Genetic diversity and population structure of three dominant myctophid fishes (<Emphasis Type="Italic">Diaphus theta,Stenobrachius leucopsarus</Emphasis>, and <Emphasis Type="Italic">S. nannochir</Emphasis>) in the North Pacific Ocean

We compared the genetic diversity of three dominant myctophid fishes in the North Pacific Ocean that have different diel vertical migration patterns on the basis of the nucleotide sequences of the mitochondrial gene for cytochrome b. No genetic structure was detected for each of these three species. The genetic diversity progressively increased for Diaphus theta, a diel migrant species showing clear diel vertical migration; Stenobrachius leucopsarus, a semi-diel migrant, in which only part of the population migrates vertically; S. nannochir, a non-diel migrant. All three species were suggested to have experienced a recent, sudden population expansion. Interspecific differences in genetic diversity might be attributable to differences in the degree of population size reduction during the glacial periods; this degree in turn corresponds to the energy demand of the fishes.     

Unravelling population structure of black musselcracker Cymatoceps nasutus: evidence for multiple populations in South African coastal waters

Genetic studies on South African marine fishes have shown that many species exist as single, well-mixed stocks throughout their core distribution. The black musselcracker or poenskop Cymatoceps nasutus is a slow-growing, late-maturing and long-lived sparid (Perciformes: Sparidae) that is endemic to South African coastal waters. Conventional tagging studies suggest that it is resident, with limited connectivity between different regions along the coast. Mitochondrial and nuclear DNA were used to investigate the genetic structure of C. nasutus along the South African coastline and showed evidence of at least two populations. Samples collected from the Tsitsikamma Marine Protected Area on the South Coast to KwaZulu-Natal on the East Coast showed no significant population structure over much of the distribution. Gene flow for C. nasutus within this region is not limited by geographic distance and appears to be unaffected by various oceanographic barriers and biogeographical boundaries. Samples from the Western Cape to the west of Tsitsikamma, however, showed moderate to substantial differentiation for both markers, which may be influenced by a temperature barrier – a coldwater ridge – along the South Coast. These data provide the first genetic assessment of this South African sparid, and suggest that a two-stock management strategy would be appropriate for the species.     

Genetic diversity and population connectivity of the sea urchin Tripneustes gratilla along the South African coast

The collector sea urchin Tripneustes gratilla has been identified as a species with potential for aquaculture production in South Africa. The species’ roe is considered a culinary delicacy in Asia and Europe. However, T. gratilla remains genetically uncharacterised in South Africa. Therefore, the purpose of this study was to provide baseline genetic information consisting of estimates of genetic diversity and population stratification, which may aid in future sustainable use of this urchin resource. A total of 22 species-specific microsatellite markers were used for the genetic characterisation of T. gratilla samples from along the South African coast, at Haga Haga, Coffee Bay, Hibberdene, Ballito Bay and Sodwana Bay. A moderate level of genetic diversity was observed, with an average number of alleles of 7.89 and an average effective number of alleles of 6.57, as well as an average observed heterozygosity of 0.55. Population differentiation tests suggest that the geographically representative samples form part of a single, large interbreeding population, with a global F_st estimate of 0.02 (p > 0.05). This finding is likely explained by high levels of gene flow between these locations caused by extensive larval dispersal during the planktonic larval stage. The panmixia observed within these natural populations of T. gratilla indicate that they could be managed as a single genetic stock.     

Mitochondrial DNA variation reveals distinct lineages in Penaeus semisulcatus (Decapoda,Penaeidae) from the Indo‐West Pacific Ocean

Penaeus semisulcatus, the green tiger prawn, is an ecologically and economically important penaeid shrimp in the Indo‐West Pacific region, especially in rice‐shrimp farming and capture fisheries in Bangladesh and Sri Lanka, respectively. Genetic variation and phylogeography of samples of this species from Bangladesh and Sri Lanka were studied utilizing different mitochondrial DNA markers, i.e. cytochrome oxidase subunit 1 (CO1), control region (CR) and 16S rRNA genes. No evidence of population structure was observed in Bangladesh, but distinct variations were found among the Sri Lankan samples (Φ_ST = 0.04, p = .002; F_ST = 0.07, p = .001), with the western sample differing from the northwestern and southern samples. The Bangladesh population had lower genetic diversity than two of the three Sri Lanka populations. The phylogeography of P. semisulcatus revealed two distinct mitochondrial DNA (mtDNA) lineages, one in the Western Pacific Ocean and second in the Indian Ocean. The Bangladesh samples showed highest levels of similarity with samples from Sri Lanka, India and Malaysia, with the Bangladesh and Sri Lanka populations sharing the most common recent ancestry. Among the Indian Ocean samples, high levels of variation were observed in the samples from Iran, indicating admixture of two distinct mtDNA lineages, one shared by the populations from the Bay of Bengal and the other possibly originating from Eastern Africa. The genetic and phylogeographic information obtained in this study will be useful in appropriate planning for management and conservation of shrimp fisheries in Bangladesh and Sri Lanka, and in the Indo‐West Pacific region.     

