Chemosymbiotic bivalves from the mud volcanoes of the Gulf of Cadiz, NE Atlantic, with descriptions of new species of Solemyidae, Lucinidae and Vesicomyidae

The chemosymbiotic bivalves collected from the mud volcanoes of the Gulf of Cadiz are reviewed. Of the thirteen species closely associated with chemosynthetic settings two Solemyidae, Solemya (Petrasma) elarraichensissp. n. and Acharax gadiraesp. n., one Lucinidae, Lucinoma asapheussp. n., and one Vesicomyidae, Isorropodon megadesmussp. n. are described and compared to close relatives of their respective families. The biodiversity and distribution of the chemosymbiotic bivalves in the Gulf of Cadiz are discussed and compared to the available information from other cold seeps in the Eastern Atlantic and Mediterranean. Although there is considerable similarity at the genus level between seep/mud volcano fields in the Eastern Atlantic and Mediterranean, there is little overlap at the species level. This indicates a high degree of endemism within chemosymbiotic bivalve assemblages.     

Length-weight relationships of seven fish species from tidepools of an intertidal rocky shore in the Gulf of Cadiz,Spain (NE Atlantic)

Length-weight relationships for seven fish species caught from tidepools in an intertidal rocky shore in the Gulf of Cadiz are presented. Fish were sampled monthly (April 2008–January 2012), using hand nets (mesh size 1.5 mm). The values of b parameter remained within the expected range of 2.5–3.5 for all fish species. An ANCOVA test was used to evaluate differences in the b growth parameter between females and males. Gobius cobitis, G. incognitus and G. paganellus showed significant differences in the parameters of the length-weight relationships according to the sex.     

Organization and microanatomy of the Sclerolinum contortum trophosome (Polychaeta, Siboglinidae)

The trophosome-an organ especially evolved to accommodate symbiotic bacteria-is a key character of the polychaete family Siboglinidae. Astonishingly, the trophosomes vary in organization and origin between the different siboglinid taxa. The trophosome of the small genus Sclerolinum was nearly unknown until now. Here we investigated the trophosome of S. contortum from the Gulf of Mexico, using light and electron microscopy. We show that this organ derives from the visceral mesoderm and propose that the trophosome of the sister clade Vestimentifera and Sclerolinum is a homologous character. Like that of juvenile vestimentiferans, the trophosome of Sclerolinum trophosome is simply organized. This study reveals that the Sclerolinum trophosome exhibits two regions that differ in the organization of host tissue and the size and shape of the symbionts. We suggest that a specific cell cycle within the symbiont-housing organ is directed along the longitudinal body axis, with a region of proliferation anteriorly and a region of degradation posteriorly. Using Raman microspectroscopy we demonstrate that the endosymbionts of S. contortum from the Gulf of Mexico contain sulfur vesicles, and we argue for a chemoautotrophic sulfur-oxidizing metabolism.     

Occurrence of Apristurus species in the Galicia Bank Seamount (NE Atlantic)

The aim of this study was to identify some of the Apristurus species by combining morphometric and genetic tools. Several specimens of the genus Apristurus were caught on the Galicia Bank Seamount (NE Atlantic), between 1460 and 1809 m depths, during a multidisciplinary survey carried out in 2011 within the framework of the INDEMARES Project. Morphometric and genetic analyses were conducted to aid the identification of the specimens collected. A total of 20 specimens were identified, of which 18 corresponded to Apristurus aphyodes (Nakaya and Stehmann, 1998 ), one to A. profundorum (Goode and Bean, 1896 ) and one to A. melanoasper Iglesias, Nakaya & Stehmann, 2004 . Genetic results based on the mtDNA COI sequences (682–690 bp fragment of the COI gene) support the identification of A. profundorum and A. melanoasper, with a bootstrap of 99 and 96%, respectively. The identification of A. aphyodes was also performed using a 499 bp fragment of the 16S mitochondrial gene. These are the first records of the Apristurus species from Galician waters, which extends their known area of distribution and provides more information on different biological and ecological aspects of this complex taxonomic group.     

