     

Parasitological diversity of the common goby, Pomatoschistus microps (Krøyer, 1838), in estuarine systems along the Portuguese coast

Parasitological diversity of the common goby Pomatoschistus microps along the Portuguese coast was studied based on samples collected using beam trawls in five coastal estuaries (Ria de Aveiro, Tagus, Sado, Mira and Guadiana estuaries) between May and July 2006. The classes Digenea, Cestoda, Nematoda and phylum Mastigophora were found, with Digenea Cryptocotyle sp. and Lecithochirium musculus being the most abundant taxa. Prevalence and mean abundance of Cryptocotyle sp. were highest in Ria de Aveiro (74%) and Mira estuary (28.05), respectively, whereas L. musculus values of both indices were highest in Sado estuary (prevalence = 64%; mean abundance = 11.77). Values varied along the Portuguese coast; the discriminant analysis based on the number of parasites showed some overlap between estuaries, although successfully separating the Sado and Mira estuaries. Overall, variations in estuaries were explained by different environmental pressures, leading to differences on host abundance and life-history of the parasites. Feeding behaviour and habitat selection therefore play a crucial role in shaping the biology, ecology and parasite assemblages of the common goby along the Portuguese coast.     

Multi-locus species tree of the chub genus <Emphasis Type="Italic">Squalius</Emphasis> (Leuciscinae: Cyprinidae) from western Iberia: new insights into its evolutionary history

The phylogenetic relationships of the genus Squalius are believed to be well established based on the mitochondrial cytochrome b gene. Here, we inferred the phylogenetic relationships of all species inhabiting most of the western Iberia river systems using a nuclear multi-locus approach and different species tree methods: concatenation and coalescent-based methods (BEST and minimize-deep-coalescence). The dataset comprised sequences of seven coding and three non-coding regions belonging to seven nuclear genes, which were chosen to cover multiple biological functions: amh, bmp4, ef1a, egr2, irbp, rh and rpl8. We provide evidence for a conflicting topology between the nuDNA species tree and the widely reported mtDNA gene tree. S. pyrenaicus is rendered paraphyletic in all nuDNA species trees, with populations of the Tagus/Colares clustering with S. carolitertii, while populations from the Guadiana, Sado and Almargem form a separate clade. Although a larger sampling size encompassing the full spectrum of Squalius populations in western Iberia is still needed to fully elucidate the phylogeography and species delimitation of this genus, our results suggest that the two S. pyrenaicus clades may represent different species.     

Total PCB-organic matter correlation in sediments from three estuarine areas of Portugal

Levels of tPCB and organic matter content were determined in surface sediments from Sado and Tagus estuaries and Ria Formosa lagoon, and in particles collected in sediment traps in Ria Formosa. Significant relationships (p<0.001) between concentrations of tPCB and organic matter content were obtained when the most contaminated samples and the less contaminated organic-rich sediments were excluded. The excluded samples originated from nearby PCB and organic matter sources. Organic matter appears, thus, the most important indicator of a sediment's sorptive capacity in these estuarine systems, except near the sources where the organic seeking effect is masked. On the basis of the relationship slopes one may conclude that Tagus and Sado estuaries are more contaminated than Alcacer channel (upper Sado estuary) and Ria Formosa.     

Feeding ecology of the lesser weever, Echiichthys vipera (Cuvier, 1829), on the western coast of Portugal

The feeding ecology of the lesser weever, Echiichthys vipera, from the adjacent coastal areas of the Douro and Tagus estuaries (Portugal) was studied between October 2000 and July 2002. The stomach contents of 246 individuals were analysed and diet was characterized by the numerical, gravimetric, occurrence and vacuity indices. Variation of feeding habits with fish length (<95 and >95 mm) and geographical area was considered. Diet of the lesser weever comprised a large variety of prey (28 species), the most important of which were crustaceans (numerical index, NI = 93.5%; occurrence index, OI = 75.6%), namely Mysidacea (especially Schistomysis sp.), Amphipoda (mainly Gammarus subtypicus) and Isopoda (Idotea spp.), and also Teleostei (mostly larval stages that posted a gravimetric index, GI = 53.0%). Diet varied with fish length, with large individuals showing a larger diversity of prey items. Furthermore, specimens from Douro also showed a higher diversity of prey items than those from Tagus. More than 50% of the stomachs were empty, being the highest vacuity values relative to smaller fishes as well as to individuals from the Tagus estuary adjacent coastal area.     

