Large Vesicomyidae (Mollusca: Bivalvia) from Cold Seeps in the Gulf
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Deep Sea Research Part II: Topical Studies in Archimer Oceanography Archive Institutionnelle de l’Ifremer November 2009, Volume 56, Issue 23, Pages 2350-2379 http://www.ifremer.fr/docelec/ http://dx.doi.org/10.1016/j.dsr2.2009.04.016 © 2009 Elsevier Ltd All rights reserved. Large Vesicomyidae (Mollusca: Bivalvia) from cold seeps in the Gulf of ailable on the publisher Web site Guinea off the coasts of Gabon, Congo and northern Angola Rudo von Cosela, * and Karine Olub, * a Muséum National d’Histoire Naturelle, Département Systématique et Evolution, USM 602 – Taxonomie – Collections, UMS 2700 CNRS, 55, Rue Buffon, 75005 Paris, France b IFREMER Centre de Brest, Département Etude des Ecosystèmes Profonds, BP 70, 29280 Plouzane, France *: Corresponding author : R. von Cosel, email address : [email protected], K. Olu, email address : [email protected] blisher-authenticated version is av Abstract: Two new genera and three new species of large Vesicomyidae are described from cold-seep sites on pockmarks and other sulfide-rich environments in the Gulf of Guinea (tropical east Atlantic) off Gabon, Congo (Brazzaville) and northern Angola, from 500 to 4000 m depth: “Calyptogena” (s.l.) regab n. sp., Wareniconcha (n.g.) guineensis (Thiele and Jaeckel 1931), Elenaconcha guiness n.g. n. sp., and Isorropodon atalantae n. sp. For two other species already taken by the R/V Valdivia in 1898, Calyptogena valdiviae (Thiele and Jaeckel 1931) and Isorropodon striatum (Thiele and Jaeckel 1931) new localities were discovered, and the species are rediscussed. E. guiness n.g. n.sp. is also recorded from off Banc d’Arguin, Mauritania, collected by commercial fishing vessels. The vesicomyid species here treated were encountered in different depth ranges along the Gabon–Congo–Angola margin, between 500 and 4000 m depth, and it was found that, in comparison with the dredge samples taken by the Valdivia expedition off southern Cameroon and off Rio de Oro (both at 2500 m), the same species occur in other depth ranges, in some cases with a vertical difference of more than 1000 m. .That means that the species are not confined to a given depth thought being typical for them and that the characteristics of the biotope are likely to play a major role in the distribution of the vesicomyids ccepted for publication following peer review. The definitive pu associated to cold seeps or other reduced environments along the West African margin. Keywords: Vesicomyidae; Cold seeps; Gulf of Guinea; Eastern Atlantic; Systematics; New Taxa Abbreviations: BZ, BIOZAIRE cruises; CP, beam trawl; KGS, Usnel type box corer; MNHN, Muséum national d’Histoire naturelle, Paris; PL, dive of the ROV; ROV, remote-operated vehicle (Victor 6000); SMNH, Swedish Museum of Natural History, Stockholm; ZMB, Museum für Naturkunde (formerly Zoologisches Museum) der Humboldt-Universität Berlin; ZR, ZAIROV cruise; lv, left valve; rv, right valve; sh, dead-collected specimen with both valves; spm, live-collected specimen; v, single valve Please note that this is an author-produced PDF of an article a 1 1. Introduction: The deep water veneroid bivalve family Vesicomyidae is quite divers and still far from well known, especially in its supraspecific taxonomy but also on species level (Krylova & Sahling, 2006, therein a detailed account of the history of vesicomyid taxonomy). The size range goes from species of millimetric size (e.g. the type species V. atlantica E.A. Smith 1885) to very large species up to 26 cm such as Calyptogena (s.l.) magnifica Boss & Turner 1980. All Vesicomyidae are associated with hydrothermal vents, cold seeps and other reducing sediments or organic remains, they are chemosymbiotic and host sulphide- oxidizing chemoautotrophic bacteria in their gills (Boss & Turner, 1980, Fiala- Medioni & Felbeck, 1990, Fisher, 1990, Bennett et al., 1994, Cosel & Salas, 2001, Krylova, 2002, Krylova & Sahling, 2006). The other known chemosymbiotic bivalve families are Solemyidae, Manzanellidae, Mytilidae, Lucinidae, Thyrasiridae and Teredinidae. Of the almost 100 described and named species of Vesicomyidae, 13 were yet reported from the eastern Atlantic, most of them from cold seeps and reducing sediments off tropical West Africa (11 species), but some also from the eastern Mediterranean and the northeastern temperate Atlantic (Boss, 1970; Cosel & Salas, 2001). The first five species of large vesicomyids from the eastern Atlantic were taken in 1898 by the German R/V Valdivia off West Africa, three species in the Gulf of Guinea and two from off Rio de Oro (NW-Africa), they were published by Thiele & Jaeckel (1931) at a time when hot-vent and cold-seep biotopes and the special ecology of the family were still not known. Only very recently, other Vesicomyidae, this time smaller species, were discovered in the West African realm: one species off Pointe-Noire in 150-200 m and three others off the Mauritanian coast in 900- 1200 m (Cosel & Salas, 2001). During recent cruises in the course of the Ifremer/TOTAL ZAIANGO and BIOZAIRE programs on cold-seep sites on pockmarks and other deep sites more or less close to the deep Congo river channel in the Gulf of Guinea (Sibuet et al., 2002; Ondréas et al., 2005; Olu-Le Roy et al, 2007a), several unknown large Vesicomyidae were taken, as well as also the species already known from the Valdivia material, for which new localities were detected. One species of these is monographed elsewhere by the present authors (Cosel & Olu, 2008), the six remaining large vesicomyid species are treated herein, 2 among them two new genera and three new species. In this new context, the Valdivia material is rediscussed. 