Deep-Sea Research I 49 (2002) 991–1005

Lucinoma kazani n. sp. (Mollusca: Bivalvia): evidenceof a living benthic community associated with a cold seep

in the Eastern Mediterranean Sea

C. Salasa,*, J. Woodsideb

aDepartment of Biolog!ıa Animal, Facultad de Ciencias, Universidad de M !alaga, 29071- M !alaga SpainbCentre for Marine Earth Sciences, Free University, De Boelelaan, 1085, 1081 HV Amsterdam, Netherlands

Received 23 August 2001; accepted 19 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 1709m. 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

Published by Elsevier Science Ltd.

Keywords: Bivalve; Lucinoma; Cold seep; Anaximander mountain; Eastern Mediterranean Sea

1. Introduction

Mud volcanoes in the Mediterranean Seaappear to have originated by tectonic compressionresulting in the extrusion of methane-rich sedi-ments (mud breccias) (Cita et al., 1996; Limonovet al., 1996; Woodside and Volgin, 1996; Wood-side et al., 1998). This mud volcanism is ofparticular interest, because it is known thatmethane, stored as gas hydrates in subsurfacesediments here (Woodside et al., 1998), could be

released by warming of Mediterranean bottom-waters (Pancost et al., 2000). Moreover, massiveeruptions at mud volcanoes could also suddenlyrelease large quantities of methane, a potentgreenhouse gas.Trenches and accretionary prisms delimit the

subduction zones, which define the edge of theAfrican plate today. The roughly 1500 km longMediterranean Ridge is an accretionary prism ofmarine sediments extending from western Greeceto southern Turkey and delimited to the south by athrust front at the southern edge of the subductionzone and to the north by the Hellenic Trench. Itformed as a result of compressional forces

*Corresponding author.

E-mail address: casanova@uma.es (C. Salas).

0967-0637/02/$ - see front matter r 2002 Published by Elsevier Science Ltd.

PII: S 0 9 6 7 - 0 6 3 7 ( 0 2 ) 0 0 0 1 0 - 9

generated by the subduction of the African platebeneath the Aegean plate (McKenzie, 1972;McClusky et al., 2000; Cita and Camerlenghi,1992; MEDINAUT/MEDINETH ShipboardScientific Parties, 2000). The collision of theAfrican and European plates, with the subductionof the oceanic part of the African plate belowEurope causes hydrothermal vents in the AegeanSea (Dando et al., 1999) and cold seeps and mudvolcanoes on the crest of the Mediterranean Ridge(Cita et al., 1996).The limestone blocks and lenses collectively

grouped under the informal name of ‘‘Calcari a

Lucina’’ (usually abbreviated CAL in the paleon-tological literature) have been recently re-inter-preted as constructions linked to chemosyntheticprocesses involving CH4 and H2S venting/seepage(Terzi, 1993; Terzi et al., 1994; Taviani, 1997).These megafaunal communities are dominated bybivalves, such as large lucinids together withmytilids and vesicomyids, which occur within theMiocene in the Apennine chain (Taviani, 1994,1996; Amadesi et al., 1997; Conti and Fontana,1997). Conti and Fontana (1998) also studied thedifferent settings of Lucina limestones to distin-guish, on the basis of the new field and composi-tional data, between autochthonous (primary) andallochthonous (secondary) lucinid bearing depos-its. Recently such a chemosynthetic environmenthas been discovered in the Pliocene of the StironeRiver, N. Italy (Taviani et al., 1997) and theQuaternary of the Eastern Mediterranean ‘‘NapoliDome’’ (Corselli and Basso, 1996). Both of themare considered deep-water cold seep communities.The purpose of this paper is to describe a living

bivalve of the family Lucinidae collected fromdeep mud volcanoes in the Anaximander Moun-tains (Eastern Mediterranean Ridge) (Fig. 1),which holds bacteriocytes with symbiotic bacteriain the gills. This confirms the presence of a livingbenthic community associated with a cold seep inthe Eastern Mediterranean Sea.