Molecular and morphological divergence in the inland silverside (<Emphasis Type="Italic">Menidia beryllina</Emphasis>) along a freshwater-estuarine interface

Populations that vary across ecological gradients or that have invaded novel habitats are important to elucidate the association between adaptive divergence and gene flow, factors that may play an important role in speciation of silverside fishes. The inland silverside, Menidia beryllina, is an ideal organism for this kind of research, displaying a great diversity in morphology among freshwater and coastal brackish habitats. Using a combination of geometric morphometrics and mitochondrial (mt) DNA, we investigated patterns of variation within and among the nominal freshwater Menidia audens and coastal M. beryllina, spanning the transition from freshwater to tidally influenced semi-brackish waters of the lower Mississippi River to brackish waters of the Lake Pontchartrain estuary. Although we found no evidence for a phylogenetic split between M. audens and M. beryllina, our results indicate that significant genetic divergence corresponds with body shape differences among the two, with a clear distinction at the interface of freshwater and brackish water. Patterns in mtDNA suggest that freshwater populations referred to as M. audens are of recent origin with evidence for habitat-based divergence compared to coastal populations referred to as M. beryllina. Our findings add to a growing body of evidence that ecological shifts, following colonization of novel habitats, may promote rapid adaptive divergence and reduced gene flow among silverside populations in adjacent environmental regimes.     

Numerical modelling of suspended sediment transport in tidal estuaries: A comparison between the Tagus (Portugal) and the Scheldt (Belgium-the Netherlands)

A numerical model to simulate the transport of suspended sediment in tidal estuaries is presented. The model is applied to the two large European estuaries the Tagus (Portugal) and the Scheldt (Belgium-The Netherlands). Calculated suspended sediment concentrations compare favourably with observations in the Tagus (r=0.84) and in the Scheldt (r=0.73). The parametrization scheme indicates that the bottom content of fine sediment is correlated with depth in the Tagus; but a different relationship is used in the Scheldt. Because of tidal range differences, average suspended sediment concentrations are lower in the Tagus (80 mg l⁻¹) than in the Scheldt (130 mg l⁻¹), but a larger relative variation between spring and neap tide concentrations may occur in the Tagus.     

Phylogenetic relationships of assimineid gastropods of the Death Valley–lower Colorado River region: relicts of a late Neogene marine incursion?

Aim A small fauna of amphibious snails (genus Assiminea Fleming, 1828) living in association with highly mineralized springs in the Death Valley–lower Colorado River region (DVLCR) is thought to be a relict of the Bouse Embayment, a putative late Miocene–early Pliocene transgression of the ancestral Gulf of California along the lower Colorado River valley. We analysed the phylogenetic relationships of this fauna using mtDNA sequence data (1171 bp) to determine whether, as would be consistent with this hypothesis, it forms a substantially divergent unit sister to marine coastal congeners. Location South‐western Great Basin and lower Colorado River region, USA. Methods Two genes [mitochondrial cytochrome c oxidase subunit I (COI) and the mitochondrial 16S ribosomal RNA gene] were sequenced for 10 populations of DVLCR assimineas (Assiminea infima Berry, 1947 ; Assiminea sp.). We also sequenced an undescribed population from a spring in the Colorado River delta; western North American Pacific Coastal Assiminea californica (Tryon, 1865); the three other congeners that live on the continent; and three Old World assimineids (outgroups). Phylogenies based on the combined data set were obtained using Bayesian methods, and divergence times were estimated using a COI molecular clock for related gastropods. Results Composite haplotypes of the DVLCR assimineas, together with that observed in the Colorado River delta population, formed a weakly supported clade that was sister to a clade composed of populations of North American Pacific and Atlantic coastal species. The genetic distance between members of these two clades was 3.46 ± 0.47% for COI and 1.69 ± 0.38% for 16S. The former clade was composed of five subunits that differed from each other by 1.29–2.84% (COI) and 0.52–1.98% (16S) sequence divergence. Main conclusions Application of the COI clock suggests that progenitors of the DVLCR fauna diverged from coastal ancestors 2.13–1.89 Ma (late Pliocene), several million years after the Bouse Embayment would have been terminated by the establishment of the lower (freshwater) Colorado River. This finding, together with shallow genetic structuring of several DVLCR lineages that are widely distributed across the topographically complex regional landscape, suggests that the Assiminea fauna of this inland area was more likely to have been founded by coastal colonists transported on water birds than through a direct connection with the sea.     

