An overview of grey mullet (Mugilidae) global occurrence and species-rich ecoregions, with indications of possible past dispersal routes within the family.

This review examines the published literature on the distribution and species richness of the family Mugilidae around six continents as well as their phylogenetic relationships in a time-calibrated tree. Three mugilid species-rich regions were identified globally, namely the Coral Triangle, southern Asia and southern Africa, all of which have between 16 and 18 morphologically recognized species. Two of the species hotspots are tropical, and only southern Africa incorporates temperate waters. The centre of mugilid evolution and then global dispersion appears to be located in the tropical waters of the Indo-Pacific. Speciation within the family was promoted mainly by plate tectonics, eustatic sea-level changes, tropical climate and high habitat diversity within this region. The number of these tropical species that radiated out to the coastal waters of Africa and America was clearly less than the number that remained in the central Indo-Pacific region. Nonetheless, access by tropical mugilid species was greater to Africa than the Americas because of the direct tropical and subtropical coastal connectivity to the former continent. This enabled more tropical mugilids to reach Africa than America, with the latter probably relying on Palaeocene "island hopping" or "rafting" to move eastwards across the Pacific Ocean. In addition, tropical mugilids were able to access warm western and central African coastal waters from the Pacific prior to the closure of the Tethys Sea gap in the eastern Mediterranean and prior to the development of the cool Benguela Current upwelling off the southwest African coast, and from America through at least two transatlantic rafting events.

Genetic diversity and structure of circumtropical almaco jack, Seriola rivoliana: tool for conservation and management.

The almaco jack, Seriola rivoliana, is a circumtropical pelagic fish of importance both in commercial fisheries and in aquaculture. To understand levels of genetic diversity within and among populations in the wild, population genetic structure and the relative magnitude of migration were assessed using mtDNA sequence data and single nucleotide polymorphisms (SNPs) from individuals sampled from locations in the Pacific and Atlantic Oceans. A total of 25 variable sites of cytochrome c oxidase subunit 1 and 3678 neutral SNPs were recovered. Three genetic groups were identified, with both marker types distributed in different oceanic regions: Pacific-1 in central Pacific, Pacific-2 in eastern Pacific and Atlantic in western Atlantic. Nonetheless, the analysis of SNP identified a fourth population in the Pacific coast of Baja California Sur, Mexico (Pacific-3), whereas that of mtDNA did not. This mito-nuclear discordance is likely explained by a recently diverged Pacific-3 population. In addition, two mtDNA haplogroups were found within the western Atlantic, likely indicating that the species came into the Atlantic from the Indian Ocean with historical gene flow from the eastern Pacific. Relative gene flow among ocean basins was low with r m < 0.2, whereas in the eastern Pacific it was asymmetric and higher from south to north (r m > 0.79). The results reflect the importance of assessing genetic structure and gene flow of natural populations for the purposes of sustainable management.

Multilocus resolution of Mugilidae phylogeny (Teleostei: Mugiliformes): Implications for the family's taxonomy.

The interrelationships among mugilids (Mugiliformes: Mugilidae) remain highly debated. Using a mitochondrial gene-based phylogeny as criterion, a revised classification with 25 genera in the Mugilidae has recently been proposed. However, phylogenetic relationships of major mitochondrial lineages remain unresolved and to gain a general acceptance the classification requires confirmation based on multilocus evidence and diagnostic morphological characters. Here, we construct a species-tree using twelve nuclear and three mitochondrial loci and infer the evolution of 71 morphological characters. Our multilocus phylogeny does not agree with previous morphology-based hypotheses for the relationships within Mugilidae, confirms the revised classification with 25 genera and further resolves their phylogenetic relationships. Using the well-resolved multilocus phylogeny as the criterion, we reclassify Mugilidae genera into three new subfamilies (Myxinae, Rhinomugilinae, and Cheloninae) and one new, recombined, subfamily (Mugilinae). The Rhinomugilinae subfamily is further divided into four tribes. The revised classification of Mugilidae is supported by morpho-anatomical synapomorphies or a combination of characters. These characters are used to erect a key to the subfamilies and genera.

A comparative study of Ligophorus uruguayense and L. saladensis (Monogenea: Ancyrocephalidae) from Mugil liza (Teleostei: Mugilidae) in southern Brazil.

