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Mollusca: Bivalvia): Evidence of a Living Benthic Community Associatedwith a Coldseep in the Eastern Mediterranean Sea

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Mollusca: Bivalvia): Evidence of a Living Benthic Community Associatedwith a Coldseep in the Eastern Mediterranean Sea Deep-Sea Research I 49 (2002) 991–1005 Lucinoma kazani n. sp. (Mollusca: Bivalvia): evidence of a living benthic community associatedwith a coldseep in the Eastern Mediterranean Sea C. Salasa,*, J. Woodsideb a Department of Biolog!ıa Animal, Facultad de Ciencias, Universidad de Malaga,! 29071- Malaga! Spain b Centre for Marine Earth Sciences, Free University, De Boelelaan, 1085, 1081 HV Amsterdam, Netherlands Received23 August 2001; accepted19 February 2002 Abstract Lucinoma kazani, a new deep-water species of Lucinidae from the Eastern Mediterranean Basin, is described and illustrated. The material was collected in the Anaximander Mountains, between Rhodes and Cyprus, Eastern Mediterranean. The first living specimens were collected during the Dutch ANAXIPROBE project in the Kazan volcano, at a depth of 1709 m. Later, during the MEDINAUT programme, both living specimens and shells were collected from several mud volcanoes at different depths in the Anaximander Mountains. This bivalve holds symbionts in the ctenidia, as do all previously studied Lucinidae. The type of habitat of this new species is gas-saturated mud, with high levels of methane, which diffuses upwards into a low-oxygen deep-water. Therefore, we consider this as evidence of a living cold seep community in the Eastern Mediterranean Sea. r 2002 Publishedby Elsevier Science Ltd. Keywords: Bivalve; Lucinoma; Cold seep; Anaximander mountain; Eastern Mediterranean Sea 1. Introduction releasedby warming of Mediterranean bottom- waters (Pancost et al., 2000). Moreover, massive Mudvolcanoes in the MediterraneanSea eruptions at mud volcanoes could also suddenly appear to have originatedby tectonic compression release large quantities of methane, a potent resulting in the extrusion of methane-rich sedi- greenhouse gas. ments (mudbreccias) (Cita et al., 1996; Limonov Trenches andaccretionary prisms delimit the et al., 1996; Woodside and Volgin, 1996; Wood- subduction zones, which define the edge of the side et al., 1998). This mud volcanism is of African plate today. The roughly 1500 km long particular interest, because it is known that Mediterranean Ridge is an accretionary prism of methane, storedas gas hydrates in subsurface marine sediments extending from western Greece sediments here (Woodside et al., 1998), could be to southern Turkey anddelimitedto the south by a thrust front at the southern edge of the subduction *Corresponding author. zone andto the north by the Hellenic Trench. It E-mail address: [email protected] (C. Salas). formedas a result of compressional forces 0967-0637/02/$ - see front matter r 2002 Publishedby Elsevier Science Ltd. PII: S 0967-0637(02)00010-9 992 C. Salas, J. Woodside / Deep-Sea Research I 49 (2002) 991–1005 generatedby the subductionof the African plate of the Dutch ANAXIPROBE project andthe beneath the Aegean plate (McKenzie, 1972; international Training Through Research pro- McClusky et al., 2000; Cita andCamerlenghi, gramme aboardthe Russian research vessel R/V 1992; MEDINAUT/MEDINETH Shipboard Gelendzhik (Woodside et al., 1997). The principal Scientific Parties, 2000). The collision of the objective of ANAXIPROBE was to determine the African andEuropean plates, with the subduction origin andsubsequent development of the Anaxi- of the oceanic part of the African plate below mander Mountains by mapping and sampling Europe causes hydrothermal vents in the Aegean them. These seamounts (Fig. 1) lie just south of Sea (Dandoet al., 1999) andcoldseeps andmud southwestern Turkey between Rhodes and Cy- volcanoes on the crest of the Mediterranean Ridge prus, andare thought to be the southwardrifted (Cita et al., 1996). geological extension of Turkey. The sampling The limestone blocks andlenses collectively strategy included the coring and dredging of ejecta groupedunder the informal name of ‘‘ Calcari a from a number of mudvolcanoes that were Lucina’’ (usually abbreviatedCAL in the paleon- discovered in 1995, as an unexpected result of a tological literature) have been recently re-inter- programme of detailed multibeam bathymetry pretedas constructions linkedto chemosynthetic (SimradEM-12D from the French research vessel processes involving CH4 andH 2S venting/seepage L’Atalante), which was also a part of the (Terzi, 1993; Terzi et al., 1994; Taviani, 1997). ANAXIPROBE project. These megafaunal communities are dominated by Two expeditions to the Eastern Mediterranean bivalves, such as large lucinids together with Sea to study the mud volcanism were carried out mytilids and vesicomyids, which occur within the within the MEDINAUT programme in the Miocene in the Apennine chain (Taviani, 1994, Anaximander Mountains, following ANAXIP- 1996; Amadesi et al., 1997; Conti and Fontana, ROBE. The first one, on the French research ship 1997). Conti andFontana (1998) also studiedthe Nadir, in November andDecember of 1998, used different settings of Lucina limestones to distin- the submersible Nautile to examine andtake site- guish, on the basis of the