Composition, temporal changes and ecological guild classification of the ichthyofaunas of large European estuaries – a comparison between the Tagus (Portugal) and the Elbe (Germany)

Fish species densities from 1025 hauls obtained with beam trawl and stow net catches from the southern Tagus and northern Elbe estuaries were analysed for two different periods of time between 1978 and 1997. The data were related with environmental factors to compare species composition and densities, contributions of different ecological guilds, structuring environmental key factors, intra‐annual cyclicity and interperiodical stability of ichthyofaunas in large European estuaries at different latitudes. Although the total number of species in both the Tagus and Elbe estuaries (Tagus: 57 species, Elbe: 58 species) was very similar, significant differences were estimated in species compositions, species densities and number of species, and individuals of the different ecological guilds between both estuaries. A total of 15 species occurred in both estuaries but only two of these species, the catadromous Anguilla anguilla and the marine estuarine opportunist Pleuronectes flesus, contributed substantially to the total abundance. Marine species together contributed more than 5% of the total number of species in the Tagus and Elbe estuaries, whereas their contribution in number of individuals amounted to 22% in the Tagus and only 9% in the Elbe. In terms of number of individuals, the ichthyofauna of the Tagus estuary was dominated by estuarine species (especially Pomatoschistus spp.) contributing 68% of the total abundance. In contrast, the fish fauna of the Elbe estuary was dominated by high numbers of anadromous species (88%) individuals, especially Osmerus eperlanus. Canonical corresponding analyses (CCA) indicated that, besides intra‐annual variables, salinity and mouth distance were two of the most important environmental factors structuring the fish communities in both estuaries. Moreover, water depth was found to be an important environmental factor in the Tagus estuary, whereas the water temperature substantially affected the ichthyofaunal composition in the Elbe estuary. In both estuaries, contributions by number of species and number of individuals of the different ecological guilds were not statistically different between the two periods considered, 1978–80 and 1995–97 for the Tagus, and 1981–86 and 1989–95 for the Elbe. Nevertheless, from 1981–86 until 1989–95 a remarkable increase of about 4.5 times in the densities of O. eperlanus was observed in the Elbe estuary, probably related to improved water quality, especially of oxygen conditions in the nurseries since 1990. This was also supported by the results of CCA, suggesting important influences of the years 1989 and 1990 on the Elbe estuary fish assemblage. Climate change as a result of global warming may be evident for the observed changes of the densities of P. flesus which decreased in the southern Tagus estuary and, in contrast, increased in the northern Elbe estuary during comparable periods of time. It is recommended to use the ecological guild classification scheme which was applied for the Tagus and Elbe estuaries for further studies of European estuarine fish assemblages.     

Estimating fish community diversity from environmental features in the Tagus estuary (Portugal): Multiple Linear Regression and Artificial Neural Network approaches

Relationships between environmental variables and diversity (Shannon‐Weaver index) of the fish communities in the Tagus estuary and adjacent coastal areas were analyzed. The focus was on the linearity or nonlinearity of these abiotic/biotic characteristics, with the aim to obtain an accurate short–medium term time‐scale diversity prediction from habitat variables alone. Multiple Linear Regressions (MLR) were used for the linear approach and Artificial Neural Networks (ANNs) for the nonlinear approach. MLR results in the external validation phase indicated a lack of model accuracy (R² = 0.0710; %SEP = 47.5868; E = ?0.0217; ARV = 1.0217; N = 43). Results of the best of the Artificial Neural Networks used in this study (12‐15‐15‐1 architecture) in the external validation phase (ANN: R² = 0.9736; %SEP = 7.8499; E = 0.9722; ARV = 0.0278; N = 43) were more accurate than those obtained with MLR. This indicates a clear nonlinear relationship between variables. In the best ANN model, nitrate concentration, depth, dissolved oxygen and temperature were the most important predictors of fish diversity in the Tagus estuary. The sensibility analysis indicated that the remaining variables (silicate, nitrite, transparency, salinity, slope, phosphate, water particulate organic matter, and chlorophyll a) played lesser roles in the model.     