Representatives of Ligophorus Euzet et Suriano, 1977 were found on the gills of Mugil liza Valenciennes caught in southern Brazil. They were identified as Ligophorus uruguayense Failla Siquier et Ostrowski de Núñez, 2009 and Ligophorus saladensis Marcotegui et Martorelli, 2009, even though specific identification proved to be difficult due to inconsistencies in some diagnostic features reported for these two species. Therefore, a combined morphological and molecular approach was used to critically review the validity of these species, by means of phase contrast and confocal fluorescence microscopical examination of sclerotised hard parts, and assessing the genetic divergence between L. saladensis, L. uruguayense and their congeners using rDNA sequences. The main morphological differences between the two species relate to the shape of the accessory piece of the penis and the median process of the ventral bar. The accessory piece in L. uruguayense is shorter than in L. saladensis, has a cylindrical, convex upper lobe and straight lower lobe (vs with the distal tip of the lower lobe turning away from the upper lobe in the latter species). The ventral bar has a V-shaped anterior median part in L. uruguayense (vs U-shaped in L. saladensis). The two species are suggested to be part of a species complex together with L. mediterraneus Sarabeev, Balbuena et Euzet, 2005. We recommend to generalise such comparative assessment of species of Ligophorus for a reliable picture of the diversity and diversification mechanisms within the genus, and to make full use of its potential as an additional marker for mullet taxonomy and systematics.

Taxonomic revision of the Southeast Asian brook barb genus Poropuntius Smith, 1931 (Teleostei, Cyprinidae) with description of a new species from Vietnam.

Molecular data from samples encompassing 22 nominal species of Poropuntius indicate that the species-level diversity in the genus has been vastly overestimated, likely due to inadequate taxon and geographic sampling and reliance on morphological characters that vary intra-specifically. The latter includes discrete mouth morphologies related to alternate feeding strategies (ecomorphs) within populations. One new species is described, Poropuntiusanlaoensis Hoàng, Pham & Tran, sp. nov., and 17 synonyms of six valid species names of Poropuntius, P.krempfi, P.alloiopleurus, P.huangchuchieni, P.laoensis, P.kontumensis, and P.deauratus, are recognised. Additional taxonomic changes in this widespread and generally poorly known genus are likely as more molecular and morphological data become available.

Mitochondrial haplotypes indicate parapatric-like phylogeographic structure in blue-spotted maskray (Neotrygon kuhlii) from the Coral Triangle region.

Phylogeographic structure was investigated in the blue-spotted maskray, Neotrygon kuhlii, focusing on the Coral Triangle region. We used as genetic marker a 519-bp fragment of the cytochrome c-oxidase subunit I (COI) gene, sequenced in a total of 147 individuals from 26 sampling locations. The parsimony network of COI haplotypes was split into seven distinct clades within the Coral Triangle region. Different clades had exclusive but contiguous geographic distributions, indicating parapatric-like phylogeographic structure. Strong genetic differences were also inferred between local populations within a clade, where reciprocal monophyly between geographically adjacent samples was observed on several instances. Nearly 25% of the total molecular variance could be ascribed to differences between geographic samples within a clade, whereas interclade variation accounted for >65% of the total variance. The strong phylogeographic structure observed within a clade can be explained by either sedentarity or female philopatry. We interpret the parapatric distribution of clades as the joint result of 1) expansion from refuge populations at times of low sea level, and 2) possible enhanced competition between individuals from different clades, or assortative mating, or hybrid zones, along lines of secondary contact. The parapatric-like structure uncovered in the present study parallels regional differences at nuclear marker loci, thus pointing to incipient speciation within Coral Triangle N. kuhlii.

Ligophorus species (Monogenea: Ancyrocephalidae) from Mugil cephalus (Teleostei: Mugilidae) off Morocco with the description of a new species and remarks about the use of Ligophorus spp. as biological markers of host populations.

Gill monogenean species of Ligophorus Euzet et Suriano, 1977 were studied from the teleost Mugil cephalus Linneaus (Mugilidae) from the Mediterranean and Atlantic coasts of Morocco. We report the presence of L. mediterraneus from both the Mediterranean and Atlantic coast and L. cephali and L. maroccanus sp. n. from the Atlantic coast only. The latter species, which is described herein as new, resembles L. guanduensis but differs from this species mainly in having a shorter penis compared to the accessory piece, a proportionally longer extremity of the accessory piece and a less developed heel. The utility of Ligophorus spp. as markers of cryptic species of the complex M. cephalus is discussed in the context of species diversity and geographical distribution of these monogeneans on this host around the world. Presence of different species of Ligophorus on M. cephalus sensu stricto from the Atlantic and Mediterranean coast of Morocco demonstrates the usefulness of these species as fine resolution markers of genetic populations of their host, which are known to inhabit those coasts.