Phylogeny of the Sepia officinalis species complex in the eastern Atlantic extends the known distribution of Sepia vermiculata across the Benguela upwelling region

Accurate species identification and biogeographic characterisation are fundamental for appropriate management of expanding cephalopod fisheries. This study addresses this topic within the common cuttlefish Sepia officinalis species complex (S. officinalis, S. hierredda and S. vermiculata), with an emphasis on occurrence in African waters. Tissue samples from the currently presumed distributions of S. vermiculata and S. hierredda (from South Africa and Ghana/Angola, respectively) were sequenced for the cytochrome c oxidase subunit I (COI) and the cytochrome b (cytb) genes of the mitochondrial genome and then compared to existing S. officinalis sequences. Three highly divergent and reciprocally monophyletic clades, corresponding to S. officinalis, S. hierredda and S. vermiculata, were resolved, representing the first molecular confirmation of the distinct species status of S. hierredda and S. vermiculata. The sequences also revealed that, contrary to expectations based on presently published information, all samples from southern Angola were S. vermiculata. These results indicate that the range of S. vermiculata extends beyond the currently described northern limit and that S. hierredda and S. vermiculata may be indiscriminately harvested in Angolan waters. Finer-scale patterns within S. vermiculata phylogeography also indicate that the Benguela Current System and/or other environmental factors serve to isolate northern and southern stocks.     

Evidence of deep genetic divergence between populations of an important recreational fishery species,Lichia amia L. 1758, around southern Africa

Lichia amia is an important coastal recreational fishery species with a cosmopolitan distribution in the eastern Atlantic. In southern Africa, it is distributed from southern Angola to northern KwaZulu-Natal in South Africa. A recent biological survey revealed differences between Angolan and South African individuals suggesting that they may represent separate stocks. As fishery management decisions should be based on accurate knowledge of population structure, an initial survey of the genetic substructuring of L. amia was conducted on samples collected in southern Angola and South Africa. Assessment of genetic diversity and population structure was conducted using a fragment of the mitochondrial Control Region. Obtained genetic diversity levels were within the expected range for marine teleosts (h = 0.867, π = 0.007), and two deeply divergent (southern Angolan and South African) populations were identified (average φ _ST = 0.78) across the Benguela Current system. These results suggest that Angolan and South African L. amia should be managed as two independent stocks. This work represents the first assessment of population genetic substructuring across the Benguela Current of a valuable coastal fishery resource, and has application for future fisheries management.     

Phylogeographic study on the chub mackerel (Scomber japonicus) in the Northwestern Pacific indicates the late Pleistocene population isolation

In the present study, the population genetic structure and historical demography of the chub mackerel, Scomber japonicus, in the Northwestern Pacific were examined based on the full‐length sequences of the mitochondrial control region and cytochrome b gene. A total of 320 individuals was sampled from 11 localities along the coast of China and Japan from August 2011 to May 2013. Two main clades representing Chinese and Japanese populations, respectively, were detected, suggesting population isolation of S. japonicus during the late Pleistocene era. The Chinese clade was further divided into two small clades and the distribution of haplotypes were not related to sampling locality, which may be a signature of secondary contact following past division. Analyses of molecular variance and pairwise F_ST revealed significant genetic differentiation between the Chinese and Japanese populations, but a lack of genetic structure for the populations along the coast of China. Both neutrality tests and mismatch distribution analysis suggested that the populations along the coast of China experienced population expansion during the late Pleistocene. Historical events, biological characteristics and other extrinsic forces such as ocean currents may all be associated with the current phylogeographic pattern of S. japonicus in the Northwestern Pacific.     