2. Material and methods

The first samples with living material werecollected in 1996 during a combined expedition

of the Dutch ANAXIPROBE project and theinternational Training Through Research pro-gramme aboard the Russian research vessel R/VGelendzhik (Woodside et al., 1997). The principalobjective of ANAXIPROBE was to determine theorigin and subsequent development of the Anaxi-mander Mountains by mapping and samplingthem. These seamounts (Fig. 1) lie just south ofsouthwestern Turkey between Rhodes and Cy-prus, and are thought to be the southward riftedgeological extension of Turkey. The samplingstrategy included the coring and dredging of ejectafrom a number of mud volcanoes that werediscovered in 1995, as an unexpected result of aprogramme of detailed multibeam bathymetry(Simrad EM-12D from the French research vesselL’Atalante), which was also a part of theANAXIPROBE project.Two expeditions to the Eastern Mediterranean

Sea to study the mud volcanism were carried outwithin the MEDINAUT programme in theAnaximander Mountains, following ANAXIP-ROBE. The first one, on the French research shipNadir, in November and December of 1998, usedthe submersible Nautile to examine and take site-specific samples, confirming the association offaults, mud volcanoes, and methane release.During this cruise additional shells and valves ofthe new species of Lucinoma were also collected inthe Anaximander Mountains. The second cruisetook place in August 1999 aboard the RussianR/V Professor Logachev with the objective to takecores to greater depths than those collected by theNautile, together with measurement of methane inthe water above the mud volcanoes (MEDI-NAUT/MEDINETH Shipboard Scientific Parties,2000).

2.1. Geological setting

Mud volcanoes in the Anaximander Mountains(Fig. 1) erupt a matrix-supported breccia, contain-ing rock clasts plucked from formations lying wellbeneath the seafloor, and are thus a simple way ofobtaining samples from the rocks forming theseamounts. The mud volcanoes are driven bygas and overpressured liquids, but it was stilla surprise to discover active gas vents (and

C. Salas, J. Woodside / Deep-Sea Research I 49 (2002) 991–1005992

clathrates) there, with their characteristic benthiccommunities (Woodside et al., 1997, 1998). Kazanmud volcano is the site where most of the materialof Lucinoma sp. was collected and the only site atwhich living material has been found. Measuredconcentrations of methane in the water columnjust above Kazan mud volcano range from 0.1 to0.9 ml/l (J.L. Charlou, pers. comm., 1999). Accord-ing to Pancost et al. (2000), indirect evidence formethane oxidation derives from the widespreadoccurrence of authigenic carbonate in mud brecciasamples and abundant 13C-depleted carbonatecrusts at the sediment–water interface (d13C valuesare as low as �95%). In addition, multiple corescollected during the Medineth cruise, in theEastern Mediterranean Ridge, including the Ka-zan mud volcano (Pancost et al., 2000), indicatethat methane concentration increases with depthby as much as four orders of magnitude in theupper 1m of the mud breccias.Among the various sampling locations were the

following four of relevance to this paper:

(1) Kazan Mud Volcano: Kazan mud volcano lieson the edge of a plateau in an area of rough

bottom topography in the Eastern Anaximan-der Mountains. It has a diameter of over 2 kmand a height of 40–50m. The sample Anaxi-probe 234G was obtained from a gravity coretaken on Kazan and containing about 120 cmof relatively recent mud breccia (i.e. there wasno oxidized mud breccia near the surface andonly a few centimetres of overlying pelagicsediments). The mud breccia was highlysaturated in gas (probably methane; studiesof the gas are in progress) and containedmany fragments of soft sediments and hardrocks up to 7 cm in size.

(2) Faulted Ridge: Faulted Ridge was originallyassumed to be a fault escarpment with debrisflows near the base and gas seeps and mudvolcanic eruptions (‘fissure’ eruption) nearthe top. Observations from Nautile duringthe 1998 MEDINAUT expedition confirmedthe presence of a fault escarpment with about500m of vertical rock outcrop (out of a totalheight of over 1000m), a slope below thesteepest upper part, and ubiquitous gas seepson the cliff face and above. The sampleAnaxiprobe 209D was taken in a dredge haul

Fig. 1. Location of survey area.

C. Salas, J. Woodside / Deep-Sea Research I 49 (2002) 991–1005 993

from the southern slope of the ridge.Although the dredge started at the bottomof the ridge in 1854m of water and proceededup slope to a depth of 1165m, it is likely thatthe benthic community sampled was nearerthe top than the bottom (i.e. closer to 1200min depth than to 1800m) because (a) it was notburied deeply in the dredged mud, (b) debrisflows down slope would make it difficult forbenthic animals to survive lower on the slope,and (c) most of the gas seeps and the shells areon upper steps in the topography according toobservations from Nautile during the subse-quent MEDINAUT expedition. Moreover,along the summit of the ridge, several pock-marks and acoustic wipe-outs suggest that gasis escaping there. The ridge itself is interpretedto be fault-controlled, with gas seeps takingplace along the fault at fractures in theoutcrop or at bedding plains of the sedimen-tary rocks. Tube worms also found in dredge209D were identified as vestimentiferans ofthe genus Lamellibrachia (E. Southward, pers.comm., 1996). They were attached to a largepiece of rock with irregular holes runningthrough it, which were inferred to be gasescape structures. The rock, which incorpo-rated fragments of mollusc shells, wasshown to be a methane-derived carbonatecrust at the location of active gas venting. Theworms were attached at one end to the wallsof the holes, and the other end projectedabout 20 cm beyond the holes. Diageneticpyrite was present in siltstone clasts fromdredge 209D, and there was also a smell ofhydrogen sulphide gas from the mud andclasts.