Contributions to the taxonomy of the mugilid genus Chelon Artedi (Teleostei: Mugilidae), with a major review of the status of C. persicus Senou, Randall & Okiyama, 1995.

New data based on multigene phylogenetic analyses using the COI, 16S, and cytb genes and subsequent molecular species delimitation revealed that the mugilid genus Chelon contains 11 species. Of these, two species, Chelon sp. A and Chelon sp. B, remained unidentified in previous studies. While Chelon sp. B seemingly is a close relative to C. dumerili (Steindachner, 1870), no more detailed information were provided for Chelon sp. A which is present along the east coast of South Africa. Genetic analyses performed in this study, revealed that specimens from the Arabian/Persian Gulf and the Red Sea are nested within one genetic lineage alongside Chelon sp. A. Morphological, morphometric, and meristic analyses of specimens from all three areas demonstrated that they belong to Chelon persicus Senou, Randall & Okyiama, 1995. Chelon persicus was originally described in the genus Chelon but was later placed either in the genus Liza or Planiliza. However, results presented herein confirm its placement in Chelon and its distribution range is extended in the Western Indian Ocean from the type locality (Arabian/Persian Gulf) west to the Red Sea and south to South Africa. Further, the uncertain validity of the recently described Chelon caeruleus Deef, 2018 from the Mediterranean Sea and a previously proposed generic character distinguishing Chelon from Planiliza and Parachelon, the shape of the paired postzygapophyses of the second vertebra, are briefly discussed.

Planiliza lauvergnii (Eydoux & Souleyet, 1850), a senior synonym of Planiliza affinis (Günther, 1861) with a re-evaluation of keeled back mullets (Mugiliformes: Mugilidae).

The taxonomic status of the keeled back mullets (Teleostei: Mugilidae) has been reinvestigated. Two nominal mugilid species having keeled backs from East Asia: Mugil lauvergnii Eydoux & Souleyet, 1850 and Mugil affinis Günther, 1861 have been re-evaluated through examination of the holotypes and fresh specimens. Comparison of morpho-meristic characters of the holotypes shows that both species are identical. Phylogenetic analysis based on mitochondrial cytochrome c oxidase 1 (CO1) confirmed morphological data by highlighting presence of a single clade from East Asia. Mugil lauvergnii (=Planiliza lauvergnii) is thus the sole keeled back mullet from East Asia and a senior synonym of Mugil affinis (=Planiliza affinis). The taxonomic status of two other keeled back mullets, Planiliza carinata and P. klunzingeri, is also contentious due to their similar morphology. Meristic and morphometric variation as well as sequence divergence between the two species are limited but phylogenetic analyses delineate well-supported clades consistent with biogeography and currently accepted taxonomy. Planiliza carinata and P. klunzingeri share a recent common ancestor in a Maximum Likelihood tree, with separate distribution ranges while P. lauvergnii formed a paraphyletic lineage. Based on present findings, we suggest maintenance of the taxonomic distinction of P. klunzingeri and P. carinata and discuss its evolutionary significance.

Plio-Pleistocene sea level and temperature fluctuations in the northwestern Pacific promoted speciation in the globally-distributed flathead mullet Mugil cephalus.

