Evolution of coral reef fish<Emphasis Type="Italic"> Thalassoma</Emphasis> spp. (Labridae). 1. Molecular phylogeny and biogeography

Wrasses in the genus Thalassoma comprise 27 recognized species that occur predominantly on coral reefs and subtropical rocky reefs worldwide. The phylogenetic relationships for 26 species were examined based on two mitochondrial genes (cytochrome b and 16S rRNA) and one nuclear intron (the first intron of the ribosomal protein S7). Two closely related species, the bird-wrasses (Gomphosus varius Lacepède, 1801 and G. caerulaeus Lacepède, 1801), were also included in the analysis. These species grouped within the genus Thalassoma. Thalassoma newtoni (Osório, 1891) from Sao Tome, which is generally synonymized with the Atlantic/Mediterranean Thalassoma pavo (Linnaeus, 1758) appears to be a valid species. Using a molecular clock, the genus was estimated to have originally diverged 8–13 million years ago, with Thalassoma ballieui (Vaillant and Sauvage, 1875) from Hawaii and Thalassoma septemfasciata Scott, 1959 from Western Australia as the ancestral species. Approximately 5–10 million years ago, a sudden burst of speciation resulted in seven clades, which were resolved with the sequence data. The terminal Tethyan event and the closing of the Isthmus of Panama were probably the major historical factors shaping the evolution of species in the genus Thalassoma. These data on the spatio-temporal pattern of speciation in the Indo-Pacific indicate that peripheral species have been generated at various times throughout the history of the genus, and that none of the widespread species are relatively young. Thus, there is no clear support for centrifugal (youngest at the periphery) versus centripetal (oldest at the periphery) modes of generation of species, two theories which have been used to account for geographic gradients in species diversity.Electronic Supplementary Material Supplementary material is available in the online version of this article at http://dx.doi.org/10.1007/s00227-003-1199-0.An erratum to this article can be found at Communicated by J.P. Grassle, New Brunswick     

Tropical fishes in a temperate sea: evolution of the wrasse <Emphasis Type="Italic">Thalassoma pavo</Emphasis> and the parrotfish <Emphasis Type="Italic">Sparisoma cretense</Emphasis> in the Mediterranean and the adjacent Macaronesian and Cape Verde Archipelagos

The northeastern Atlantic and the Mediterranean Sea share geological histories and display great faunal affinities. The majority of the Mediterranean species have Atlantic origins, with a few species with tropical affinities. These include the parrotfish Sparisoma cretense and the wrasse Thalassoma pavo that are restricted to the subtropical northeastern Atlantic, the Macaronesian archipelagos (Azores, Madeira, and Canaries) and the southern Mediterranean. The Pleistocene glaciations have been described as having different effects on the fauna of the two regions. During glacial peaks, Mediterranean waters remained warmer than those of the adjacent Atlantic. Within the eastern Atlantic, the effects of Pleistocene glaciations were differentiated. Here, we perform a comparative analysis focusing on T. pavo and S. cretense populations from the northeastern Atlantic and the Mediterranean to assess the effects of Pleistocene glaciations in these two species. Sequences from the mitochondrial control region were obtained and analyzed combining phylogeographic and demographic approaches. Gene flow between Atlantic and Mediterranean populations was shown to be very high. The Mediterranean populations of T. pavo and S. cretense showed high levels of genetic diversity, even in the eastern basin, pointing to an ancient colonization event. This suggests that both species must have been able to persist in the Mediterranean during the cold Pleistocene periods. Historical migration estimates revealed a Mediterranean towards Atlantic trend in the case of T. pavo, which may reflect the re-colonization of areas in the Atlantic by fish that survived the cold phases in relatively warmer Mediterranean refugia. Our data also showed that within the Macaronesian Archipelagos, migrations occurred from Madeira towards the Azores, for both T. pavo and S. cretense, thus supporting a post-glacial colonization of the Azores by fish that persisted in the warmer region of Madeira. Similar geographic distributions, thermal affinities, and means of dispersion for T. pavo and S. cretense resulted in a similar response to the effects of Pleistocene glaciations, as evidenced by identical phylogeographic patterns.     