(3) Kula Mud Volcano: Several small mud volca-noes lie on a triangular plateau of limitedextension at the northern corner of the easternAnaximander Mountains. Kula is the largest,measuring about 1600m across and about100m high. It is remarkably circular anddome shaped rather than conical. Observa-tions from Nautile indicate that it has resumedactivity in the recent geological past after along period of dormancy. Recent mud flowsare observed only over a small area about

200–300m across at the summit. Gas hydrateswere sampled from Kula mud volcano duringthe 1996 ANAXIPROBE expedition of R/VGelendzhik (Woodside et al., 1998) and the1999 MEDINAUT sampling expedition ofR/V Professor Logachev, and so were shellsand tube worms.

(4) Amsterdam Mud Volcano: Amsterdam is thelargest mud volcano studied. It is about 3 kmin diameter and rather flat, with a relief ofabout 20m, except for a small actively ventingparasitic cone on the western side, which has adiameter of about 350m and a height of about90m. Large volumes of erupted material haveflowed downhill to the south from Amsterdamto fill a small basin with over 12 km3 of mudbreccia. Concentrations of methane in thewater column above Amsterdam are as highas 13.5 ml/l and, gas hydrates were sampledfrom the central area of the mud volcanoduring the 1999 MEDINAUT expedition ofR/V Professor Logachev.

The lucinid material was collected by differ-ent bottom sampling techniques at differentlocations in the Anaximander Mountains duringthe ANAXIPROBE Programme expedition onR/V Gelendzhik (Woodside et al., 1997). The mainsampling device was a large diameter (14 cminternal diameter) gravity corer with a length of6.6m and a weight of about 1500 kg. Thesecondary sampling device was a pipe dredge witha diameter of 75 cm and a length of 1m.

2.2. Material examined of Lucinoma kazani n. sp.

ANAXIPROBE Programme

St. 209D (35125.900N–30124.550E to 35127.100N–30124.610E; 1854–1165m deep; Faulted Ridge):One complete shell (38.4mm length� 35.0mmheight), selected as holotype, in MNHN, Paris.St. 234G (35125.730N–30133.640E; 1709m deep;

Kazan Mud Volcano): One live collected specimen(37mm length� 33.4mm height), selected asparatype, in MNHN, Paris. Two shells (24.2mmlength� 23.7mm; 33.7mm length� 29.1mmheight), both selected as paratypes, in NMNH,Leiden.

C. Salas, J. Woodside / Deep-Sea Research I 49 (2002) 991–1005994

MEDINAUT Programme

St. MN10 (33143.50N–241410E; 1700m deep;Kazan mud volcano): 5 living juveniles and 90valves.St. MN12 (33143.50N–241410E; 1700m deep;

Kazan mud volcano): 4 juvenile shells and 33valves.St. MN9 (35127.10N–30124.40E; 1300m deep;

Faulted Ridge): 1 shell and 6 valves.St. MN11 (351260N–30127.50E; 1630m deep;

Kula mud volcano): 1 valve.St. MN13 (351200N–30116.50E; 2030m deep;

Amsterdam mud volcano): 5 valves.Comparative material examined (all from

MNHN, Paris, except where otherwise stated):Lucinoma borealis: Palermo (Italy), coll. Lamy:

1 shell; Messina (Italy), coll. Lamy: 2 shells;Ajaccio (Corsica), coll. Jousseaume: 3 shells;Portugal, Algarve, Sagres (37100.70N, 08155.00W,12–17m): several juvenile specimens; Brittany,Finist"ere, Pointe Noire: 1 shell, leg. Gofas; Ros-coff: 1 shell, leg. Gofas; Western Norway, Storresoilfield, 25–34m: 11 specimens, leg. AKVAPLAN-NIVA Programme (Swedish Museum of NaturalHistory, Stockholm); Azores, ‘‘Bia-cores’’ expedi-tion st. 81 (39100.50N, 28104.50W, 142m): 1juvenile specimen and 1 valve.

Lucinoma ignota Locard (=L. borealis ): Gulf ofCadiz ( Southern Spain): 1 valve, syntype from‘‘Talisman’’ 1883 sta. 2 (361530N, 081320W, 99m).

Lucinoma filosa (Stimpson, 1851): Philadelphia:1 shell, leg. Cope; South of Newfoundland, 323–322m: 7 shells, leg. von Cosel.