BACKGROUND: The study of speciation in the marine realm is challenging because of the apparent absence of physical barriers to dispersal, which are one of the main drivers of genetic diversity. Although phylogeographic studies using mitochondrial DNA (mtDNA) information often reveal significant genetic heterogeneity within marine species, the evolutionary significance of such diversity is difficult to interpret with these markers. In the northwestern (NW) Pacific, several studies have emphasised the potential importance of sea-level regression during the most recent glaciations as a driver of genetic diversity in marine species. These studies have failed, however, to determine whether the period of isolation was long enough for divergence to attain speciation. Among these marine species, the cosmopolitan estuarine-dependent fish Mugil cephalus represents an interesting case study. Several divergent allopatric mtDNA lineages have been described in this species worldwide, and three occur in sympatry in the NW Pacific. RESULTS: Ten nuclear microsatellites were surveyed to estimate the level of genetic isolation of these lineages and determine the role of sea-level fluctuation in the evolution of NW Pacific M. cephalus. Three cryptic species of M. cephalus were identified within this region (NWP1, 2 and 3) using an assignment test on the microsatellite data. Each species corresponds with one of the three mtDNA lineages in the COI phylogenetic tree. NWP3 is the most divergent species, with a distribution range that suggests tropical affinities, while NWP1, with a northward distribution from Taiwan to Russia, is a temperate species. NWP2 is distributed along the warm Kuroshio Current. The divergence of NWP1 from NWP2 dates back to the Pleistocene epoch and probably corresponds to the separation of the Japan and China Seas when sea levels dropped. Despite their subsequent range expansion since this period of glaciation, no gene flow was observed among these three lineages, indicating that speciation has been achieved. CONCLUSIONS: This study successfully identified three cryptic species in M. cephalus inhabiting the NW Pacific, using a combination of microsatellites and mitochondrial genetic markers. The current genetic architecture of the M. cephalus species complex in the NW Pacific is the result of a complex interaction of contemporary processes and historical events. Sea level and temperature fluctuations during Plio-Pleistocene epochs probably played a major role in creating the marine species diversity of the NW Pacific that is found today.

Natal origin and migration pathways of Mekong catfish (Pangasius krempfi) using strontium isotopes and trace element concentrations in environmental water and otoliths.

To improve our knowledge of the migration pathway of a highly threatened fish species along the Mekong River, strontium isotope ratios (87Sr/86Sr) and 18 trace element concentrations were measured in the water and in the otoliths of an anadromous catfish, Pangasius krempfi, to infer its natal origin and potential migration pathways. Water was sampled at 18 locations along the mainstream, tributaries and distributaries of the Mekong River. To check for accuracy and precision, measurements of the 87Sr/86Sr ratios and trace element concentrations were then compared in two laboratories that use different analytical methods. Differences in trace element concentrations between locations were not significant and could not, therefore, be used to discriminate between migration pathways. However, the Mekong mainstream, tributaries and distributaries could all be discriminated using Sr isotopes. The 87Sr/86Sr profiles recorded in P. krempfi otoliths showed that there were three contingents with obligate freshwater hatching and variable spawning sites along the Mekong mainstream, from Phnom Penh (Cambodia) to Nong Khai (Thailand) or further. After hatching, the fish migrated more or less rapidly to the Mekong Delta and then settled for most of their lifetime in brackish water. Spawning habitats and migration routes may be threatened by habitat shifts and the increasing number of hydropower dams along the river, especially the contingents born above Khone Falls (Laos). The conservation of P. krempfi, as well as other migratory fish in the Mekong River, requires agreements, common actions and management by all countries along the Mekong River. This study highlighted the importance of using both Sr/Ca and 87Sr/86Sr ratios to understand life history of anadromous fishes as the 87Sr/86Sr ratio in the water was shown to be less effective than the Sr/Ca ratio in identifying movements between different saline areas.

Differential expression of the heat shock protein Hsp70 in natural populations of the tilapia, Sarotherodon melanotheron, acclimatised to a range of environmental salinities.

BACKGROUND: The relationship between environmental variation and induction of heat shock proteins (Hsps) has been much documented under experimental conditions. However, very little is known about such induction in natural populations acclimatised to prevailing environmental conditions. Furthermore, while induction of stress proteins has been well documented in response to environmental contaminants and thermal stressors, little is known about whether factors, such as extreme salinity, are also potential inductors. The black-chinned tilapia Sarotherodon melanotheron is unusual for its ability to colonise estuarine environments in West Africa that are characterised by extremely high salinities. The relationships between mRNA levels of the 70 kDa heat shock protein (Hsp70) and Na+, K+-ATPase1alpha (Naka) in the gills, environmental salinity, and a life-history trait (condition factor) were investigated in wild populations of this species sampled from three locations in the Saloum estuary, at salinities ranging from 40 to 100 psu. RESULTS: The highest Hsp70 and Naka mRNA levels, and the poorest condition factors were recorded in the most saline sampling site (100 psu). The Hsp70 and Naka mRNA were correlated amongst themselves and showed a direct positive correlation with environmental salinity, and a negative correlation with fish condition factor. Thus, the Hsp70 is constitutively overexpressed by S. melanotheron acclimatised to extreme hypersalinity. CONCLUSIONS: These results indicate that, although S. melanotheron can colonise extremely saline environments, the overexpression of Hsp70 combined with the higher Naka mRNA expression reveals that this represents a chronic stress. The induction of Hsp70 was, therefore, a biomarker of chronic hyper-osmotic stress which presumably can be linked to the impaired growth performance and precocious reproduction that have been demonstrated in the populations at the extremely saline sites.