Population genetic structure of the stalked barnacle <Emphasis Type="Italic">Pollicipes pollicipes</Emphasis> (Gmelin, 1789) in the northeastern Atlantic: influence of coastal currents and mesoscale hydrographic structures

Within its distribution range in the northeastern Atlantic, the stalked barnacle Pollicipes pollicipes shows a well-defined pattern of genetic variation, comprising (a) a subtropical/temperate northern assemblage, made up of populations distributed between 47°N and 28°N along the French, Iberian, North African and Canary Islands coastlines, and (b) a single isolated and highly divergent tropical population in the Cape Verde Islands (16°N), at the southernmost limit of the species’ distribution. However, within the northern assemblage several populations show a level of genetic differentiation that allows rejection of the hypothesis of genetic homogeneity. The congruence observed between genetic and hydrographic patterns suggests a crucial role of hydrodynamics, and of the dispersal of the planktonic larvae, in the determination of population structure. Along the southern European Atlantic coast, the Iberian Poleward Current and mesoscale hydrographic structures are, respectively, facilitating gene flow at the regional level and genetic differentiation at the local level. On the Atlantic coast of North Africa, the homogenizing equatorward flow of the Canary Current does not extend as far as the Cape Verde Islands. A demographic expansion is dated to the late Pleistocene, preceding the Eemian interglacial, and is oldest in the case of the long-standing Cape Verde population, sustained by a stable tropical habitat. The divergence between the Cape Verde population and the remaining populations is thus ancient, and suggests that oceanic current patterns may constitute a generalized physical barrier to the dispersal of marine organisms between Cape Verde and the rest of Macaronesia.     

Evolution of coral reef fish<Emphasis Type="Italic"> Thalassoma</Emphasis> spp. (Labridae). 1. Molecular phylogeny and biogeography

Wrasses in the genus Thalassoma comprise 27 recognized species that occur predominantly on coral reefs and subtropical rocky reefs worldwide. The phylogenetic relationships for 26 species were examined based on two mitochondrial genes (cytochrome b and 16S rRNA) and one nuclear intron (the first intron of the ribosomal protein S7). Two closely related species, the bird-wrasses (Gomphosus varius Lacepède, 1801 and G. caerulaeus Lacepède, 1801), were also included in the analysis. These species grouped within the genus Thalassoma. Thalassoma newtoni (Osório, 1891) from Sao Tome, which is generally synonymized with the Atlantic/Mediterranean Thalassoma pavo (Linnaeus, 1758) appears to be a valid species. Using a molecular clock, the genus was estimated to have originally diverged 8–13 million years ago, with Thalassoma ballieui (Vaillant and Sauvage, 1875) from Hawaii and Thalassoma septemfasciata Scott, 1959 from Western Australia as the ancestral species. Approximately 5–10 million years ago, a sudden burst of speciation resulted in seven clades, which were resolved with the sequence data. The terminal Tethyan event and the closing of the Isthmus of Panama were probably the major historical factors shaping the evolution of species in the genus Thalassoma. These data on the spatio-temporal pattern of speciation in the Indo-Pacific indicate that peripheral species have been generated at various times throughout the history of the genus, and that none of the widespread species are relatively young. Thus, there is no clear support for centrifugal (youngest at the periphery) versus centripetal (oldest at the periphery) modes of generation of species, two theories which have been used to account for geographic gradients in species diversity.     

Genetic analysis of tuna populations,<Emphasis Type="Italic"> Thunnus thynnus thynnus</Emphasis> and<Emphasis Type="Italic"> T</Emphasis>.<Emphasis Type="Italic"> alalunga</Emphasis>

The genetic population structures of Atlantic northern bluefin tuna ( Thunnus thynnus thynnus) and albacore ( T. alalunga) were examined using allozyme analysis. A total of 822 Atlantic northern bluefin tuna from 18 different samples (16 Mediterranean, 1 East Atlantic, 1 West Atlantic) and 188 albacore from 5 samples (4 Mediterranean, 1 East Atlantic) were surveyed for genetic variation in 37 loci. Polymorphism and heterozygosity reveal a moderate level of genetic variability, with only two highly polymorphic loci in both Atlantic northern bluefin tuna ( FH* and SOD- 1*) and albacore ( GPI- 3* and XDH*). The level of population differentiation found for Atlantic northern bluefin tuna and albacore fits the pattern that has generally been observed in tunas, with genetic differences on a broad rather than a more local scale. For Atlantic northern bluefin tuna, no spatial or temporal genetic heterogeneity was observed within the Mediterranean Sea or between the East Atlantic and Mediterranean, indicating the existence of a single genetic grouping on the eastern side of the Atlantic Ocean. Very limited genetic differentiation was found between West Atlantic and East Atlantic/Mediterranean northern bluefin tuna, mainly due to an inversion of SOD- 1* allele frequencies. Regarding albacore, no genetic heterogeneity was observed within the Mediterranean Sea or between Mediterranean and Azores samples, suggesting the existence of a single gene pool in this area.     