2.3. Preparation for SEM observation

To observe the location of the bacteriocytes inthe ctenidia and the bacterial morphology byscanning electron microscopy (SEM), small piecesof gills of L. kazani and of L. borealis were taken.The gill tissues were washed in distilled water anddehydrated through a graded ethanol series; thesamples were then critical-point dried. After this,the pieces of gills were mounted on stubs, coatedby argon-sputtered gold-palladium, to be viewedin the SEM and photographed.The qualitative and quantitative compositional

analysis were carried out in a JEOL JSM-6400

scanning electron microscope, with LINK ANA-LITICAL 6508 electronic microsonde and withLINK ANALITICAL EXL for the detection oflight elements and microanalysis. For the elemen-tal analysis, the pieces of gills were coated byargon sputtered silver instead of gold, because thepeak generated by gold would mask the sulphurpeak in the spectrum.

3. Systematics

3.1. Family Lucinidae

Genus Lucinoma Dall, 1901Type species: Lucina filosa Stimpson, 1851, by

original designation.The species collected in the Anaximander

Mountains is referred to the genus Lucinoma, ataxon originally introduced as subgenus of Pha-

coides Blainville 1825. Shells are usually large,lentiform, concentrically lamellose or striated andwith periostracum. The cardinal teeth are welldeveloped, with 2a (LV) and 3b (RV) bifid; thelateral teeth are obsolete or absent. The innermargin is entire, and there is no pallial sinus.In European waters, the genus Lucinoma is

represented by only one species L. borealis (Linn!e,1767), which ranges from the north of Norway(H�isaeter, 1986) to Morocco (Pasteur-Humbert,1962) and Mauritania (Cosel, pers. comm.), andinto the Western Mediterranean, where it is rare(Lamy, 1920, p. 198). There are no records fromthe Eastern Mediterranean even in the bettersurveyed areas such as the Levantine coast (Barashand Danin, 1992), Greece (Zenetos, 1996), orMalta (Cachia et al., 1993).

3.2. Lucinoma kazani new species (Figs. 2 and 3,

6–8, 12–14 and 17)

Type material: Holotype: One shell (38.4mmlength� 35.0mm height), from ANAXIPROBE96, St. 209D, deposited in Mus!eum Nationald’Histoire Naturelle, Paris, (MNHN). Paratypes:a live collected specimen (37mm length� 33.4mmheight) from ANAXIPROBE 96, St. 234G, depo-sited in MNHN; two shells (24.2mm length� 23.7mm height; 33.7mm length� 29.1mm height)

C. Salas, J. Woodside / Deep-Sea Research I 49 (2002) 991–1005 995

from ANAXIPROBE 96, St. 234G, deposited inNational Museum of Natural History, Leiden.

Type locality: Anaximander Mountains, southof southwestern Turkey, Eastern MediterraneanSea (35125.900N–30124.550E to 35127.100N–30124.610E; 1854–1165m deep).

Description: Shell medium-size, convex, thickand solid, equivalve. Outline subcircular, slightly

longer than high; inequilateral. Posterior margincurved, antero-dorsal margin forming an obtuseangle, ventral margin gently curved. Umbossomewhat prominent directed inwards and proso-gyrous, just in front of the vertical midline. A

Fig. 2. Lucinoma kazani n. sp., internal view of the holotype,

left valve. Actual length 38.4mm.

Fig. 3. Lucinoma kazani n. sp., internal view of the holotype,

right valve. Actual length 38.4mm.

Fig. 4. Lucinoma borealis (Linn!e, 1767), internal view of a

specimen from subtidal sands near Pointe du Minou, Brittany,

left valve. Actual length 31mm.

Fig. 5. Lucinoma borealis (Linn!e, 1767), internal view of a

specimen from subtidal sands near Pointe du Minou, Brittany,

right valve. Actual length 31mm.

C. Salas, J. Woodside / Deep-Sea Research I 49 (2002) 991–1005996

slight and ill-defined radial depression from thebeginning of ligament to the postero-ventralcorner and another one less defined, almostparallel to the anterior dorsal margin.

Sculpture of fine and irregularly spaced con-centric lamellae, more prominent near the ventralmargin. Outer surface of shell white. Periostracumthin, cream on the umbos, brownish-beige from

Fig. 6. Lucinoma kazani n. sp., holotype, external view of the

left valve.

Fig. 7. Lucinoma kazani n. sp., holotype, dorsal view.

Fig. 8. Lucinoma kazani n. sp., prodissoconch of a juvenile shell

from MEDINAUT expedition, Kazan mud volcano St. MN10.

Scale bar=100mm.