Acanthopagrus datnia (Hamilton, 1822), a senior synonym of Acanthopagrus longispinnis (Valenciennes, 1830) (Perciformes: Sparidae).

The taxonomic status of the Bay of Bengal nominal sparid species Coius datnia Hamilton, 1822 and Acanthopagrus longispinnis (Valenciennes, 1830) are reviewed and investigated both morphologically and genetically. Because of inadequate description and no type specimen, Coius datnia has recently been considered to belong to Sparidentex, a genus without molarifom teeth. Critical examination of the original description and examination of specimens from the type locality and adjacent areas reveal that Coius datnia belongs to Acanthopagrus, a genus with an inner series of molars. Furthermore, examination of specimens previously recognized as Acanthopagrus longispinnis (Valenciennes 1830), and recent collection of fresh specimens from lower Ganges estuary in Bangladesh, show that morphological differences between Acanthopagrus datnia and A. longispinnis are minor, and they are genetically identical. The longer second anal-fin spine in A. longispinnis (>21% SL) is, in fact, a feature of some younger A. datnia. Accordingly, A. datnia is regarded as a senior synonym of A. longispinnis, and is distinguished from its congeners by the presence of 12 dorsal-fin spines (rarely 11 or 13), 3½ scale rows between the fifth dorsal-fin spine base and lateral line, pelvic and anal-fins pale yellow to yellow with black streaks present in the interradial membranes of anal-fin rays, and caudal fin grey or yellowish grey. A neotype (and neogenotype) has been designated for Acanthopagrus datnia (Hamilton, 1822).

Biodiversity inventory of the grey mullets (Actinopterygii: Mugilidae) of the Indo-Australian Archipelago through the iterative use of DNA-based species delimitation and specimen assignment methods.

DNA barcoding opens new perspectives on the way we document biodiversity. Initially proposed to circumvent the limits of morphological characters to assign unknown individuals to known species, DNA barcoding has been used in a wide array of studies where collecting species identity constitutes a crucial step. The assignment of unknowns to knowns assumes that species are already well identified and delineated, making the assignment performed reliable. Here, we used DNA-based species delimitation and specimen assignment methods iteratively to tackle the inventory of the Indo-Australian Archipelago grey mullets, a notorious case of taxonomic complexity that requires DNA-based identification methods considering that traditional morphological identifications are usually not repeatable and sequence mislabeling is common in international sequence repositories. We first revisited a DNA barcode reference library available at the global scale for Mugilidae through different DNA-based species delimitation methods to produce a robust consensus scheme of species delineation. We then used this curated library to assign unknown specimens collected throughout the Indo-Australian Archipelago to known species. A second iteration of OTU delimitation and specimen assignment was then performed. We show the benefits of using species delimitation and specimen assignment methods iteratively to improve the accuracy of specimen identification and propose a workflow to do so.

Scolopsis igcarensis Mishra, Biswas, Russell, Satpathy amp; Selvanayagam, 2013, a junior synonym of S. vosmeri (Bloch, 1792) (Perciformes: Nemipteridae).

Scolopsis igcarensis Mishra, Biswas, Russell, Satpathy Selvanayagam, 2013 was described from specimens collected from coastal waters of southern India and Sri Lanka. A comparison of recently collected specimens from Bangladesh, initially identified as S. igcarensis, with Scolopsis vosmeri (Bloch, 1792) showed morphological differences between the two species are minor, and that specimens of S. igcarensis in fact represent juvenile and subadult colour forms of S. vosmeri. Underwater and aquarium observations, as well as molecular data based on the COI barcode region, support this conclusion. Accordingly, S. igcarensis is regarded as a junior synonym of S. vosmeri, which is redescribed herein. Phylogenetic analysis of COI barcodes of Scolopsis specimens produced in this study, together with those available from GenBank, indicate S. vosmeri is part of a species complex which includes two additional cryptic sister species that require further taxonomic investigation.

Hordes of Phages in the Gut of the Tilapia Sarotherodon melanotheron.

