Analysis of meristic and mitochondrial DNA variation in Syngnathus abaster (Teleostea: Syngnathidae) from two western Mediterranean lagoons

Syngnathus abaster is a euryhaline pipefish distributed in the Mediterranean, Black Sea, and the north-eastern Atlantic. Although its populations are characterised by high morphological plasticity, neither congruent information about the morphological differentiation of S. abaster populations from the Mediterranean lagoons is available, nor population genetic surveys have been so far performed. In this context, the aims of our study were as follow: i) to describe the variation at nine meristic characters of S. abaster from two western Mediterranean brackish-water areas: the Tunis north lagoon (Tunisia) and the Mauguio lagoon (France); ii) to analyse sequences of four mitochondrial DNA regions in order to evaluate the occurrence of genetic variation between the two areas, if any. The morphological survey revealed a subdivision into two distinct groups: the first included the Tunisian specimens, the second the French ones. Genetic analysis evidenced the occurrence of a sharp genetic structuring with high levels of genetic differentiation between Tunisian and French S. abaster populations. Results suggest that the evolutionary forces driven by the different biogeographical and ecological conditions of the two Mediterranean brackish-water areas have boosted the morphological and genetic divergence of the populations here analysed. The scarce potential of long-distance dispersal of S. abaster may have also enhanced the divergence retrieved.     

Fin whale (Balaenoptera physalus) population identity in the western Mediterranean Sea

Archival bottom‐mounted audio recorders were deployed in nine different areas of the western Mediterranean Sea, Strait of Gibraltar, and adjacent North Atlantic waters during 2006–2009 to study fin whale (Balaenoptera physalus) seasonal presence and population structure. Analysis of 29,822 recording hours revealed typical long, patterned sequences of 20 Hz notes (here called “song”), back‐beats, 135–140 Hz notes, and downsweeps. Acoustic parameters (internote interval, note duration, frequency range, center and peak frequencies) were statistically compared among songs and song notes recorded in all areas. Fin whale singers producing songs attributable to the northeastern North Atlantic subpopulation were detected crossing the Strait of Gibraltar and wintering in the southwestern Mediterranean Sea (Alboran basin), while songs attributed to the Mediterranean were detected in the northwest Mediterranean basin. These results suggest that the northeastern North Atlantic fin whale distribution extends into the southwest Mediterranean basin, and spatial and temporal overlap may exist between this subpopulation and the Mediterranean subpopulation. This new interpretation of the fin whale population structure in the western Mediterranean Sea has important ecological and conservation implications. The conventionally accepted distribution ranges of northeastern North Atlantic and Mediterranean fin whale subpopulations should be reconsidered in light of the results from this study.     

Syntaxonomic analysis of theAndropogon-rich grasslands (Hyparrhenietalia hirtae) in the western Mediterranean region

Using the Braun-Blanquet approach, a syntaxonomical revision of the orderHyparrhenietalia hirtae in the western Mediterranean region was made. Various syntaxa were studied with relevés from the Iberian Peninsula, southern France, Sardinia, Sicily and adjacent islands, Balearic Islands, Canary Islands and Morocco. One alliance (Hyparrhenion hirtae), 8 associations (one of them new:Lotononido lupinifoliae-Hyparrhenietum sinaicae) and several subassociations and variants were distinguished. A complete classification of theHyparrhenietalia hirtae in the studied area with ecological and biogeographical diagnoses is given.     

Tectonic vicariance versus Messinian dispersal in western Mediterranean ground beetles