Fig. 9. Lucinoma borealis (Linn!e, 1767), same specimen as

Figs. 4 and 5, external view of the left valve.

Fig. 10. Lucinoma borealis (Linn!e, 1767), same specimen as

Figs. 4 and 5, dorsal view.

Fig. 11. Lucinoma borealis, prodissoconch of a juvenile from

Algarve. Scale bar=100mm.

C. Salas, J. Woodside / Deep-Sea Research I 49 (2002) 991–1005 997

the middle part of the valve to the margins.Ligament external, long and sunken, reachingalmost to the posterior margin. Escutcheon notvisible. Lunule well delimited, lanceolate,E2/3 the

length of the anterior dorsal margin and brownishin colour.Prodissoconch semicircular, relatively large

(240 mm in diameter) without particular sculptureand very eroded by the reduced environment(Fig. 8).Hinge plate relatively narrow, slightly curved.

Right valve with two cardinals, the anterior (3a)simple and the posterior (3b) bifid; one smallknob-like anterior lateral (Ia); posterior lateralobsolete. Left valve with two cardinals, theanterior one (2a) bifid and the posterior one (4b)simple; a knob-like anterior lateral (IIa), ratherprominent; posterior lateral obsolete.Well delimited adductor muscle scars; the

anterior one longer than the posterior one andrelatively large, with the proximal part sunken inthe hinge plate, under the anterior lateral; and witha rather long ventral extension, slightly divergingfrom the pallial line. Posterior adductor scarsubelliptical in outline. Small and elliptical ante-rior pedal muscle scar, close to the anterior frontof the anterior adductor muscle scar. Pallial line

Fig. 12. Lucinoma kazani n. sp., SEM pictures of a critical-

point dried fragment of the gills, general view of the gill’s edge

in transverse section. Scale bar=1mm.

Fig. 13. Lucinoma kazani n. sp., SEM pictures of a critical-

point dried fragment of the gills, several bacteriocytes. Scale

bar=10mm.

Fig. 14. Lucinoma kazani n. sp., SEM pictures of a critical-

point dried fragment of the gills, an open bacteriocyte showing

the symbiotic bacteria. Scale bar=10mm.

Fig. 15. Lucinoma borealis, SEM pictures of a critical-point

dried fragment of the gills, general view of the gill’s edge in

transverse section. Scale bar=1mm.

Fig. 16. Lucinoma borealis, SEM pictures of a critical-point

dried fragment of the gills, bacteriocytes. Scale bar=10mm.

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well defined and beginning near the anteriorborder of the anterior adductor scar. Inside ofthe shell white, with oblique scars about themiddle of the valves, and with slight ridgesreflecting the exterior depressions of the shell.

Inner margin of the valves smooth. Long and finepallial blood vessel impressions, crossing obliquelythe valves from the postero-dorsal to the antero-ventral side.

Anatomy: An elongated cylindrical foot, with asmall posterior heel, is present.Ctenidia large and thickened, composed only of

the inner demibranchs. Labial palps minute.Fig. 12 (SEM level) shows a transversal sectionin part of the large demibranch of L. kazani withthe bacteriocytes, which contain the intracellularsymbionts. These can be observed in Fig. 13 asbulges set very close each other. Fig. 14 shows oneopen bacteriocyte, in which the intracellularsymbionts (bacteria) can be observed. The size ofthe observed bacteria ranges between ca. 1 and3 mm, showing different shapes, from round toelliptical forms. The qualitative and quantitativecompositional analysis of some of these symbiontsfrom different bacteriocytes (Table 1) shows apercentage of organic carbon higher than 90% anda percentage of sulphur lower than 1%, thusindicating that they are living organisms and notsulphur deposits. The symbionts are smaller thanthe cystine-rich granules (up to 6.5 mm) describedin Codakia orbicularis by Frenkiel and Mou.eza(1995).No mantle gills, either near the anterior

adductor or near a mantle septum, have beenobserved. Mantle edge smooth along most of itsextension, fused only a short distance anterior tothe inhalant aperture. The mantle folds are bluntand bear a single row of minute papillae on eachside of the inhalant aperture; there are no papillaearound the exhalant aperture (Fig. 17). The ‘‘ex-halant siphon’’ (according to the terminology of

Table 1

Qualitative and quantitative elemental analysis of bacteria from several bacteriocytes of the ctenidia of Lucinoma kazani n. sp.