new fieldandcomposi- specific samples, confirming the association of tional data, between autochthonous (primary) and faults, mudvolcanoes, andmethane release. allochthonous (secondary) lucinid bearing depos- During this cruise additional shells and valves of its. Recently such a chemosynthetic environment the new species of Lucinoma were also collectedin has been discovered in the Pliocene of the Stirone the Anaximander Mountains. The second cruise River, N. Italy (Taviani et al., 1997) andthe took place in August 1999 aboardthe Russian Quaternary of the Eastern Mediterranean ‘‘Napoli R/V Professor Logachev with the objective to take Dome’’ (Corselli andBasso, 1996). Both of them cores to greater depths than those collected by the are considered deep-water cold seep communities. Nautile, together with measurement of methane in The purpose of this paper is to describe a living the water above the mudvolcanoes (MEDI- bivalve of the family Lucinidae collected from NAUT/MEDINETH ShipboardScientific Parties, deep mud volcanoes in the Anaximander Moun- 2000). tains (Eastern Mediterranean Ridge) (Fig. 1), which holds bacteriocytes with symbiotic bacteria 2.1. Geological setting in the gills. This confirms the presence of a living benthic community associatedwith a coldseep in Mudvolcanoes in the AnaximanderMountains the Eastern Mediterranean Sea. (Fig. 1) erupt a matrix-supportedbreccia, contain- ing rock clasts pluckedfrom formations lying well beneath the seafloor, andare thus a simple way of 2. Material and methods obtaining samples from the rocks forming the seamounts. The mudvolcanoes are drivenby The first samples with living material were gas andoverpressuredliquids,but it was still collected in 1996 during a combined expedition a surprise to discover active gas vents (and C. Salas, J. Woodside / Deep-Sea Research I 49 (2002) 991–1005 993 Fig. 1. Location of survey area. clathrates) there, with their characteristic benthic bottom topography in the Eastern Anaximan- communities (Woodside et al., 1997, 1998). Kazan der Mountains. It has a diameter of over 2 km mudvolcano is the site where most of the material anda height of 40–50 m. The sample Anaxi- of Lucinoma sp. was collectedandthe only site at probe 234G was obtainedfrom a gravity core which living material has been found. Measured taken on Kazan andcontaining about 120 cm concentrations of methane in the water column of relatively recent mudbreccia (i.e. there was just above Kazan mudvolcano range from 0.1 to no oxidized mud breccia near the surface and 0.9 ml/l (J.L. Charlou, pers. comm., 1999). Accord- only a few centimetres of overlying pelagic ing to Pancost et al. (2000), indirect evidence for sediments). The mud breccia was highly methane oxidation derives from the widespread saturatedin gas (probably methane; studies occurrence of authigenic carbonate in mudbreccia of the gas are in progress) andcontained samples andabundant 13C-depleted carbonate many fragments of soft sediments and hard crusts at the sediment–water interface (d13C values rocks up to 7 cm in size. are as low as À95%). In addition, multiple cores (2) Faulted Ridge: FaultedRidge was originally collected during the Medineth cruise, in the assumedto be a fault escarpment with debris Eastern Mediterranean Ridge, including the Ka- flows near the base andgas seeps andmud zan mudvolcano (Pancost et al., 2000), indicate volcanic eruptions (‘fissure’ eruption) near that methane concentration increases with depth the top. Observations from Nautile during by as much as four orders of magnitude in the the 1998 MEDINAUT expedition confirmed upper 1 m of the mudbreccias. the presence of a fault escarpment with about Among the various sampling locations were the 500 m of vertical rock outcrop (out of a total following four of relevance to this paper: height of over 1000 m), a slope below the steepest upper part, andubiquitous gas seeps (1) Kazan Mud Volcano: Kazan mudvolcano lies on the cliff face andabove. The sample on the edge of a plateau in an area of rough Anaxiprobe 209D was taken in a dredge haul 994 C. Salas, J. Woodside / Deep-Sea Research I 49 (2002) 991–1005 from the southern slope of the ridge. 200–300 m across at the summit. Gas hydrates Although the dredge started at the bottom were sampledfrom Kula mudvolcano during of the ridge in 1854 m of water and proceeded the 1996 ANAXIPROBE expedition of R/V up slope to a depth of 1165 m, it is likely that Gelendzhik (Woodside et al., 1998) and the the benthic community sampledwas nearer 1999 MEDINAUT sampling expedition of the top than the bottom (i.e. closer to 1200 m R/V Professor Logachev, andso were shells in depth than to 1800 m) because (a) it was not andtube worms. buried deeply in the dredged mud, (b) debris (4) Amsterdam Mud Volcano: Amsterdam is the flows down slope would make it difficult for largest mud volcano studied. It is about 3 km benthic animals to survive lower on the slope, in diameter and rather flat, with a relief of and(c) most of the gas seeps andthe shells are about 20 m, except for a small actively venting on upper steps in the topography according to parasitic cone on the western side, which has a observations from Nautile during the subse- diameter of about 350 m and a height of about quent MEDINAUT expedition.
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