The complex geological history of the western Mediterranean region conceals the interpretation of the evolutionary history of its current fauna, as similar distribution patterns may have very different temporal and geographic origins. Particularly intriguing are some subterranean species in islands, which origin is usually difficult to interpret as their strongly modified morphologies obscure their relationships. We studied subterranean taxa and their likely relatives of two groups of ground beetles in the western Mediterranean: the Duvalius lineage (“isotopic” Trechini) and Molopina (Pterostichini). We included specimens from the islands of Mallorca, Sardinia and Sicily, plus mainland Europe and North Africa. Phylogenetic relationships were reconstructed with a combination of mitochondrial and nuclear data, and divergence dates were estimated with Bayesian methods using the same a priori molecular evolutionary rates for the same gene fragments in the two groups. In the Duvalius lineage, the subgenus Trechopsis, including all the highly modified cave or nivicolous species, was found to be polyphyletic: the species from Mallorca was found to be of Pleistocene origin and sister to the less modified species of subgenus Duvalius from the same island, whereas the Algerian species of Trechopsis were, on the contrary, related to the Sicilian Duvalius, indicating a northern colonisation route during the late Pliocene. Molopina was divided into three main lineages: the genera Abax, Percus, and the Molops groups of genera. The basal diversification of the latter was dated within a temporal window (35–25 Ma) fully congruent with the tectonic opening of the western Mediterranean basin and included six main lineages with uncertain relationships among them: the epigean genera (a) Molops and (b) Tanythrix; and the subterranean (c) Typhlochoromus (Eastern Alps), (d) Speomolops (Sardinia), (e) Henrotius (Mallorca) and (f) a strongly supported clade including the Pyrenean genera Zariquieya, Oscadytes and Molopidius. Despite the similar distribution of some of their subterranean taxa, the two studied groups show a strongly contrasting origin and mode of diversification. While the Duvalius lineage had a recent origin, with complex colonisation patterns and widespread morphological convergence among the subterranean species, the subterranean Molopina had an ancient vicariant origin resulting from the tectonic opening of the western Mediterranean basin.     

Phenetic,biogeographical, and evolutionary study ofOrnithogalum subg.Heliocharmos (Hyacinthaceae) in the western Mediterranean basin

A macromorphological study is made on taxa of the genusOrnithogalum subg.Heliocharmos in North Africa, Spain, and France. The results obtained are consistent with data from cytogenetics, reproductive biology and strategies of reproduction. They allow the retention of two species:O. algeriense andO. umbellatum. A biogeographical and phylogenetic interpretation of the subgenus is proposed for the western Mediterranean. Theoretical views on phenetics are discussed.     

The occurrence of the Japanese species Dinosoma lophiomi Toman, 1973 (Digenea: Hemiuridae) in Lophius piscatorius (Teleostei) from the western Mediterranean

Hemiurid worms from the stomach of the angler Lophius piscatorius off Corsica are described and considered to belong to a Japanese species, Dinosoma lophiomi Toman, 1973, which was originally described from an Indo-Pacific lophiid host. The apparent disjunctive distribution of this species and apparent differences in the terminal genitalia between the European and Japanese specimens are discussed. This is the first record of Dinosoma Manter, 1934 from the Mediterranean Sea. The validity of diagnostic features of Adinosoma Manter, 1947 is commented upon, and lists of records of species of Dinosoma and digeneans from L. piscatorius in the Western Mediterranean are included.     

Parasites and the island syndrome: the colonization of the western Mediterranean islands by Heligmosomoides polygyrus (Dujardin, 1845)

Aim Populations of free‐living vertebrates on islands frequently differ from their mainland counterparts by a series of changes in morphometric, life‐history, behavioural, physiological and genetic traits, collectively referred to as the ‘island syndrome’. It is not known, however, whether the ‘island syndrome’ also affects parasitic organisms. The present study establishes the colonization pattern of the Mediterranean islands by the nematode Heligmosomoides polygyrus, a direct and specific parasite of rodent hosts of the Apodemus genus, and evaluates the effects of island colonization by this species on two components of the island syndrome: the loss of genetic diversity and the enlargement of the ecological niche. Location Heligmosomoides polygyrus was sampled on seven western Mediterranean islands ? Corsica, Crete, Elba, Majorca, Minorca, Sardinia and Sicily ? as well as in 20 continental locations covering the Mediterranean basin. Methods The mitochondrial cytochrome b gene (690 base pairs) was sequenced in 166 adult H. polygyrus individuals sampled in the 27 continental and island locations. Phylogenetic reconstructions in distance, parsimony, maximum likelihood and Bayesian posterior probabilities were carried out on the whole cytochrome b gene data set. The levels of nucleotide, haplotype and genetic divergence (Kimura two‐parameter distance estimator) diversities were estimated in each island population and in the various continental lineages. Results Phylogenetic reconstructions show that the mainland origins of H. polygyrus were continental Spain for the Balearic Islands (Majorca, Minorca), northern Italy for the Tyrrhenian Islands (Corsica, Sardinia, Elba), southern Italy for Sicily, and the Balkan region for Crete. A comparison of island H. polygyrus populations with their mainland source populations revealed two characteristic components of the island syndrome in this parasite. First, island H. polygyrus populations display a significant loss of genetic diversity, which is related (r² = 0.73) to the distance separating the island from the mainland source region. Second, H. polygyrus exhibits a niche enlargement following insularization. Indeed, H. polygyrus in Corsica is present in both A. sylvaticus and Mus musculus domesticus, while mainland H. polygyrus populations are present exclusively in Apodemus hosts. Main conclusions Our results show that H. polygyrus has undergone a loss of genetic diversity and a niche (host) enlargement following colonization of the western Mediterranean islands. To our knowledge, this study provides the first evidence for components of the ‘island syndrome’ in a parasitic nematode species.     