Elements Ba 1 Ba 2 Ba 3 Ba 4 Ba 5 Ba 6 Ba 7 Ba 8

C 93.43 96.15 93.35 95 90.8 93.95 95.45 96.46

O 5.2 2.1 5.4 3.3 7.75 3.6 2.43 1.02

Ca 0.51 0.67 0.36 0.53 0.41 1.01 0.91 1.26

Si 0.11 0.07 0.12 0.13 0.19 0.12 0.1 0.1

P 0.31 0.45 0.24 0.35 0.42 0.44 0.42 0.5

S 0.34 0.4 0.43 0.49 0.34 0.66 0.49 0.55

Mg 0.11 0.16 0.21 0.24 0.2 0.1

Fig. 17. Lucinoma kazani n. sp. Posterior mantle apertures of a

paratype from Kazan mud volcano, Anaxiprobe sta.234G.

Exhalant aperture (Ea) and inhalant aperture (Ia).

C. Salas, J. Woodside / Deep-Sea Research I 49 (2002) 991–1005 999

Allen, 1958) is retracted, giving the appearance ofa funnel turned inwards on the preserved speci-men.

Distribution: The species is known only from theAnaximander Mountains, Eastern MediterraneanBasin.

Habitat: The live specimens were taken on theKazan mud volcano at a depth of 1700–1709m, ina mud breccia deposit that was highly saturated ingas and contained many fragments of softsediments and hard rocks up to 7 cm in size. Othershells and valves were dredged from the southernslope of Faulted Ridge, a fault escarpmentexposing several hundreds of metres of verticalrock outcrop with gas seeps evident at beddingplanes and fractures. They were found, in theupper part of a dredge haul which started at adepth of 1854m and finished at 1165m, in rockswith irregular holes running through them, whichwere inferred to be authigenic carbonates with gasescape structures.

Etymology: The name refers to the Kazan mudvolcano in the Anaximander Mountains, the mudvolcano at which living specimens were found(Figs. 2 and 3, 6–8, 12–14 and 17).

4. Discussion

4.1. Taxonomic comparison

The shell of L. kazani resembles that of L.

borealis (Figs. 4–5, 9–11), but the umbos are moreprominent and directed more inwards and for-wards; the lunule is distinctly broader and betterdelimited; the hinge plate is slightly larger and lesscurved than in L. borealis. The hinge is alsodifferent, L. kazani having a rather prominent,knob-like anterior lateral in each valve, whereasthese are obsolete in L. borealis. Another impor-tant difference is the anterior adductor scar, whichis broader in L. kazani, with the proximal partnearly twice the size of a L. borealis of comparablesize. Moreover, the angle between the ventralextension of the anterior adductor and the pallialline is greater (ca. 251) than in L. borealis (ca. 151).We have examined the figured syntype of Lucina

ignota Locard, 1898, described from the Gulf of

Cadiz (Southern Spain) and agree with Lamy(1920) that it is L. borealis.The gills of the two species seem to be different,

L. kazani having shorter and larger ctenidia thanL. borealis (Fig. 15). However, according toHentschel et al. (2000), the phenotype of the gillsof the Lucinoma aequizonata changes, in colourand size, in relation to their ability to dispose ofsulphur from the environment. So, it could be thatthe phenotypic differences of the gills betweenL. kazani and L. borealis were related to theirnutritional condition. Both species hold symbiontsin bacteriocytes, inside the ctenidial cells, but thebacteriocytes are set very close together inL. kazani (Fig. 13), whereas in L. borealis theyare more separated by intercalary cells (Fig. 16).The labial palps of L. kazani are smaller than

those of L. borealis, but the feet in the two speciesare rather similar, with a moderately pronouncedheel.The posterior apertures, derived from the fusion

of the inner folds of the mantle, are different. Theinhalant aperture in L. kazani shows extremelysmall papillae in both sides of the middle fold(Fig. 17), but in L. borealis there are large papillae,as illustrated by Allen (1958, p. 434) and confirmedon our specimens from Norway. Moreover, themantle folds in L. borealis are large and appressedto each other, whereas in our specimen ofL. kazani these are blunt and remain apart oneither side of the aperture. The exhalant aperturesare more similar in the two species; they have nopapillae and appear as a funnel with a roundedaperture, directed inwards on the fixed specimensbut capable of being stretched out a long distancein living specimens (Allen, 1958, p. 438). However,the extension of the exhalant aperture along themargin is greater in L. borealis.The habitats of the two species are also quite

different. According to Jeffreys (1863), Lucinoma

borealis in the British Isles lives in muddy graveland sand, from spring tide low-water mark to 82fathoms (ca. 150m), and in one locality off theMull of Galloway in 110–145 fathoms (ca. 200–265m). Allen (1958) reports this species fromZostera beds at low-water mark in the Roscoffarea, France, and in the sublittoral zone (Ply-mouth Sound, Firth of Clyde, and Bay of