Distribution and diversity of the Syllidae (Annelida: Polychaeta) from the Mexican Gulf of Mexico and Caribbean

The objective of this study was to contribute to the knowledge of the syllids distributed in the eastern coasts of Mexico (Gulf of Mexico and Caribbean) taking into consideration their composition, distribution and diversity. Sampling was done at 77 stations including 49 in soft bottoms (terrigenous and carbonate sediments), and 28 in hard bottoms (coralline substrates) sampled in nine reef zones included in three reef systems (Veracruz, Campeche and Cozumel). In all 2084 individuals were collected (13.5% in soft sediments and 86.4% in coralline areas). Forty-five species were identified: 31 in soft bottoms and 38 in corals, with 21 species common to both types of environments. In soft bottoms, the highest species richness and diversity were found in the Carbonate Zone, while syllids were uncommon in the Terrigenous Zone. In hard bottoms, the highest density, species richness and diversity values were found in the Veracruz Reef System, followed by the Campeche Bank and Cozumel Island Reef Systems. The dominant species in soft bottoms were Haplosyllis spongicola, Exogone lourei, Typosyllis cornuta and T. papillosus, whereas T. alosae, T. corallicola, T. cornuta and H. spongicola dominated in hard bottoms. The importance and success of the syllids in both soft bottoms and reef environments, particularly in those of the Gulf of Mexico is emphasized.     

Digenean species diversity in teleost fishes from the Gulf of Gabes, Tunisia (Western Mediterranean)

This study is the first attempt to survey the diversity of fish digeneans in the Gulf of Gabes (southern coast of Tunisia). A total of 779 fishes belonging to 32 species were sampled. 53 species of Digenea belonging to 15 families were recorded. Among these species, 24 are reported for the first time from the coast of Tunisia. We report one new host record, Lecithochirium sp. from Sardinella aurita. The Hemiuridae is the dominant family. A host-parasite list is presented with the information on the prevalence, abundance and mean intensity of each species collected. The diversity of Digenea is compared with other localities in the Mediterranean Sea and the northern east of Tunisia. The Gulf of Gabes shows the lowest diversity linked to the anthropogenic activities and impact of exotic species. The use of Digenea as indicators of the state of the ecosystem is discussed.     

Surveillance of Viruses in Wild Fish Populations in Areas around the Gulf of Cadiz (South Atlantic Iberian Peninsula)

This report describes a viral epidemiological study of wild fish around the Gulf of Cadiz (southwestern Iberian Peninsula) and is focused on infectious pancreatic necrosis virus (IPNV), viral hemorrhagic septicemia virus (VHSV), and viral nervous necrosis virus (VNNV). One fish species (Chelon labrosus) was sampled inside the gulf, at the mouth of the San Pedro River. Another 29 were sampled, in three oceanographic campaigns, at sites around the Bay of Cadiz. The fish were processed individually and subjected to isolation in cell culture and molecular diagnosis. VHSV was not isolated from any species. Thirteen IPNV-type isolates were obtained from barracuda (Sphyraena sphyraena), axillary seabream (Pagellus acarne), common two-banded seabream (Diplodus vulgaris), common pandora (P. erythrinus), Senegal seabream (D. bellottii), and surmullet (Mullus surmuletus). Six VNNV isolates were obtained from axillary seabream, common pandora, black seabream (Spondyliosoma cantharus), red mullet (Mullet barbatus), Lusitanian toadfish (Halobatrachus didactylus), and tub gurnard (Chelidonichtys lucerna). In the river mouth, viruses were detected only after reamplification, obtaining prevalence percentages of IPNV and VNNV (44.4 and 63.0%, respectively) much higher than those observed in the oceanographic campaigns (25.7 and 19.6%, respectively). The opposite results were obtained in the case of VHSV after reamplification: 11.1% in the river mouth and 43.6% in the oceanic locations. Analyzing the results with respect to the proximity of the sampling sites to the coast, an anthropogenic influence on wild fish is suggested and discussed. The type of viruses and the presence of natural reassortants are also discussed.     