Molecular systematics and biogeography of the western Mediterranean stonefly genus Tyrrhenoleuctra (Insecta, Plecoptera)

The stonefly genus Tyrrhenoleuctra includes species living in western Mediterranean temporary freshwater streams, sometimes also at sea level, a very unusual habitat for most Plecoptera. Traditional morphological approaches proved unsuccessful in drawing both taxonomic and phylogenetic patterns, thus hampering discussion of biogeographical patterns for this interesting group. We aimed at: (a) assessing the taxonomic status of populations of Tyrrhenoleuctra covering the geographic range of the genus; (b) studying the phylogenetic relationships among the recognized species; and (c) describing biogeographic patterns. We used phylogenetic analyses to infer the phylogenetic history of this group of stoneflies based on a combined data set of 1666 bp including fragments of the 12S ribosomal (12S) and cytochrome oxidase I (CO‐I) mtDNA genes, with maximum likelihood and Bayesian methods. Two main clades have been identified: a Sardo‐Corsican one, including Tyrrhenoleuctra zavattarii, and an Ibero‐Maghrebian one including four lineages of unkown taxonomic rank from the Balearic Islands (Maiorca), from northern Africa (Ceuta) and southernmost Spain (Algeciras), and a complex preliminarily referred herein to T. minuta (Klapálek, 1901), which includes two lineages, one from Cordoba, and one from Sierra de Grazalema (El Cerro) and Portugal (Tellhares) respectively. Dating the nodes by fixing the split of the Ibero‐Maghrebian clade from the Sardo‐Corsican one at 29 million years ago (Mya), yielded dates referring to the major geological events in the Mediterranean basin. Estimated molecular evolutionary rates ranged from 0.02–0.09% per million years (my) in the T. zavattarii lineages, to 0.2–0.7% per MY in the Ibero‐Maghrebian clade. The phylogenetic pattern emerged from the present study is congruent with the known paleo‐history of the western Mediterranean basin, with the divergence of the two main Tyrrhenoleuctra lineages corresponding to the split of the Sardo‐Corsican microplate from the Iberian block. Vicariance events have characterized the history of this stonefly group along its entire biogeographical history. Surprisingly low evolutionary rates, previously supposed by Fochetti (1991, 1994) and Fochetti et al. (2004) based on nuclear markers (allozymes), have been herein found also in mitochondrial markers.     

Leptodiscaceans (Noctilucales, Dinophyceae) from the Pacific Ocean: First records of Petalodinium and Leptodiscus beyond the Mediterranean Sea

Records of dinoflagellates of the family Leptodiscaceae (Noctilucales) from the Kuroshio Current, Philippine, Celebes, Sulu, South China Seas and the western and central Equatorial Pacific Ocean are described. Scaphodinium mirabile was the most common leptodiscacean. Two specimens that differed from the type species of Scaphodinium were found: one specimen showed a highly bifurcate proximal extremity and another showed two dissimilar proboscides from the distal extremity. Another unidentified leptodiscacean showed an arrowhead-shaped contour with the margins folded. Six specimens of Petalodinium porcelio were found, being the first record beyond the Mediterranean-Black Seas. Six specimens were tentatively assigned to the genus Leptodiscus, being the first record beyond the western Mediterranean Sea. The folded specimens that ranged from 90 to 120 μm in diameter and with a prominent flagellum were tentatively considered to be young specimens of Leptodiscus. The abundance of the leptodiscaceans is underestimated in the world's oceans.     