2. Material and methods The material was obtained during the Ifremer-TOTAL collaboration projects ZAIANGO and BIOZAIRE on cold seeps and at deep stations in the vicinity of the Congo channel off the coast of Gabon, Congo and northern Angola. It was collected during the cruises ZAIROV (P.I. H. Ondréas), BIOZAIRE 1, BIOZAIRE 2 (P.I. M. Sibuet) and BIOZAIRE 3 (P.I. A. Khripounoff) by the R/V "Atalante" with the ROV Victor 6000 and by means of an USNEL type box corer (“KGS” in sample lists) for macrofauna. During BIOZAIRE 3, a 6 m-beam trawl (“CP” in sample lists) dedicated to megafauna sampling was employed. The material was sorted on board and in the Laboratoire Environnement Profond of IFREMER, Brest. The sampling sites include different pockmarks characterized by methane-rich fluid emissions. The most known cold-seep site is a giant isolated pockmark, named REGAB, situated 8 km north of the Congo channel at a depth of 3150 m (Fig. 1). It is related to a deep buried paleo-channel identified by previous geophysical studies (Gay et al., 2003). It was investigated by the ROV Victor, giving the ground thrusting for fluid escape features (Ondréas et al., 2005), chemical characterisation of seeps (Charlou et al., 2004) and chemosynthetic communities distribution (Olu-Le Roy et al., 2007a). Large quantities of methane are released in the water column and methane hydrates were sampled in the sediment and on the seafloor. Two species of large Vesicomyidae co-occur with the mussel Bathymodiolus aff boomerang (Olu-Le Roy et al., 2007b) and the siboglinid tubeworm Escarpia southwardae Andersen, Hourdez, Marie, Jollivet, Lallier & Sibuet 2004 as large size symbiont bearing species forming habitats for macrofauna assemblages. A smaller pockmark, ASTRID, situated 80 km north to the Congo channel and about 100 km from the REGAB site at a depth of 2820 m, was explored during only one Victor dive. It was colonised by bushes of E. southwardae, and some clusters of empty vesicomyid shells were also observed. Very few samples and environmental data are available for this site. 3 The second main cold-seep site, named GUINESS, is situated along the Gabon margin between 580 and 670 m depth. It comprises several pockmarks, smaller and less active than that of the REGAB site. Victor’s dives revealed that most of them were presently inactive with carbonate precipitation and old bivalve shells, but seepages were observed on two of the pockmarks sustaining small (about 1m diameter) but dense vesicomyid clusters covered by microbial mats. Contrary to the REGAB site, where both large vesicomyid species typical for that site co-occur in the same clusters, the clusters sampled in the GUINESS area were monospecific. The two vesicomyid species occurring on GUINESS were sampled on two different pockmarks situated 2 km apart. Four pockmarks previously identified from multibeam backscatter were surveyed during a Victor dive in the MPS 1 area situated about 50 km north to the Congo canyon (fig 1) at a depth of about 1500 m. However, no living chemosynthetic communities but only empty vesicomyid shells were observed on the bottom. Several other pockmarks were surveyed and sampled by the ROV Victor at the “BSR” (Bottom Simulating Reflector) Congo site, corresponding to the area where this reflector, visible on seismic profiles and underlying the hydrate layer, reaches the seafloor, between 750 and 800 m depth. Further samples were obtained at two sites, situated close (ZD) or far (ZC) from the Congo channel. These deep stations, which were sampled using the beam trawl and the USNEL box corer, are situated along the 4000 m isobath respectively at 200 km and 15 km south of the Congo channel. Initially, they had been chosen to evaluate the influence of channel overflows on the benthic communities and not for a particular geological setting favouring methane seepages. Of the live-collected vesicomyid specimens, an important part was already dissected on board and the tissues were preserved and used for biochemistry purpose or molecular research. A part of the remaining specimens were frozen before preserving which made the tissues more fragile. The specimens with soft parts are now part of the MNHN collection (for some species only few material) and will be used for detailed anatomical work in the future.