Introduced Pacific oysters Crassostrea gigas in South Africa: demographic change,genetic diversity and body condition

Successful aquaculture species are often chosen for their fast growth rates and fecundity, which are also characteristics of invasive species. The Pacific oyster Crassostrea gigas, which constitutes 80% of global oyster trade, has been confirmed as invasive in 17 of the 66 countries where it is cultured. The single study of its status in South Africa reported populations in six South Coast estuaries in 2001, dropping to three sites in 2003. We resurveyed these estuaries, visited others in the Eastern and Western Cape provinces, and sampled oyster tissue for molecular analyses of population structure. Pacific oysters have disappeared from Knysna and, following our collections, possibly also from the GouKou Estuary. Between 2003 and 2012, the Breede Estuary population decreased by 87%, from an estimated 184 206 to 23 760 individuals. Within this estuary, oysters 12 km upriver had denser shells and higher body condition indices than did those within 1.4 km of the river mouth, presumably reflecting higher availability of suspended organic matter. However, low salinity over most of the species’ range in the estuary probably inhibits recruitment. New populations of Pacific oysters in the Swartkops and Kaaimans estuaries urgently require monitoring and eradication. Haplotype (h) and nucleotide (π) diversities across all oyster populations sampled (h = 0.2300 [SD 0.0595], π = 0.0006 [SD 0.0007]) were lower than those of co-occurring indigenous Cape rock oysters Striostrea margaritacea from the GouKou and Breede estuaries (h = 0.9076 [SD 0.0386], π = 0.00589 [SD 0.00347]). Pacific oysters either have been introduced to South African estuaries infrequently, or have experienced genetic bottlenecks following river floods or human exploitation, or both. Populations growing outside culture infrastructure are restricted to estuaries in the Eastern and Western Cape provinces, with no evidence of occurrence in fully marine shelf environments. Given the species’ considerable socio-economic importance, estuarine and coastal surveillance coupled with aquaculture zoning are required to integrate biodiversity and food security considerations.     

Significant population genetic structuring of the holoplanktic scyphozoan Pelagia noctiluca in the Atlantic Ocean

Pelagia noctiluca is thought to have a global distribution, yet our understanding of genetic connectivity across the range of this problem animal is poor. Here, we investigate the genetic structure of populations off southern Africa using mitochondrial COI and nuclear ITS1 and ITS2 genes, and compare the results to recent work conducted in the North-East Atlantic. Analyses showed significant differentiation between the southern and northern Atlantic population groups (COI Φ _st = 0.72, ITS2 Φ _st = 0.23, p < 0.001), which suggests historical rather than contemporary gene flow. Southern African samples showed high haplotypic (h = 1) and low nucleotide (π = 0.008) diversity, similar to those from Europe. Phylogenetic analyses suggest South African samples to have diverged earlier than those from the northern Atlantic.     

南海长棘海星物种有效性探讨

南海长棘海星暴发已严重威胁到该海域珊瑚礁生态系统健康,乃至整个南海生物多样性。针对南海长棘海星拉丁学名混用、中文名不统一的现状,我们采集了中沙群岛济猛暗沙海域长棘海星样品,结合长棘海星此前物种分类和分布的研究结果,对南海长棘海星物种有效性进行探讨。结果表明：所有长棘海星序列明显分为4个类群,各类群间遗传距离范围为0.087 5~0.104 7,达到了种间差异水平。南海长棘海星与长棘海星的太平洋种聚类到一起,实为太阳长棘海星(Acanthaster solaris),与其余3个种存在明显的种间差异。太阳长棘海星中2个支系间的遗传距离为0.005 3,在COⅠ基因层面属于种内差异。南海长棘海星物种有效性的研究结果为后续开展其遗传特征与适应性机制、种群分布与扩散机制、种群暴发机制等内容的研究提供科学依据。     

Evidence for multiple paternity and confirmation of an Indo-Pacific origin of blacktip shark Carcharhinus limbatus occurring in South Africa

The blacktip shark Carcharhinus limbatus is a cosmopolitan species found in warm-temperate, subtropical and tropical waters around the world. The research here aimed to assess whether multiple paternity exists in South African C. limbatus and to confirm phylogeographic patterns previously observed within the species. A minimum and maximum frequency of 50% and 71% multiple paternity, respectively, were observed in 14 litters genotyped with five microsatellite markers. Based on the mitochondrial control region, relatively high nucleotide and haplotype diversity characterised the South African sampling population, and pairwise φ_ST values indicated that it significantly differed from the populations of the Pacific and the western Atlantic oceans. The haplotype network showed that the South African samples were grouped closely with the Australian, Indo-Pacific and West African C. limbatus samples, which is suggestive of an Indo-Pacific origin for this population. This study is the first to report multiple paternity in this species. Furthermore, the results reveal that C. limbatus from South Africa is genetically diverse and phylogeographically distinct from most other C. limbatus populations.     