Deep‐water fish assemblages in the central‐western Mediterranean (south Sardinian deep‐waters)

This paper explores structure and spatial distribution of fish assemblages in an area of the central‐western Mediterranean Sea (south Sardinian deep‐waters) at depths between 546 and 1598 m. A total of 67 species (12 chondrichthyes and 55 teleosteans) were sampled. Multivariate analysis showed a clear pattern of zonation. Three main assemblages were identified within the vertical range investigated: the first situated in the shallower area between 546 and 699 m, the second group between 720 and 1099 m, and the third between 1145 and 1598 m. Abundance values declined with increasing depth. Highest biomass values were found at depths of 720–1099 m with the presence of larger species such as Galeus melastomus, Mora moro, Trachyrhynchus scabrus and Alepocephalus rostratus. Species richness decreased with depth. The deepest bottoms of the central‐western Mediterranean Sea shelter an ichthyofauna dominated by small to medium‐sized species living in a food‐scarce environment in which some large mobile fishes are widespread.     

Larval development and spawning of Citharus linguatula (Linnaeus, 1758) in the western Mediterranean

Larval development from the early stages to metamorphosis ofthe Pleuronectiform Citharus linguatula is described and illustratedfor the first time; until now only its eggs and newly hatchedlarvae were known. The specimens used in this study were collectedon plankton samples taken from the Catalan continental shelf(western Mediterranean) on two surveys made in September andOctober 1983. The descnption of larval development includesmorphological and morphometnc characters and pigmentation. Relationshipsbetween various parts of the body and standard length duringdevelopment are also considered. Information of their seasonaland spatial distribution along the studied area is provided.     

The opecoelidae (Digenea) of sparid fishes of the western Mediterranean. II. Pycnadenoides yamaguti, 1938 and Pseudopycnadena Saad-Fares & Maillard, 1986

The genera Pycnadenoides Yamaguti, 1938 and Pseudopycnadena Saad-Fares & Maillard, 1986 are redefined and their validity discussed. Two forms, currently considered to be Pycnadenoides senegalensis Fischthal & Thomas, 1972, one from Lithognathus mormyrus and Diplodus vulgaris off Corsica (Scandola) and one from Sparus aurata off Provence and Languedoc, are described separately, as some preliminary biochemical studies suggest that they might be distinct. This species is commented upon in relation to Distomum mormyri Stossich, 1885, D. umbrinae Stossich, 1885, Pycnadenoides pagrosomi Yamaguti, 1938 and P. ghanensis Fischthal & Thomas, 1968. Pseudopycnadena fischthali Saad-Fares & Maillard, 1986 is redescribed from Diplodus vulgaris, D. annularis and D. sargus off Corsica (Scandola).

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Résumé Les auteurs redéfinissent et discutent la validité des genres Pycnadenoides Yamaguti, 1938 et Pseudopycnadena Saad-Fares & Maillard, 1986. Deux formes considérées actuellement comme Pycnadenoides senegalensis Fischthal & Thomas, 1972, l'une parasite de Lithognathus mormyrus et de Diplodus vulgaris des côtes de Corse (Scandola), l'autre parasite de Sparus aurata des côtes de Provence et du Languedoc, sont décrites séparément. Des études biochimiques préliminaires suggèrent qu'il pourrait s'agir de deux espèces différentes. Cette espèce est discutée par rapport à Distomum mormyri Stossich, 1885, D. umbrinae Stossich, 1885, Pycnadenoides pagrosomi Yamaguti, 1938 et P. ghanensis Fischthal & Thomas, 1968. Pseudopycnadena fischthali Saad-Fares & Maillard, 1986 est redécrit à partir d'exemplaires provenant de Diplodus vulgaris, D. annularis et D. sargus des côtes de Corse (Scandola).