The metatrochophore of a deep-sea hydrothermal vent vestimentiferan (Polychaeta: Siboglinidae)

Vestimentiferans (Siboglinidae, Polychaeta) live as juveniles and adults in an obligate mutualistic association with thiotrophic bacteria. Since their development is aposymbiotic, metatrochophores of vestimentiferans from the East Pacific Rise colonizing deep-sea hydrothermal vents are infected with the specific symbiont, develop the trophosome, and reduce their digestive system. To gain insight into the anatomy and ultrastructure and to compare this stage with metatrochophores from other siboglinids, we serial sectioned and reconstructed three specimens using light and transmission electron microscopy. The metatrochophore was composed of a prostomium, a small peristomium, two chaetigers (or two chaetigers and one additional segment without chaetae), and a minute pygidium. A digestive system and an intraepidermal nervous system were developed. Larval organs such as the prototroch, the neurotroch, and an apical organ were present, along with juvenile/adult organs such as tentacles, uncini, pyriform glands, and the anlage of the nephridial organ. We propose that in vestimentiferans, the vestimentum is the head arising from the prostomium, peristomium, and the anterior part of the first chaetiger. In frenulates, in contrast, the head is composed on the one hand of the cephalic lobe arising from the prostomium and on the other of the forepart developing from the peristomium and the anterior part of the first chaetiger. In frenulates the muscular septum between the forepart and trunk develops later than the first two chaetigers. Since this septum has no counterpart in vestimentiferans, the forepart-trunk border of frenulates is not considered homologous with the vestimentum-trunk border in vestimentiferans. The obturacular region in vestimentiferans does not appear to be a body region but rather the head appendages arising from the first chaetiger. In contrast, the tentacles in frenulates are prostomial head appendages. In both taxa, the trunk is the posterior part of the first chaetiger, and the opisthosoma is the following chaetigers and the pygidium. Comparisons with other polychaetes suggest that two larval segments are autapomorphic for the monophyletic Siboglinidae.     

Late glacial to recent deep-sea benthic foraminifera from the northeastern Atlantic (Cadiz Gulf) and western Mediterranean (Alboran Sea): Paleooceanographic results

Deep-sea benthic foraminifera of the region of water exchange between the Mediterranean and the Atlantic Ocean, at the Gibraltar Strait, show significant water-mass relationships for the past 18,000 years.Cores used in this benthic foraminifera analysis were selected, according to the context of the distribution of various deep bottom water masses, on each side of the Gibraltar Strait:

• 1.(1) on the Atlantic side of the Strait, two cores (KS 8228, KS 8229) are located in the present day NADW, and one (KC 8221) in the upper part of the present Mediterranean outflow water,
• 2.(2) on the Mediterranean side of Gibraltar, two cores (KC 8241, SU 8107) are located in the deep water mass, and one (KS 8230) in the intermediate water mass.

In the two deep basins (1500–2800 m depth in the Gulf of Cadiz, 1200–1300 m depth in the Alboran Sea), the paleooceanographic changes appear to be in an opposite way for the past 18,000 years. The Alboran basin shows a paleooceanographic evolution from a well-oxygenated, nutrient-rich environment at about 18,000 yr B.P. to a nutrient-poor, oxygen-depleted environment from 13,000 to the present time; moreover, for the time-span synchronous with the well-known development of sapropels in the eastern Mediterranean basins between 10,000 and 7000 yr B.P., the faunal assemblage shows most unusual characteristics implying drastic environmental conditions. Conversely, in the Gulf of Cadiz, the environment pass as from a biotope occupied by an oxygen-depleted, nutrient-poor water mass at about 18,000 years B.P. to a biotope occupied by NADW since the Younger Dryas; this agrees with previous data obtained in the northeastern Atlantic Ocean (Caralp, 1987).In the epibathyal zones (550–800 m depth), on both sides of the Gibraltar Strait, paleohydrographic changes do not seem so important. According to the present and very late Holocene assemblages, which are similar on both sides of the Strait with a strong east-west flow, two other stratigraphic episodes have shown the same conditions of water exchanges: the end of the isotopic stage 2 and the Younger Dryas. Conversely, during the last glacial maximum, the Bølling-Allerød and the lower Holocene, westward water fluxes were probably lower. At no time, the hypothesis of a reversal or a stop in the east-west exchanges between Mediterranean and Atlantic Ocean may be justified.     