Stock structure and demographic history of the Indo-West Pacific mud crab Scylla serrata

The increasing exploitation of mangrove forests, without any sustainable planning, has been seriously compromising the survival of this ecosystem and of its exclusive resources. Scylla serrata is one of the most commercially exploited crabs inhabiting mangroves and estuaries of the Indo-Pacific region. This species is extensively harvested, mainly for selling to the tourist market, and, as a consequence, its populations are in constant decline. The aim of the present study was to assess the level of genetic exchange of S. serrata within the Western Indian Ocean (WIO), using a population genetic structure approach. To achieve this goal, we reconstructed the intra-specific geographic pattern of genetic variation by partial sequencing the mitochondrial DNA cytochrome oxidase subunit I, in samples from seven mangrove sites of the WIO. Our data set then encompassed all the sequences for the same genetic marker deposited in Genbank and corresponding to samples from South East Asia, Australia and some Pacific Islands: this allowed us to estimate the level of connectivity among S. serrata populations within its distribution area.     

Phylogeography of the rocky intertidal periwinkle Echinolittorina paytensis through a biogeographic transition zone in the Southeastern Pacific

Barriers to dispersal are recognized to play an important role in the differentiation of populations and ultimately in speciation. In the southeast Pacific, on the northern coast of Peru, a transition zone between the Peruvian and Panamic marine biogeographic provinces exists. Here, the convergence between two contrasting surface currents could generate a barrier effect for the larval dispersal of meroplanktonic invertebrates, which could in turn generate differentiated populations or genetic lineages on both sides of the transition zone. To address this, we studied to Echinolittorina paytensis, an abundant rocky intertidal periwinkle that spans both biogeographic provinces. A total of 95 individuals from Ecuador (2°19′S) to central Peru (7°31′S), covering the Panamic Province, the Peruvian Province, and the transition zone between, were collected. The mitochondrial markers cytochrome c oxidase I and 16SrRNA were sequenced in order to investigate phylogeography and genetic structuring. In general, no genetic structuring was found across the transition zone, suggesting this biogeographic boundary would not be acting as a barrier in this species. Factors such as a high larval dispersal capability and the occurrence of El Niño–Southern Oscillation events such as El Niño are discussed.     

Toward a mechanistic understanding of larval dispersal: insights from genomic fingerprinting of the deep-sea hydrothermal vent tubeworm Riftia pachyptila

Larval dispersal is critical for the maintenance of species populations in patchy and ephemeral hydrothermal vent habitats. On fast‐spreading ridges, such as the East Pacific Rise, rates of habitat turnover are comparable to estimated lifespans of many of the inhabiting species. Traditionally, dispersal questions have been addressed with two very different approaches, larval studies and population genetics. Population genetic studies of vent‐endemic species have been informative for determining whether patterns of dispersal are suggestive of stepping stone or island models and estimating rates of gene flow (effective migrants per generation) over broad geographic ranges. However, these studies leave fundamental questions unanswered about the specific mechanisms by which larvae disperse and species maintain their populations and biogeographic ranges. With the goal of examining genetic structure and elucidating alternative larval dispersal mechanisms, we employed a genomic DNA fingerprinting technique, amplified fragment length polymorphisms (AFLPs). To assess the potential utility of AFLPs, and genetic structure of the hydrothermal vent tubeworm Riftia pachyptila, genomic ‘fingerprints’ were recovered from 29 individuals from five vent fields spanning a distance of up to c. 5000 km along the East Pacific Rise. In contrast to previous population genetic studies that found little to no genetic structure using allozymes and mitochondrial DNA, genetic analyses of 630 polymorphic AFLP loci identified distinct subclades within R. pachyptila populations. Significant levels of differentiation were observed among populations from all vent regions as well as within each region. Discrete assemblages of tubeworms separated by as little as c. 400 m within a given vent region were genetically distinguishable and cohorts (based on size‐frequency distribution) within an aggregation were found to be most closely related. These results suggest that mechanisms of larval dispersal act to retain cohort fidelity in R. pachyptila.     