Preliminary studies conducted on the human gastro-intestinal tract have revealed that enteric viral communities play a preponderant role in microbial homeostatis. However to date, such communities have never been investigated in the fish gut. Herein, we examined the main ecological traits of viruses in the digestive tract of a euryhaline fish, the tilapia Sarotherodon melanotheron. Individuals were collected at 8 different sites in Senegal covering a salinity gradient from 3 to 104‰, and showing large disparities in their organic pollutant concentrations. Results showed that the gut of S. melanotheron is home to a highly abundant viral community (0.2-10.7 × 109 viruses ml-1), distinct from the surrounding water, and essentially composed of phages of which a substantial proportion is temperate (the fraction of lysogenized cells-FLC ranging from 8.1 to 33.0%). Also, a positive and significant correlation was detected between FLC and the concentrations of polycyclic aromatic hydrocarbon in sediment, while no clear relationships were found between salinity and any of the microbial parameters considered. Finally, our data suggest that virus-bacteria interactions within the fish intestine are likely sensitive to the presence of particular xenobiotics, which may compromise the balance in the gut microbiota, and subsequently affect the health of their host.

DNA barcoding post-larvae can improve the knowledge about fish biodiversity: an example from La Reunion, SW Indian Ocean.

The aim of this study was to demonstrate that fish larvae identified using their COI sequences offer a unique opportunity for improving the knowledge of local fish richness. Fish larvae were sampled at the end of their pelagic phase using light-traps set off the West Coast of La Reunion Island, southwestern Indian Ocean, once per month from October 2014 to March 2015. Among the 5174 larvae caught, 214 morphologically different specimens were selected, 196 successfully barcoded, giving a total of 101 different Barcode Index Numbers (BINs). Among these BINs, 55 had never been recorded in La Reunion exclusive economic zone (EEZ), and 13 were new for the BOLD database. Even if the sampling effort for collecting fish post-larvae during this study was relatively low, it allowed adding at least nine new species to an updated checklist of fishes of La Reunion EEZ.

Cytochrome b barcoding, molecular systematics and geographic differentiation in rabbitfishes (Siganidae).

The fish genus Siganus (Siganidae) is widely distributed in the coastal habitats of all the tropical Indo-Pacific, with 28 nominal species recognized so far, based on general morphology and coloration patterns. A mitochondrial phylogeny of 16 Siganidae species, based on the partial nucleotide sequences of the cytochome b gene, was produced. Individual haplotypes of given nominal species generally clustered at the extremity of long branches, thus validating the current taxonomy. However, S. lineatus haplotypes formed a paraphyletic group including S. guttatus, while S. fuscescens haplotypes were apparently splitted in two groups, calling for further investigation. S. woodlandi and S. argenteus formed a monophyletic group, as expected from their close morphological relatedness, although they were separated by a substantial, 14.5-16.3% nucleotide distance. Among eight species sampled from different locations across the Indo-West Pacific, S. argenteus and S. spinus showed the lowest degree of geographic differentiation, a result that correlated well with their extended pelagic larval stage. Fixation index estimates were high in all six other species tested (S. doliatus, S. fuscescens, S. lineatus, S. puellus, S. punctatus, S. vulpinus). The cytochrome b gene fragment chosen here proved useful as a barcode in Siganidae.

Diversity and distribution of cryptic species within the Mugil cephalus species complex in Vietnam.

Mugil cephalus sensu lato is a globally distributed complex of cryptic species whose distribution range and evolutionary history remains largely unknown. In the North West (NW) Pacific three species have been identified genetically among fish described morphologically as M. cephalus. Their distribution ranges are largely parapatric and has been proposed to mirror different thermal preferences. To date, few samples have been analyzed from South China Sea, which limits inferences on the evolutionary history of the species complex. We sampled fish identified morphologically as M. cephalus along Vietnamese shores and characterized them using the sequence polymorphism of two mitochondrial genes, the cytochrome oxidase I and cytochrome b. This demonstrated that all three species described in the NW Pacific are present in both northern and southern Vietnamese waters. Although the difference in species abundance reflects those observed in the NW Pacific, no phylogeographic pattern was revealed. In addition, no population structure was observed whatever the species or the distribution range considered, which indicates a significant level of gene flow that maintains genetic homogeneity of the three species. It is also conceivable that each species experienced a recent population expansion from a single ancestral population. Finally we suggest that if the cold waters of the NW Pacific present a physiologic challenge leading to the almost parapatric distribution of the three species, then it is likely that the warm surface temperatures of the South China Sea negate this barrier.