New Fabriciola species (Polychaeta, Sabellidae, Fabriciinae) from the eastern Atlantic, with a description of sperm and spermathecal ultrastructure

New species in the genus Fabriciola Friedrich are described. Fabriciola is defined by a single synapomorphy, the presence of non-vascularized, ventral filamentous appendages. Otherwise the genus is variable, with differences among species in chaetae, eyes, and peristomial organization. The new species, Fabriciola flammula, F. liguronis and F. parvus , reflect this heterogeneity. Fabriciola liguronis has spermathecae in the radiolar crown of females, the first record of these structures in Fabriciola , but F. flammula and F. parvus lack spermathecae. Sperm ultrastructure in all three species share apomorphies with other fabriciins, but vary in other respects. Fabriciola may be paraphyletic, but material of the type species F. spongicola (Southern, 1921) has been unobtainable so far, and detailed study of this species is required to define the genus Fabriciola adequately.     

Elasmobranch species richness,fisheries, abundance and size composition in the Azores archipelago (NE Atlantic)

ABSTRACT

Elasmobranchs are a vulnerable resource, more susceptible to overfishing than most teleosts, and their assessment is complicated due to a general lack of information about their fisheries, biology and ecology. This study aimed to analyse all fishery and survey data available for elasmobranchs caught over the past c. 25 years around the Azores (NE Atlantic) to provide a baseline information, which can be used to inform stock assessment and management strategies. Elasmobranch species covered pelagic, benthopelagic and demersal habitats, from shallow to deep-water strata in areas around the islands and seamounts. These species are taken accidentally as by-catch of three main fisheries: swordfish fishery, black scabbardfish fishery and demersal bottom longline fishery. The latter represents one of the most important fishing activities in the Azores, and frequent elasmobranch by-catches include Raja clavata, Galeorhinus galeus, Deania calcea, D. profundorum, Etmopterus pusillus and E. spinax. A slight reduction in the abundance indices of these species was observed, despite the implemented technical measures (e.g. minimum size, zero catch). Little is known about resource dynamics for the Azorean region and no analytical assessments have been conducted. This study highlights the vulnerability to overfishing of these resources and the urgent need to develop management strategies.     

Recent benthic foraminiferal assemblages in deep high-energy environments from the Gulf of Cadiz (Spain)

The benthic environment in the Gulf of Cadiz, north-eastern Atlantic, is strongly affected by the Mediterranean outflow water undercurrent (MOW) which flows northwards along the western Iberian Margin at 500–1500 m water depth. Foraminiferal census counts of living and dead assemblages from 27 surface samples ranging from 103 to 1917 m water depth, and the examination of hard substrates reveal a close correlation of the fauna with the local hydrography and sediment facies. Four different faunal groups are separated by factor analysis of the living fauna. Assemblage 1 contains typical lower slope species and dominates samples from the lower MOW core layer and in the North Atlantic deep water below. Shelf edge foraminifera are common in assemblage 2a which shows the highest proportions in samples from 103 to 272 m. Assemblage 2b is dominated by upper slope species and suspension-feeders that are frequent in the upper MOW core layer and in distal settings between 396 and 901 m. Species from assemblage 3 prefer epibenthic habitats and are recorded with high proportions exclusively in the immediate flow paths of the upper MOW between 496 and 881 m. Colonisation structures and species composition of epibenthic assemblages from the proximal facies largely differ from those in distal settings. In general, epibenthic foraminifers only use elevated substrates under the influence of near-bottom flow. Under high current velocities, epibenthic foraminifers prefer large and heavy objects. They colonise high attachment levels where a maximum yield of advected food particles can be achieved. In distal settings at lower flow velocities, the elevation height does not exceed 20 mm above the surrounding sediment surface. This level is related to a hydrologic transition layer with high concentrations of suspended particles. The comparison of microhabitat preferences and faunal structure under high and low current velocities reveal that substrate stability may be a confining environmental variable for endobenthic and shallow epibenthic foraminifers. The observations also indicate that the preferential settling height of epibenthic foraminifera is related to the highest lateral flux rates of food particles within reach from the sea floor. A dynamic selection of elevated microhabitats is only used by 7.8% of all species recognised in the Gulf of Cadiz area.     