Southern limit of distribution of the soft-shell clam <Emphasis Type="Italic">Mya arenaria</Emphasis> on the Atlantic East Coast

The invasive soft-shell clam Mya arenaria was recorded for the first time in the Tagus estuary, central Portugal. Propagules of the soft-shell clam were probably unintentionally introduced into the Tagus estuary through ballast water. Mya arenaria was observed in muddy sands in the upper intertidal zone. The assemblage in which the soft-shell clam was observed was composed of typical estuarine species such as the gastropod Hydrobia ulvae, the bivalve Scrobicularia plana and several polychaete species. The population density of Mya arenaria in the finding area was 40 ind m⁻² with a fresh biomass of 163.5 g FW m⁻².     

DDT concentrations in surficial sediments of three estuarine systems in Portugal

Levels of p,p-DDT and its metabolites in surficial sediments of Tagus and Sado Estuary and lagoon Ria Formosa varied one or two orders of magnitude. In general, p,-DDE and p,-DDD were the dominant metabolites and p,-DDT was a minor component. While concentrations of p,-DDT, p,-DDD and p,-DDE were hardly related to loss on ignition, concentrations of total DDT (tDDT) in surficial sediments of each studied system were positively correlated to that parameter. Higher contaminated samples were excluded from these relationships: a few sediment samples from the Tagus outlet channel, most of the sediments from the upper Sado Estuary, and some particulate matter samples collected in sediment traps in the adjoining area to the Ria Formosa. Concentration of tDDT appears, thus, to be related to sedimentary organic matter of these estuarine systems except near the actual sources of pollution. On the basis of the relationship slopes, one may conclude that lower Tagus Estuary is more contaminated than Ria Formosa and lower Sado Estuary.     

Community shift of ammonia-oxidizing bacteria along an anthropogenic pollution gradient from the Pearl River Delta to the South China Sea

The phylogenetic diversity and abundance of ammonia-oxidizing beta-proteobacteria (beta-AOB) was analyzed along an anthropogenic pollution gradient from the coastal Pearl River Delta to the South China Sea using the ammonia monooxygenase subunit A (amoA) gene. Along the gradient from coastal to the open ocean, the phylogenetic diversity of the dominant genus changed from Nitrosomonas to Nitrosospira, indicating the niche specificity by these two genera as both salinity and anthropogenic influence were major factors involved. The diversity of bacterial amoA gene was also variable along the gradient, with the highest in the deep-sea sediments, followed by the marshes sediments and the lowest in the coastal areas. Within the Nitrosomonas-related clade, four distinct lineages were identified including a putative new one (A5-16) from the different sites over the large geographical area. In the Nitrosospira-related clade, the habitat-specific lineages to the deep-sea and coastal sediments were identified. This study also provides strong support that Nitrosomonas genus, especially Nitrosomonas oligotropha lineage (6a) could be a potential bio-indicator species for pollution or freshwater/wastewater input into coastal environments. A suite of statistical analyses used showed that water depth and temperature were major factors shaping the community structure of beta-AOB in this study area.     

Population structure and genetic diversity of trout (<Emphasis Type="Italic">Oncorhynchus mykiss</Emphasis>) above and below natural and man-made barriers in the Russian River,California