Completely disjunct mitochondrial DNA haplotype distribution without a phylogeographic break in a planktonic developing gastropod

Planktonic developing organisms are generally assumed to be good dispersers showing little genetic structuring in neutral markers. At first glance, this also applies to the planktonic developing periwinkle Tectarius striatus, an endemic gastropod from Macaronesia (i.e. Azores, Madeira, Canary Islands and Cape Verde Islands), where the only sign of genetic structuring hitherto is provided by a non-significant allozyme/RAPD heterogeneity between the Cape Verde Islands and the other archipelagos. However, partial sequences of the mitochondrial cytochrome b and cytochrome oxidase I genes now show that the Cape Verde Islands and the three other archipelagos have no haplotypes in common, whereas the latter three do share several haplotypes. Nevertheless, this highly disjunct haplotype distribution does not entail a phylogeographic break separating the haplotypes of both areas in two reciprocally monophyletic groups. This remarkable geographic and phylogenetic structuring may be explained by assuming that T. striatus colonized the Macaronesian archipelagos in periods when sea levels were lower (and/or volcanic activity was higher), so that seamounts peaked above sea level and could act as stepping-stones. Yet, after the last glacial period seamounts submerged, thus preventing further stepping-stones mediated dispersal of T. striatus between the Cape Verde Islands and the other archipelagos, while not affecting dispersal among the latter because of their closer proximity and connectivity. Hence, these contrasting patterns of neutral genetic variation in T. striatus show that genetic structuring in planktonic developing species may be far more complex than is usually assumed.     

Phylogeography of two Atlantic squirrelfishes (Family Holocentridae): exploring links between pelagic larval duration and population connectivity

Genetic surveys of reef fishes have revealed high population connectivity within ocean basins, consistent with the assumption that pelagic larvae disperse long distances by oceanic currents. However, several recent studies have demonstrated that larval retention and self-recruitment may be higher than previously expected. To assess connectivity in tropical reef fishes, we contribute range-wide mtDNA surveys of two Atlantic squirrelfishes (family Holocentridae). The blackbar soldierfish, Myripristis jacobus, has a pelagic juvenile phase of about 58 days, compared to about 71 days (~22% longer) in the longjaw squirrelfish, Holocentrus ascensionis. If the pelagic duration is guiding dispersal ability, M. jacobus should have greater population genetic structure than H. ascensionis. In comparisons of mtDNA cytochrome b sequences from 69 M. jacobus (744 bp) and 101 H. ascensionis (769 bp), both species exhibited a large number of closely related haplotypes (h=0.781 and 0.974, π=0.003 and 0.006, respectively), indicating late Pleistocene coalescence of mtDNA lineages. Contrary to the prediction based on pelagic duration, M. jacobus has much less population structure (φ_ST=0.008, P=0.228) than H. ascensionis (φ_ST=0.091, P<0.001). Significant population partitions in H. ascensionis were observed between eastern, central and western Atlantic, and between Brazil and the Caribbean in the western Atlantic. These results, in combination with the findings from 13 codistributed species, indicate that pelagic larval duration is a poor predictor of population genetic structure in Atlantic reef fishes. A key to understanding this disparity may be the evolutionary depth among corresponding taxonomic groups of “reef fishes”, which extends back to the mid-Cretaceous and encompasses enormous diversity in ecology and life history. We should not expect a simple relationship between pelagic larval duration and genetic connectivity, among lineages that diverged 50–100 million years ago.     