Phylogeny of the mud worm genus Marenzelleria (Polychaeta, Spionidae) inferred from mitochondrial DNA sequences

The taxonomy of the cryptic morphospecies of the mud worm genus Marenzelleria is particularly difficult and the phylogenetic relationship within the genus is unknown. Herein we reconstructed the phylogeny of all five species of this genus using sequence data of three mitochondrial genes (16SrDNA, cytochrome b, cytochrome oxidase subunit I) from 104 specimens out of 26 populations. For the three invasive species of the genus, Marenzelleria neglecta, M. viridis and M. arctia , individuals from native populations as well as from recently invaded populations were included. Nuclear 18S rDNA sequences were used to evaluate the appropriate outgroup taxon among several spionid polychaete species. The results supported the monophyly of Marenzelleria , and Malacoceros fuliginosus was found to be a suitable outgroup for the analysis of the mitochondrial gene segments. All phylogenetic reconstructions revealed a basal position of M. arctia and M. wireni , which have primarily Arctic distribution, with M. arctia obtaining the most basal position. Together with the present-day distribution of the species, this indicates an origin of the genus in the Arctic region. The relationship of the species M. neglecta , M. viridis and M. bastropi could not be resolved sufficiently due to genealogical discordance that might reflect relatively young cladogenetic events.     

Sexual reproduction in three hermaphroditic deep-sea Caryophyllia species (Anthozoa: Scleractinia) from the NE Atlantic Ocean

The reproductive biology and gametogenesis of three species of Caryophyllia were examined using histological techniques. Caryophyllia ambrosia, Alcock 1898, C. cornuformis, Pourtales 1868, and C. sequenzae, Duncan 1873, were collected from the Porcupine Seabight and Rockall Trough in the NE Atlantic Ocean. These three ahermatypic solitary corals inhabit different depth ranges: C. cornuformis – 435–2000 m, C. sequenzae – 960–1900 m, and C. ambrosia – 1100–3000 m. All three species are hermaphroditic. Hermaphroditism in these species was found to be cyclical, with only one sex of gametes viable in any individual at any point in time, although gametes of both sexes were found together within a single mesentery. Once the viable gametes are spawned, the next sex of gametes continues to grow until mature, and so gametogenesis is a continuous cycle. Oocytes and spermacysts in all species increased in density towards the actinopharynx. Maximum fecundity for C. sequenzae was 940 oocytes per polyp, and for C. ambrosia 2900 oocytes per polyp. Fecundity could not be established for C. cornuformis. In all three species, individuals were asynchronous within populations, and production of gametes was quasi-continuous throughout the year. All species are hypothesised to have lecithotrophic larvae owing to their large oocyte sizes (C. cornuformis max – 350 μm; C. sequenzae max – 430 μm; C. ambrosia max – 700 μm). Both the average oocyte size and fecundity increased in species going down the depth gradient of the NE Atlantic.     

A cladistic analysis of Siboglinidae Caullery, 1914 (Polychaeta, Annelida): formerly the phyla Pogonophora and Vestimentifera