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Mentone, British Isles). Dando et al. (1986) foundL. borealis in Zostera beds at shallow depth, inmedium sand sediment (Salcombe estuary, southcoast of Devon), in fine sand (Yealm estuary), andin muddy sediments and black mud (PlymouthSound). The species is very abundant in fjords(Tunberg, 1984), in coarse shell sand and in siltyfine sand at shallow depth. Along the West coastof Brittany (France), L. borealis may reachdensities of 1500 specimens per m2 in someseagrass beds (Monnat, 1970, cited in Le Pennecet al. 1988). There are records from deep water(Dautzenberg and Fischer, 1906: 1530m) from theAzores; however, these are based on valvescollected around volcanic islands with very steepslopes, and specimens have been carried down-slope. Conversely, L. kazani is a genuine deep-water species, with living specimens collected inbathyal depths.Among the species of lucinids indicated from the

different Italian ‘‘Calcari a Lucina’’ deposits(Conti, 1997), Dentilucina persolida Sacco, 1901,from the flysch deposits of Outer Marnoso-Arenacea (proximal facies), Parma Foothills,resembles Lucinoma kazani in outline and sculp-ture, but the anterior adductor muscle scar ofD. persolida shows greater similarity to that ofLucinoma borealis, being longer, narrower andmore parallel to the pallial line than the anterioradductor muscle scar of L. kazani.Corselli and Basso (1996) found a Quaternary

cold seep Molluscan thanatocoenosis on the top ofthe Napoli Dome, a mud volcano located on theEastern Mediterranean Ridge, at about 1950mdepth. The valves collected have been identified bythe authors as Myrtea sp. (Lucinidae), Vesicomya

sp. 1 and Vesicomya sp. 2 (Vesicomyidae),belonging to two families typical of cold seepcommunities. However, they mention that thoughliving specimens might be present, they were notactually found on the Napoli Dome.

4.2. Symbiosis in Lucinidae

The presence of symbiotic sulphide-oxidizingbacteria in all the species of Lucinidae that havebeen investigated (Felbeck et al., 1981; Cava-naugh, 1983; Berg and Alatolo, 1984; Fisher and

Hand, 1984; Schweimanns and Felbeck, 1985;Dando et al., 1985, 1986; Reid and Brand, 1986;Distel and Felbeck, 1987; Cary et al., 1989;Hickman, 1994) has stimulated the interest in thebasic biology of living lucinids and in theirphylogeny. According to Taylor and Glover(2000) bacteria have been reported from at least30 species of Lucinidae, representing 18 differentgenera from several clades.Dando et al. (1985) have found that Lucinoma

borealis, Myrtea spinifera (Montagu, 1803) andThyasira flexuosa (Montagu, 1803), living in acommunity with the pogonophoran Siboglinum

fiordicum Webb, had enzymes characteristic of thesymbiosis in their ctenidia. Subsequentely, Dandoet al. (1986) studied the gills of L. borealis andobserved the presence of numerous prokaryotes inspecialized cells (bacteriocytes) in the subfilamen-tar region of the gills, together with high concen-trations of elemental sulphur and of a c-typecytochrome. Recently, Gros et al. (2000) havedescribed sulphur-oxidizing endosymbiosis in thelucinid Divaricella quadrisulcata (d’Orbigny, 1842),from Guadeloupe Island.Bacteria from the ctenidia of Lucinoma borealis,

Myrtea spinifera, Thyasira flexuosa and T. sarsi

(Philippi, 1845) were isolated and cultured byWood and Kelly (1989). The bacteria were eitherautotrophs, obtaining energy from oxidation ofinorganic sulphur compounds or methylotrophs,using methanol or methylamine as the source ofcarbon and energy. Wood and Kelly proposed thatthese bacteria are symbiotic with the animals, butHickman (1994) pointed out that the authors wereunable to demonstrate that the bacteria werehoused within the ctenidia. On the other hand,Cary et al. (1989) indicated that cultivations invitro of the symbionts were not successful, andsymbiont-free hosts were not as yet available.This metabolic pathway and the methano-

trophic nature of the symbionts have beendemonstrated only for a seep-mussel from theLouisiana slope (Fisher et al., 1987). These authorstested a variety of animals from the same area,which harbour chemosynthetic bacteria (the seep-mussel, 2 vestimentiferans, 1 pogonophoran, 2vesicomyids and 1 lucinid), for the presence ofmethanol deshidrogenase activity, and only the gill