Genetic structuring of immature loggerhead sea turtles (Caretta caretta) in the Mediterranean Sea reflects water circulation patterns

The analysis of mitochondrial DNA in loggerhead sea turtles (Caretta caretta) from eight foraging grounds in the Mediterranean and the adjoining Atlantic revealed deep genetic structuring within the western Mediterranean. As a consequence, the foraging grounds off the North-African coast and the Gimnesies Islands are shown to be inhabited mainly by turtles of the Atlantic stocks, whereas the foraging grounds off the European shore of the western Mediterranean are shown to be inhabited mainly by turtles from the eastern Mediterranean rookeries. This structuring is explained by the pattern of sea surface currents and water masses and suggests that immature loggerhead sea turtles entering the western Mediterranean from the Atlantic and the eastern Mediterranean remain linked to particular water masses, with a limited exchange of turtles between water masses. As the north of the western Mediterranean comprises mostly individuals from the highly endangered eastern Mediterranean rookeries, conservation plans should make it a priority to reduce the mortality caused by incidental by-catch in these areas.     

Genetic divergence between east and west Atlantic populations of<Emphasis Type="Italic"> Actinia</Emphasis> spp. sea anemones (Cnidaria: Actiniidae)

The sea anemone Actinia equina was considered a highly variable species with a wide geographical distribution, but molecular systematic studies have shown that this wide distribution may be the result of the lumping of cryptic species. In this work enzyme electrophoresis was used to analyse the genetic variability of A. equina from the Atlantic coasts of Europe and Africa, as well as the relationships between those populations and other species of the genus. Samples of A. equina from the United Kingdom and France were compared with supposedly conspecific populations from South Africa and a recently described species from Madeira, Actinia nigropunctata. The South African and Madeiran populations were genetically very divergent from each other (genetic identity, I=0.15), as well as from the A. equina population from the United Kingdom and from the other species studied (I between 0.23 and 0.78), indicating that the African population of A. equina belongs to a different species. High genetic similarities were found between west Atlantic A. bermudensis and east Atlantic A. sali, in spite of the great geographical distance between them, and between the European A. equina and A. prasina. The results confirm the presence of the new species A. nigropunctata from Madeira and suggest a new species within the A. equina species group, from South Africa.Communicated by O. Kinne, Oldendorf/Luhe     

Phylogeography and evolution of the triplefin <Emphasis Type="Italic">Tripterygion delaisi</Emphasis> (Pisces,Blennioidei)

The genus Tripterygion (Risso 1826) is restricted to the eastern Atlantic and the Mediterranean, and comprises only three species. T. melanuros and T. tripteronotus are essentially endemic to the Mediterranean, while Tripterygion delaisi occurs in the Atlantic and in the Mediterranean. Two subspecies of T. delaisi have been described (T. d. xanthosoma in the Mediterranean and T. d. delaisi in the Atlantic). Several scenarios have been proposed for the evolution of T. delaisi subspecies, but so far its subspeciation process is not clear. In this study we present a population survey of T. delaisi including specimens from the two recognized subspecies. We combined a phylogeographic approach with estimates of the direction of migration (between the Atlantic and the Mediterranean) and of the coalescence time of the two subspecies, using polymorphic mitochondrial and nuclear genes. The results of this study clearly support the existence of two Tripterygion delaisi clades, one in the eastern Atlantic islands and another in the Atlantic coasts of Europe and in the Mediterranean. Historical migration between the islands and Western Europe plus Mediterranean was restricted, and showed a westbound trend, with a higher number of migrants going from the Western Europe plus Mediterranean into the islands. We estimated the time of coalescence of both groups of T. delaisi to be more recent than the onset of Pleistocene glaciations (1.7 Mya). Our results are consistent with previous hypothesis that consider successive dispersal events of a Tripterygion ancestor from the western African coast colonizing the Atlantic islands and the Mediterranean, promoting the evolutionary divergence between these areas.     

Spatial–temporal pattern of bluefin tuna (Thunnus thynnus L. 1758) gonad maturation across the Mediterranean Sea