     

On the first record of the genus Octopoteuthis (Cephalopoda: Octopoteuthidae) in the Sardinian Channel (central western Mediterranean Sea)

A specimen of the rare squid genus Octopoteuthis (Cephalopoda: Octopoteuthidae) from 1600 m in the Sardinian Channel (central western Mediterranean) is described. Meristic and morphological data are provided. The specimen was a spent female and details of the reproductive system and implanted spermatangia are reported. Genetic analysis of this specimen and comparison with those available for species in this genus were carried out. Due to issues associated with the taxonomy of the genus Octopoteuthis, it was not possible to identify the specimen to the species level. The taxonomy of the genus is discussed.     

Molecular evidence for the occurrence of Contracaecum rudolphii A (Nematoda: Anisakidae) in shag Phalacrocorax aristotelis (Linnaeus) (Aves: Phalacrocoracidae) from Sardinia (western Mediterranean Sea)

Specimens of Contracaecum rudolphii Hartwich, 1964 (Nematoda: Anisakidae) from Phalacrocorax aristotelis (Linnaeus) from the Archipelago of La Maddalena (Sardinia, western Mediterranean Sea) were characterised genetically and compared with C. rudolphii A sensu D’Amelio et al. 1990 and C. rudolphii B sensu D’Amelio et al. 1990 from Phalacrocorax carbo sinensis (Blumenbach) from north-eastern Italy, and with C. rudolphii C sensu D’Amelio et al. 2007 from Phalacrocorax auritus (Lesson) from west-central Florida, USA. The sequencing of the small subunit of the mitochondrial ribosomal RNA gene (rrnS) and by polymerase chain reaction–restriction fragment length polymorphism (PCR-RFLP) analysis of the same gene and of the internal transcribed spacers (ITS) of nuclear ribosomal DNA (rDNA) allowed the identification of all specimens of C. rudolphii from P. aristotelis as C. rudolphii A. The results confirmed that the definition of genetic markers, following the analysis of nuclear ribosomal and mitochondrial DNA, provides quick and practical diagnostic tools for the detection of the 3 sibling species of C. rudolphii. The occurrence of C. rudolphii in P. aristotelis is reported for the first time from the Mediterranean area, improving the picture of the dispersal patterns of the populations of these piscivorous birds, and confirming the existence of different and isolated populations between the North and South European waters.     

Nomenclatural remarks and typification of Dasya species (Dasyaceae,Rhodophyta) from the western Mediterranean

New data on synonymy and typification of Dasya species of the western Mediterranean are provided, including: D. baillouviana (S. G. Gmel.) Mont., D. corymbifera J. Agardh, D. hutchinsiae Harv., D. ocellata (Gratel.) Harv., D. rigescens Zanardini, D. rigidula (Kütz.) Ardiss., D. sessilis Yamada and D. punicea (Zanardini) Menegh. in Zanardini. Data on synonymy and a brief nomenclatural story are shown for each species, a lectotype and a neotype are selected respectively for D. rigidula and D. punicea and clarification of neotype designation is discussed for D. corymbifera.     

Balearic insular isolation and large continental spread framed the phylogeography of the western Mediterranean Cheirolophus intybaceus s.l. (Asteraceae)

Recent Quaternary geological and climate events have shaped the evolutionary histories of plant species in the Mediterranean basin, one of the most important hotspots of biodiversity. Genetic analyses of the western Mediterranean Cheirolophus intybaceus s.l. (Asteraceae) based on AFLP were conducted to establish the relationships between its close species and populations, to reconstruct the phylogeography of the group and to analyse potential unidirectional versus bidirectional dispersals between the Ibero‐Provençal belt and the Balearic Islands. AFLP data revealed two main genetic groups, one constituted by the Balearic populations and Garraf (NE Iberia) and the other formed by the remaining mainland populations that were further sub‐structured into two geographically separated subgroups (SE + E Iberia and NE Iberia + SW France). Genetic diversity and spatial structure analyses suggested a mid‐Pleistocene scenario for the origin of C. intybaceus in southern Iberia, followed by dispersal to the north and a single colonisation event of the Balearic archipelago from the near Dianic NE Iberian area. This hypothesis was supported by paleogeographic data, which showed the existence of terrestrial connections between the continent and the islands during the Middle–Late Pleistocene marine regressions, whereas the more recent single back‐colonisation of the mainland from Mallorca might be explained by several hypotheses, such as long‐distance dispersal mediated by migratory marine birds or sea currents.