C. Salas, J. Woodside / Deep-Sea Research I 49 (2002) 991–1005 1001

tissue from the mytilid contained detectable levelsof this enzyme; therefore, only the mytilid sym-bionts are capable of methylotrophy.The presence of symbionts in the ctenidia of the

new species (Figs. 12–14) is consistent with: (1) allthe studied Lucinidae, which are symbiotic withsulphur-oxidizing bacteria (Reid and Brand, 1986;Hickman, 1994); in particular L. borealis (Dandoet al., 1985, 1986; Wood and Kelly, 1989); (2) thehabitat of this new species, gas-saturated mud withinferred or observed high levels of sulphide andlow-oxygen deep-water, is usually occupied bysymbiotic organisms, including the lucinids. In thiscase the gas released upwards in the Kazan mudvolcano is methane (J. L. Charlou, 1999, pers.comm.), but in most cases the methane appears tobe associated with the zone of sulphate reduction,resulting in the generation of inorganic carbon andhydrogen sulphide. Such geochemical profiles areexpected if methane is oxidized by a consortium ofmethanogens and sulphate-reducing bacteria (Pan-cost et al., 2000). Moreover, the cores obtainedduring the 1999 Medineth cruise on the EasternMediterranean Ridge indicate that sulphate re-ducers are abundant and active in the sedimentsfrom mud volcanoes (Pancost et al., 2000); and (3)the presence in the community of vestimentiferans,which are obligate symbiotic metazoans (Childresset al. 1984; Fisher and Childress, 1984; Jones,1985), typical of cold seep and vent communities.Therefore, the occurrence of living endosym-

biont-bearing organisms, such as Lucinoma kazani

described here and Vestimentiferans (studies inprogress by E. Southwards, pers. comm.) in theAnaximander Mountains is evidence of a livingcold seep community in the Eastern Mediterra-nean basin.Extant faunal assemblages that are specific of

hydrothermal vents have not been found to date inthe Mediterranean, although fossil hydrothermalvent fauna are reported in ancient massivesulphide deposits on Cyprus (Dando et al.,1999). Recent hydrothermal sites in the Mediter-ranean are scattered and situated in shallow water(o200m). It is likely that the animals whichnormally inhabit the nearshore reducing sedimentsare unable to tolerate the prevailing high tempera-tures, salinities, and sulphide and metal concentra-

tions around the vents (Dando et al., 1995;Thiermann et al., 1997). Therefore, living chemo-symbiotic communities in the Mediterranean seemto be restricted to cold seep environments (such asfound in the Anaximander Mountains, presentpaper) or shallower reducing sediments (such asthe substrate of marine sea grasses).

Acknowledgements

We gratefully acknowledge the funding pro-vided to the second author by the NetherlandsGeosciences Foundation (Stichting GOA) to carryout the ANAXIPROBE Project (GOA Project750.195.02) and the Training Through Researchcruise TTR-6 during which the samples wereobtained. In this regard, we thank also thescientists, officers, and crew of R/V Gelendzhik

and especially M.K. Ivanov and his team fromMoscow State University. We are indebted to thecaptains and crews of the R/V Nadir, submersibleNautile and R/V Prof. Logachev. The MEDI-NAUT expedition was a jointly funded French–Dutch research project carried out on board R/VNadir and the submersible Nautile. The authorsare grateful to Jean Paul Foucher (co-chiefscientist of the project together with J. Woodside),Myriam Sibuet (responsible for biological sam-ples) and all the shipboard scientific party, whotook the biological samples and gas measurementsduring the MEDINAUT cruises. J.L. Charlou isthanked for the data on concentration of methanein the water column above Kazan mud volcano.MEDINAUT was partially supported by theResearch Council for Earth and Life Sciences(ALW) with financial aid from the NetherlandsOrganisation for Scientific Research (NWO) onthe Dutch side and by Ifremer on the French side.We are also indebted to Jos!e Pascual and AntonioRamirez (University of M!alaga) for the qualitativeand quantitative elemental analysis on the SEMpreparations. The first author thanks M. Segonzac(CENTOB, French National Sorting Center,IFREMER, Brest) and M. Sibuet (DRO/EPEcologie Abyssale; IFREMER, Brest) for entrust-ing her with the study of the collected material.Karine Olu is thanked for the preparation and

C. Salas, J. Woodside / Deep-Sea Research I 49 (2002) 991–10051002

transfer of additional bivalve samples. H.C. Salasis grateful to Prof. D. Doumenc (Laboratoire deBiologie des Invert!ebr!es Marins et Malacologie,MNHN, Paris) for the facilities provided herduring her stay in his department during thepresent study. We would like to thank Serge Gofas(University of M!alaga) and Tiphaine Zitter (FreeUniversity, Amsterdam) for help with the illustra-tions, and Rudo von Cosel (MNHN) and JohnTaylor (NHM, London) for their critical readingof the manuscript.

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