Atlantic bluefin tuna (BFT; Thunnus thynnus) is a migrating species straddling the North Atlantic Ocean and Mediterranean Sea. It is assumed that this species is divided into a western and an eastern stock, which spawn in the Gulf of Mexico and the Mediterranean Sea, respectively. To learn more about the reproductive behavior of the eastern BFT stock, we tracked gonadal development in adult fish that were sampled between April and July during three consecutive years (2003–2005). Sampling campaigns were carried out using common fishing methods at selected locations within the Mediterranean Sea, namely Levantine Sea, Malta, and Balearic Islands. An additional sampling point, Barbate, was situated northwest of the Straits of Gibraltar along the Atlantic coast. Morphometric parameters such as the total body mass (M _B) and the weights of the gonads (M _G) were recorded, and the respective gonadosomatic index (GSI) values were calculated. The data collected revealed two important trends: (1) GSI values are higher in fish caught in the eastern rather than the western locations across the Mediterranean Sea, and (2) the GSI reaches maximum values between late May and early June in Levantine Sea (eastern Mediterranean Sea), and only 2 and 4 weeks later in the central (Malta) and western (Balearic Islands) locations, respectively. The advanced gonadal development in BFT correlates well with higher sea surface temperatures. Our findings also distinguish the northern Levantine Sea BFT population (mean M _B 78.41 ± 4.13 kg), and the Barbate BFT population with the greatest M _B (all fish sampled > 100 kg). These data reflect a situation in which the eastern Mediterranean basin may function as a habitat for young BFT, until they gain a larger M _B and are able to move to the Atlantic Ocean. However, the existence of genetically discrete BFT populations in the Mediterranean Sea cannot be ruled out.     

Fine-scale distribution of juvenile Atlantic and Mediterranean loggerhead turtles (Caretta caretta) in the Mediterranean Sea

Loggerhead turtles nesting in the Mediterranean Sea exhibit remarkable genetic structuring. This paper tests the hypothesis that young loggerhead turtles from different rookeries do not distribute homogeneously among the major Mediterranean foraging grounds, due to a complex pattern of surface currents. We extracted long fragments of mitochondrial DNA from 275 stranded or bycaught juvenile turtles from six foraging grounds (Catalano-Balearic Sea, Algerian basin, Tyrrhenian Sea, Adriatic Sea, northern Ionian Sea and southern Levantine Sea). We used a Bayesian mixed-stock analysis to estimate the contributions from rookeries in the Mediterranean, the North-west Atlantic and Cape Verde to the studied foraging grounds. Differences were found in the relative contribution of juvenile turtles of Atlantic and Mediterranean origin to each foraging ground. A decreasing proportion of Atlantic juveniles was detected along the main surface current entering the Mediterranean, with a high prevalence of turtles from eastern Florida in the Algerian basin and lower numbers elsewhere. In regard to the turtles of Mediterranean origin, juveniles from Libya prevailed in central and western Mediterranean foraging grounds other than the Algerian basin. Conversely, the Adriatic Sea was characterised by a large presence of individuals from western Greece, while the southern Levantine Sea was inhabited by a heterogeneous mix of turtles from the eastern Mediterranean rookeries (Turkey, Lebanon and Israel). Overall, the distribution of juveniles may be related to surface circulation patterns in the Mediterranean and suggests that fisheries might have differential effects on each population depending on the overlap degree between foraging and fishing grounds.     

Global phylogeography of<Emphasis Type="Italic"> Cassiopea</Emphasis> (Scyphozoa: Rhizostomeae): molecular evidence for cryptic species and multiple invasions of the Hawaiian Islands

The upside-down jellyfish Cassiopea is a globally distributed, semi-sessile, planktonically dispersed scyphomedusa. Cassiopea occurs in shallow, tropical inshore marine waters on sandy mudflats and is generally associated with mangrove-dominated habitats. Controversy over the taxonomy of upside-down jellyfishes precedes their introduction to the Hawaiian Islands during the Second World War, and persists today. Here we address the global phylogeography and molecular systematics of the three currently recognized species: Cassiopea andromeda, C. frondosa, and C. xamachana. Mitochondrial cytochrome c oxidase I (COI) sequences from Australia, Bermuda, Fiji, the Florida Keys, the Hawaiian Islands, Indonesia, Palau, Panama, Papua New Guinea, and the Red Sea were analyzed. Highly divergent COI haplotypes within the putative species C. andromeda (23.4% Kimura 2-parameter molecular divergence), and shared haplotypes among populations of two separate putative species, C. andromeda and C. xamachana from different ocean basins, suggest multiple anthropogenic introductions and systematic confusion. Two deeply divergent Oahu haplotypes (20.3%) from morphologically similar, geographically separate invasive populations indicate long-term (14–40 million years ago) reproductive isolation of phylogenetically distinct source populations and cryptic species. Data support at least two independent introductions to the Hawaiian Islands, one from the Indo-Pacific, another from the western Atlantic/Red Sea. Molecular phylogenetic results support six species: (1) C. frondosa, western Atlantic (2) C. andromeda, Red Sea/western Atlantic/Hawaiian Islands (3) C. ornata, Indonesia/Palau/Fiji (4) Cassiopea sp. 1, eastern Australia (5) Cassiopea sp. 2, Papua New Guinea and (6) Cassiopea sp. 3, Papua New Guinea/Hawaiian Islands.Communicated by P.W. Sammarco, Chauvin     