Phylogeography of the mud crab (Scylla serrata) in the Indo‐West Pacific reappraised from mitochondrial molecular and oceanographic clues: transoceanic dispersal and coastal sequential colonization

The widespread mud crab, Scylla serrata, of the Indo‐West Pacific is an excellent model species to demonstrate how the colonization history of a species can be influenced by complex oceanographic conditions. Through the combination of ecological data (fossil records and paleo‐oceanographic conditions) and molecular data (coalescent simulations, network analysis, and nucleotide diversity tests), the phylogeographic history of S. serrata was re‐analyzed. Based on the analysis of mtDNA cytochrome oxidase I sequences, two major clades were identified for S. serrata, including a widespread clade (Clade I) with three disjunct geographic clusters (IA, IB and IC) and an endemic Northwest Australian clade (Clade II). Moreover, a significant phylogeographic structure corresponding to four subpopulations was revealed: Northwest Australia, West Indian Ocean, Red Sea‐South China Sea and West Pacific. A colonization history of a Northwest Australia origin for S. serrata followed by westward transmarine dispersal across the Indian Ocean for Clade I and sequential colonization from the West Indian Ocean to Red Sea‐South China Sea and West Pacific was corroborated. The Pleistocene fluctuations of paleo‐oceanographic conditions including surface circulations and physical topography in the Indo‐West Pacific might be responsible for the wide distribution, colonization history and genetic divergence of this species.     

Genetic connectivity among populations of two congeneric direct-developing whelks varies across spatial scales

We examined the phylogeographic structure of two congeneric, direct-developing, intertidal whelks endemic to New Zealand, Cominella virgata and C. maculosa. A total of 855 specimens were collected from 23 rocky shore locations and we used them to determine the level of DNA sequence variation at the mitochondrial DNA (mtDNA) cytochrome c oxidase subunit I gene. Cominella virgata and C. maculosa have relatively high levels of haplotype diversity but low levels of nucleotide diversity. Sample sites formed regional groups that were composed of genetically differentiated populations, a finding consistent with their limited dispersal abilities. Cominella virgata showed higher levels of regional differentiation than C. maculosa, and there were also species-specific differences in genetic patterns within some regions. Both species exhibited genetic homogenisation among proximate sites within groups, suggesting some dispersal does occur between areas that are not connected by contiguous rocky shore habitat.     

Phylogeny and phylogeography of the land hermit crab Coenobita purpureus (Decapoda: Anomura: Coenobitidae) in the Northwestern Pacific Region

Hermit crabs of the family Coenobitidae comprise the land hermit crabs, with 16 Coenobita species, and the coconut crab Birgus latro. They are terrestrial but spend their marine life as planktonic larvae. Some coenobitid crabs are widely distributed in the Indo‐West Pacific region, and some species occupy narrower ranges. To improve our knowledge of coenobitid crab speciation and geographical distribution patterns, we examined the phylogenetic relationship between Coenobita purpureus, which has a narrow distribution in the Northwestern Pacific, and its more widely distributed coenobitids including B. latro, Coenobita brevimanus, Coenobita cavipes, Coenobita perlatus, Coenobita rugosus and Coenobita violascens based on the mtDNA cytochrome c oxidase subunit I (COI) and 16S rDNA genes. We also assessed the phylogeography of Co. purpureus populations based on the COI gene. Our phylogenetic and phylogeographical analyses revealed that (i) Co. purpureus clustered with Co. rugosus and (ii) the Co. purpureus population in the Ogasawara Islands was genetically distinct from those in other regions, which showed genetic panmixia. It has been hypothesized that Co. purpureus evolved in the isolated landmasses of the Ryukyu region during the Pliocene, and that its population expanded and colonized the Ogasawara Islands in the late Pleistocene. Further phylogeographical studies on Coenobita species with relatively narrow distributions coupled with characterization of their phylogenetic relationships with widely distributed congeners will advance our knowledge of the speciation and geographical distribution history of coenobitid crabs.     