The effects of landscape features on gene flow in threatened and endangered species play an important role in influencing the genetic structure of populations. We examined genetic variation of trout in the species Oncorhynchus mykiss at 22 microsatellite loci from 20 sites in the Russian River basin in central California. We assessed relative patterns of genetic structure and variation in fish from above and below both natural (waterfalls) and man-made (dams) barriers. Additionally, we compared sites sampled in the Russian River with sites from 16 other coastal watersheds in California. Genetic variation among the 20 sites sampled within the Russian River was significantly partitioned into six groups above natural barriers and one group consisting of all below barrier and above dam sites. Although the below-barrier sites showed moderate gene flow, we found some support for sub-population differentiation of individual tributaries in the watershed. Genetic variation at all below-barrier sites was high compared to above-barrier sites. Fish above dams were similar to those from below-barrier sites and had similar levels of genetic diversity, indicating they have not been isolated very long from below-barrier populations. Population samples from above natural barriers were highly divergent, with large F _st values, and had significantly lower genetic diversity, indicating relatively small population sizes. The origins of populations above natural barriers could not be ascertained by comparing microsatellite diversity to other California rivers. Finally, below-barrier sites farther inland were more genetically differentiated from other watersheds than below-barrier sites nearer the river’s mouth.     

The intersection of ecological risk assessment and plant communities: an analysis of <Emphasis Type="Italic">Agrostis</Emphasis> and <Emphasis Type="Italic">Panicum</Emphasis> species in the northeastern U.S.

Ecological risk assessments for grass species with novel traits are advisable, or required, in order to identify potential environmental harms prior to large-scale cultivation. Credible risk assessments are built upon knowledge of the communities that could be negatively affected by crop-to-wild gene flow, new weeds, or invasive plants. This study focused on two cultivated grasses with different life histories: the exotic, weedy Agrostis stolonifera (creeping bentgrass) and the native Panicum virgatum (switchgrass). Vascular plant communities were analyzed in 190 transects (50 m) in ten habitat types across two ecoregions (inland and coastal) in the northeastern U.S. Ordination plots and dendrogram analysis showed clustering of inland plant community assemblages within habitat types, while coastal plant communities were similar across the habitats studied. Agrostis and Panicum species had unequal distribution across the habitat types and ecoregions. Agrostis species were more common in the inland ecoregion and habitats receiving moderate management or disturbance events. In both ecoregions, A. stolonifera had high co-occurrence values with other exotic Agrostis species, suggesting potential for interspecific gene flow. P. virgatum was most common in inland roadside and wasteland habitats, but was distributed equally in the three coastal habitats. Co-occurrence between P. virgatum and congenerics was infrequent, although one transect had both P. virgatum and the state-listed coastal species Panicum amarum. This is the first study to characterize Agrostis and Panicum plant communities and distribution providing the basis for ecological risk assessments, coexistence-strategies, and geographic exclusion zones.     

Modelling the influence of nutrient loads on Portuguese estuaries

The effects of implementing Directive 91/271/EEC of 21 May 1991 (Waste Water Treatment Plan Directive) and Directive 91/676/EEC of 12 December (Nitrates Directive) are analysed in 7 Portuguese estuaries (Minho, Lima, Douro, Mondego, Tagus, Sado and Guadiana) and two coastal lagoons (Ria de Aveiro and Ria Formosa), with a modelling approach. MOHID Water Modelling System was used to perform simulations with three nitrogen load scenarios for each system: a reference scenario, a 50% nitrate removal by agriculture scenario and another with a 100% nutrients removal by waste water treatment plants (WWTP). It is shown that the interaction between hydrodynamic and ecological processes is an important feature to study trophic problems in estuaries. Ecological processes such as primary production only occur inside the system if the residence time of water is high enough to enable organismal activity and if the adequate conditions are found (e.g. light, nutrients, temperature). From the model results it is possible to conclude: (i) in systems with short residence time a reduction in nutrient load will only produce a decrease in nutrient transit and will not affect the system’s global ecological status (e.g. Douro Estuary); (ii) in systems with long residence time the effects will range from significant, when primary production is mostly limited by nutrients (e.g. Ria de Aveiro), to non-significant, when primary production in the system is light-limited (e.g. Tagus Estuary).     