Atlantic-Mediterranean and within-Mediterranean molecular variation in <Emphasis Type="Italic">Coris julis</Emphasis> (L. 1758) (Teleostei,Labridae)

Sequence variation in the mitochondrial control region was studied in the Mediterranean rainbow wrasse (Coris julis), a species with pronounced pelagic larval phase inhabiting the Mediterranean Sea and the adjacent coastal eastern Atlantic Ocean. A total of 309 specimens from 19 sampling sites were analysed with the aim of elucidating patterns of molecular variation between the Atlantic and the Mediterranean as well as within the Mediterranean Sea. Phylogeographic analyses revealed a pronounced structuring into a Mediterranean and an Atlantic group. Samples from a site at the Moroccan Mediterranean coast in the Alboran Sea showed intermediate frequencies of “Mediterranean” and “Atlantic” haplotypes. We recognised a departure from molecular neutrality and a star-like genealogy for samples from the Mediterranean Sea, which we propose to have happened due to a recent demographic expansion. The results are discussed in the light of previous studies on molecular variation in fish species between the Atlantic and the Mediterranean and within the Mediterranean.     

Phylogeographic differentiation of storm petrels (<Emphasis Type="Italic">Hydrobates pelagicus</Emphasis>) based on cytochrome<Emphasis Type="Italic"> b</Emphasis> mitochondrial DNA variation

We evaluated mitochondrial DNA (mtDNA) sequence variation in a 910 bp region of the cytochrome b gene of the storm petrel, Hydrobates pelagicus. Samples from birds collected from five populations in the North Atlantic Ocean and the Mediterranean Sea were investigated. Genetic differentiation within the Mediterranean basin was low but high in the Atlantic. Strong differences were noted between the Atlantic and the Mediterranean populations, confirming the distinction of the subspecies H. p. pelagicus and H. p. melitensis for the Atlantic and Mediterranean seabirds, respectively. Divergence between the two subspecies probably resulted from paleogeographic changes in the Strait of Gibraltar, which was likely the route used by H. pelagicus to invade the Mediterranean Sea. Current and past demography and ecology of the storm petrel is regarded as an explanation for the level of differentiation observed within each oceanic basin. We compare the phylogeographic pattern of the storm petrel to other seabirds that breed in the same regions.Communicated by S. A. Poulet, RoscoffC.C. and L.B. contributed equally to this paper.     

Genetic structure of the flounders Platichthys flesus and P. stellatus at different geographic scales

The genetic structure of the flounders Platichthys flesus L. and P. stellatus Pallas was investigated on different spatial scales through analysis of allozyme variation at 7 to 24 polymorphic loci in samples collected from different regions (Baltic Sea, North Sea, Brittany, Portugal, western Mediterranean, Adriatic Sea, Aegean Sea and Japan) in 1984 to 1987. No geographic variation was evident within a region. Some pattern of differentiation by distance was inferred within the Atlantic, while the Mediterranean comprised three geographically isolated populations and was itself geographically isolated from the Atlantic (fixed allele differences at up to three loci were found among P. flesus populations from the Atlantic, the western Mediterranean, the Adriatic Sea, the Aegean Sea and also P. stellatus from the coast of Japan). Sea temperature during the reproductive period probably acts as a barrier to gene flow between populations. Genetic distances among European flounder populations (P. flesus) were higher than, or of the same magnitude as, the genetic distance between Pacific (P. stellatus) and European flounder populations, suggesting that P. flesus is paraphyletic and/or there is no phylogenetic basis to recognising P. stellatus as a different species. The divergence between P. flesus and P. stellatus was thus inferred to be more recent than the divergence between the present P. flesus populations from the NE Atlantic and eastern Mediterranean. The eastern Mediterranean populations are thought to originate from the colonisation of the Mediterranean by a proto-P. flesus/P. stellatus ancestor, whereas the present western Mediterranean population has undergone a more recent colonisation event by P. flesus. Patterns of mitochondrial DNA variation, established on a smaller array of P. flesus samples, were in accordance with the geographic patterns inferred from the allozyme survey. In addition, they supported the hypothesis of a two-step colonisation of the western Mediterranean. These results contribute to our understanding of the biogeography of the Mediterranean marine fauna, especially the group of boreal remnants to which P. flesus belongs. Received: 7 February 1997 / Accepted: 26 March 1997     