The invasive Asian green mussel Perna viridis in South Africa: all that is green is not viridis

The Asian green mussel Perna viridis is an invasive Indo-Pacific species recently reported from South African harbours. To verify the invasion, a phylogenetic (and morphological) analysis of green-shelled mussels (n = 39), found in six South African harbours, was conducted using the mitochondrial cytochrome c oxidase subunit I gene (COI). Estimates of genetic distances using the neighbour-joining analysis identified P. viridis only from Durban Harbour. All other green mussels were more than 3.2% divergent from P. viridis and were identified as green-shelled variants of indigenous P. perna. The only reliable morphological differences distinguishing the two species were the poorly developed mantle papillae and the wavy pallial line in P. viridis. The confirmed occurrence of P. viridis in a South African harbour suggests that there is a possible threat of the species becoming established and then spreading onto the open coast and competing with indigenous P. perna.     

Determining the correct identity of South African Marthasterias (Echinodermata: Asteroidea)

Marthasterias glacialis are found in the cool-temperate waters of the north-eastern Atlantic Ocean, in the subtropical waters of the Mediterranean Sea and along the south-western tip of Africa. The South African Marthasterias population includes two morphotypes, a smooth, spineless rarispina form and a spiny africana form, that have been described as separate species, subspecies, or forma by various authors over the past century. To test whether these two morphotypes represent separate species, and whether either, or both, are conspecific with the North-East Atlantic species, 78 Marthasterias were collected from the Cape Peninsula of South Africa. Morphological comparisons between individuals of the two forms showed no significant clustering of samples, indicating that there is no morphological separation of the two South African forms into distinct groupings. The africana and rarispina forms were also shown to be genetically indistinguishable, using a fragment of the mitochondrial cytochrome c oxidase subunit I (COI) gene and the nuclear internal transcribed spacer 1 (ITS1). In addition, the COI sequences were also compared to those from European specimens, and phylogenetic reconstruction and intra- and interspecific levels of divergence suggested that the South African specimens form a single group that is genetically distinct from the European M. glacialis. Although the allopatric distribution, high genetic divergence (more than 3% for the COI fragment) and morphological differences suggest that the South African form should be raised to species status under the name Marthasterias africana, further work must assess an independent genetic marker (nuclear) to support raising the COI clade to species level. True M. glacialis have a spine-armament pattern of a series of three or more regular spine rows down the length of each arm, whereas M. africana are either covered in many irregularly spaced spines, or have an extraordinarily bare surface with only two spine rows per arm. Marthasterias africana may also have an actinal spine simulating the presence of a third inferomarginal spine. This work tentatively resolves the taxonomic dispute, elucidates the separation and amalgamation of the two African forms and suggests a single, uniquely South African Marthasterias species that might be distinct from the north Atlantic M. glacialis, although further analyses to test reproductive isolation between the North-East Atlantic and South African forms are required.     

Genetic structure of the reef grouper Epinephelus merra in the West Indian Ocean appears congruent with biogeographic and oceanographic boundaries

The reef fauna connectivity of the West Indian Ocean (WIO) is one of the least studied globally. Here we use genetic analyses of the grouper Epinephelus merra (Bloch 1793) to determine patterns of connectivity and to identify barriers to dispersal in this WIO marine area. Phylogeographic and population‐level analyses were conducted on cytochrome b sequences and microsatellites (13 loci) from 557 individuals sampled in 15 localities distributed across the West Indian Ocean. Additional samples from the Pacific Ocean were used to benchmark the WIO population structure. The high level of divergence revealed between Indian and Pacific localities (of about 4.5% in sequences) might be the signature of the major tectonic and climatic changes operating at the Plio‐Pleistocene transition, congruently with numerous examples of Indo‐Pacific speciation. In comparison, the E. merra sequences from the Indian Ocean constitute a monophyletic clade with a low average genetic distance (d < 0.5%). However both genetic markers indicated some structure within this ocean. The main structure revealed was the isolation of the Maldives from the WIO localities (a different group signature identified by clustering analysis, great values of differentiation). Both marker types reveal further significant structure within the WIO, mainly the isolation of the Mascarene Islands (significant AMOVA and isolation‐by‐distance patterns) and some patchy structure between the northernmost localities and within the Mozambique Channel. The WIO genetic structure of E. merra appeared congruent with main biogeographic boundaries and oceanographic currents.