Mitochondrial and nuclear genetic structure in <Emphasis Type="Italic">Rhodeus ocellatus</Emphasis> (Teleostei: Cyprinidae) with approximate Bayesian computation

The rosy bitterling, Rhodeus ocellatus, is a small freshwater fish belonging to the family Cyprinidae. This species lives in ponds where freshwater mussels are abundant and female lays eggs inside mussels. To understand whether the reproduction mode influenced the phylogeography, our study examined the genetic structure of R. ocellatus using sequences from the mitochondrial DNA (cytochrome b gene and control region) and nuclear DNA (the first intron of ribosomal protein S7 gene). In total, 213 specimens were collected from twelve populations in south of Yangtze River, including Yangtze River, Taiwan and Hainan Islands. The phylogenetic analyses based on mitochondrial and nuclear sequences both showed the lack of a population genetic structure, but all results, including the approximate Bayesian computation approaches, showed that these two markers revealed incongruent historical signals. Our study found that (1) the discordance between these two markers were accounted for admixtures by introduced; (2) the phylogeographic pattern of R. ocellatus and that of other freshwater fish were identical; (3) the reproduction mode was due to that the gene flows among populations were limited; and (4) R. ocellatus colonized in south of the Yangtze River (including) after southeast coastal districts formed and before Taiwan Island reached its present shape by comparing our results with that of the previous studies.     

Further Analysis of Anammox Bacterial Community Structures Along an Anthropogenic Nitrogen-Input Gradient from the Riparian Sediments of the Pearl River Delta to the Deep-Ocean Sediments of the South China Sea

The community structures of anammox bacteria in sediments along an anthropogenic inorganic nitrogen input gradient were further delineated with the newly available information incorporated. Anammox bacterial 16S rRNA gene-amplified sequences retrieved from riparian sediments of the Pearl River, Mai Po coastal wetland, and the South China Sea (SCS) sediments were compiled, compared and analyzed. Results indicated that the community structures of anammox bacteria varied from the upstream of the Pearl River to deep-ocean sediment of the SCS along the anthropogenic input grandient. Mai Po wetland had the most diverse anammox bacteria, followed by the shallow SCS, deep SCS and the Pearl River. Genera of the anammox bacteria Kuenenia and Brocadia showed higher proportion in the riparian sediments of the Pearl River, while those of Kuenenia and Scalindua dominated the Mai Po coastal wetland. The Scalindua subclusters showed apparent segregation in coastal wetland (S. zhenghei-III and S. wagneri), shallow SCS (S. zhenghei-I and S3) and deep SCS (S. zhenghei-I, S2 and S. arabica). Pearson correlation analysis indicated nitrogen species [NH₄⁺ and ∑(NO₂^?+NO₃^? )] negatively correlated with the diversity indices of anammox bacteria. Canonical correspondence analysis (CCA) showed that salinity, inorganic nitrogen [NH₄+, ∑(NO₂^?+NO₃^?)], and ratio of NH₄⁺/∑(NO₂^? +NO₃^?) significantly affected the bacterial community compositions. Results collectively support that the community composition of anammox bacteria can serve as a bio-indicator to the anthropogenic terrestrial N input or pollution.     

Genetic structure analysis of mantis shrimp <Emphasis Type="Italic">Oratosquilla oratoria</Emphasis> based on mitochondrial DNA control region sequence

The genetic relationships of mantis shrimp Oratosquilla oratoria between the coastal waters of China and Japan were not well studied. To reveal the genetic differentiation and genetic structure among populations, we collected populations of mantis shrimp O. oratoria from the coastal waters of China and Japan to analyze the mtDNA control region variation. A total of 309 individuals of O. oratoria were collected from 13 localities (11 from China and 2 from Japan) and a segment of mitochondrial DNA control region was sequenced. Three hundred nine haplotypes were defined, yielding a very high haplotype diversity and nucleotide diversity. Two lineages of O. oratoria were revealed and displayed strong differences in the geographical distribution. In the coastal waters of China, the geographic distribution of the two lineages was completely separated by the Yangtze River estuary; however, the lineages showed geographic sympatry in two populations from Japan. Based on the lineage distribution, three groups were defined. There was no significant genetic differentiation among the populations within the three groups, indicating high gene flow within each group. Significant and negative values for Tajima D and Fu’s Fs tests, and mismatch distributions for two lineages indicated population expansion. The present result confirmed that the freshwater outflow from the Yangtze River formed a physical barrier and affected gene exchange. The different distribution patterns of the two lineages in coastal waters of China and Japan indicated that the larvae of O. oratoria were transferred from China to the coastal waters of Japan with a one-way gene flow.