Phylogeography of the calanoid copepods <Emphasis Type="Italic">Calanus helgolandicus</Emphasis> and <Emphasis Type="Italic">C. euxinus</Emphasis> suggests Pleistocene divergences between Atlantic,Mediterranean, and Black Sea populations

Molecular systematic analyses of marine taxa are crucial for recording ocean biodiversity, so too are elucidation of the history of population divergence and the dynamics of speciation. In this paper we present the joined phylogeography of the calanoid copepod Calanus helgolandicus (Claus 1863) from the North East (NE) Atlantic and the Adriatic Sea and the closely related C. euxinus (Hulsemann 1991) from the Black Sea based on sequences of a mitochondrial Cytochrome Oxidase subunit I (COI) fragment. Coalescent-based Bayesian methods and minimum spanning networks are used to reconstruct the history of population divergence. Our results reveal that copepod populations from all three basins share a great number of haplotypes and demonstrate a close genetic affinity of C. euxinus with C. helgolandicus. The data do not support significant genetic structuring among samples within seas. Coalescent analyses suggest divergences between NE Atlantic, Mediterranean, and Black Sea populations dating back to the middle Pleistocene, with the NE Atlantic–Mediterranean divergence being the earliest and the Mediterranean–Black Sea divergence the most recent. These middle Pleistocene dates are much older than the estimated dates of colonisation of the Mediterranean and Black Seas based on paleoclimatic scenarios. Our results do not rule out that the assumed colonisations took place but they indicate that the populations colonising the Mediterranean and the Black Sea were already, and have since remained, diverged. The chaetognath Sagitta setosa, which has a comparable distribution pattern and feeds upon the copepods, provides a unique opportunity to compare phylogeographic patterns and distinguish among alternative hypotheses. The dates produced in this paper are in agreement with those estimated elsewhere for S. setosa. We propose that a great deal of the genetic make-up of marine planktonic populations comprises divergences that date back to long before the last glacial maximum. We consider questions on the taxonomic status of C. euxinus to remain open. However, its high genetic affinity to the C. helgolandicus calls for further investigation.     

Mitochondrial DNA lineages in the European populations of mussels (<Emphasis Type="Italic">Mytilus</Emphasis> spp.)

Marine mussels (Mytilus spp.) belong to a group of benthic species crucial to coastal ecosystems in Europe and are important for the cultivation industry. In the present study, the nuclear adhesive protein marker (Me15/16) was used for identification of Mytilus species in coastal areas, on a large geographic scale in Europe. Pure M. edulis populations were found in the White Sea and Iceland. M. edulis, M. trossulus and their hybrids were found in the Baltic Sea and the North Sea (Oosterschelde, The Netherlands). M. galloprovincialis, M. edulis and their hybrids occurred in Ireland. M. galloprovincialis populations were observed in the Sea of Azov (Black Sea), the Mediterranean and Portugal. The mitochondrial (mt) DNA coding-region ND2-COIII was studied by PCR (polymerase chain reaction) and RFLP (restriction fragment length polymorphism) assay methods. The mtDNA control region was studied by PCR. Substantial differentiation in the frequency of female haplotypes among the studied populations in Europe was observed. Despite isolation between the Mytilus taxa on a macro-geographic scale, considerable mitochondrial gene flow occurred between populations, with introgression in hybrid zones on a more local geographic scale in Europe. MtDNA of the Atlantic Iberian (Portugal) population of M. galloprovincialis was more similar to mtDNA in populations of M. galloprovincialis and M. edulis from the Atlantic coasts of the Ireland and M. edulis from the North Sea, than to M. galloprovincialis from the Mediterranean. Lower polymorphism of mtDNA in populations of the Baltic and Azov Sea mussels in comparison with other European populations was observed and can be explained by the recent history of both seas after the Pleistocene glaciation. In the M. galloprovincialis population from the Azov Sea, the presence of the male-inherited (M) genome was demonstrated for the first time by sequencing the control region and was observed at high frequency. Possible influence of mussel culture on geographic distribution of the Mytilus taxa in Europe is discussed.