It has been proposed in recent years that the phyla Pogonophora and Vestimentifera are a derived clade of polychaete annelids. It has also been proposed that if this clade belongs among polychaetes, then the taxon name Pogonophora is misleading and should revert to a name first formulated for the group, Siboglinidae Caullery, 1914. This recommendation is adopted in this paper, and a cladistic study using terminals of 'generic' rank in the former Pogonophora (including Vestimentifera) is undertaken. The purpose of this is to assess which taxon names should now be used for clades within Siboglinidae, and to provide a revised taxonomy, based on phylogenetic principles. Another major aim is to assess the position of the vestimentiferan clade within Siboglinidae. The results show that Vestimentifera is the sister group to Sclerolinum, and this clade is then sister group to Frenulata, i.e. the remaining Siboglinidae. The results suggest that all taxa within Siboglinidae that are not genera or species are redundant, except for the following: Siboglinidae is defined as the first polychaete, and all its descendants, to have an gut occluded by expanded endoderm filled with chemoautotrophic bacteria, as seen in the holotype of Riftia pachyptila Jones, 1981. Monilifera can be defined based on apomorphy-based system such that it is the first siboglinid, and all its descendants, to have rings of chaetae (uncini) in the opisthosoma, as seen in the holotype of Sclerolinum magdalenae Southward, 1972. Vestimentifera can be denned as the first siboglinid and all its descendants to have a vestimentum as seen in the holotype of Riftia pachyptia. Frenulata is defined as the siboglinid, and all its descendants, to have a mid-trunk girdle, as seen in the holotype of Siboglinum weberi Caullery, 1914. The taxa of generic rank are not defined here since their monophyly was not investigated.     

Depth-dependent gene flow in Gulf of Mexico cold seep Lamellibrachia tubeworms (Annelida,Siboglinidae)

Lamellibrachia vestimentiferan tubeworms form aggregations at hydrocarbon cold seeps in the deep Gulf of Mexico (GoM), creating structures that provide living space for other fauna. In the GoM, three Lamellibrachia taxa vary in morphology and depth ranges: Lamellibrachia luymesi (300–950 m), Lamellibrachia sp. 1 (950–2,604 m), and Lamellibrachia sp. 2 (1,175–3,304 m). While Lamellibrachia sp. 2 is consistently identified as a separate species, L. luymesi and sp. 1 cannot be discriminated using barcoding markers cytochrome oxidase subunit 1 (COI) and large ribosomal subunit rDNA (16S). To determine if limited gene flow was a factor in the formation of these taxa, we employed more quickly evolving markers, including mitochondrial cytochrome B (CYTB), hemoglobin subunit B2 intron (HbB2i), and six polymorphic microsatellites; microsatellites were amplified across 45 L. luymesi and sp. 1 individuals. Additionally, we used microsatellites to ask whether populations of Lamellibrachia sp. 1 and sp. 2 show evidence of significant structure. Despite a lack of resolution seen with CYTB and HbB2i, L. luymesi and sp. 1 form genetically differentiated clusters at the cross-amplified microsatellites. Furthermore, we find no evidence for population structure for either Lamellibrachia sp. 1 or sp. 2 across the GoM.     

A new species of Lepidasthenia (Polychaeta, Polynoidae) from Canary Islands

Lepidasthenia medanensis spn, is described from subtidal sandy bottom and beds of Cymodoceanodosa on the south west coast of Tenerife, Canary Islands.     

Phylogenetic relationships among NE Atlantic Plocamionida Topsent (1927) (Porifera, Poecilosclerida): under-estimated diversity in reef ecosystems

Background

Small and cryptic sponges associated with cold-water coral reefs are particularly numerous and challenging to identify, but their ecological and biochemical importance is likely to compete with megabenthic specimens.

Methodology/Principal Findings

Here we use a combination of the standard M1M6 and I3M11 partitions of the COI fragment, partial rDNA 28S sequences and morphology to delineate small encrusting Plocamionida species. In total, 46 specimens were retrieved from seven shallow to deep-water coral locations, crossing 3,000 km along the European margins. Our work provides evidence that the Plocamionida ambigua f. tylotata and f. grandichelata can be considered valid species, whereas Plocamionida ambigua f. tornata corresponds to the species P. ambigua. Within the monophyletic group of Plocamionida, P. microcionides is shown as really divergent from the other taxa, and four putative new Plocamionida species are suggested.

Conclusions/Significance

This study shows that the use of molecular and morphological information in an integrative approach is a powerful tool for the identification of sponge species, and suggests that an under-estimated biodiversity of sponges occurs in cold-water coral reefs.