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VENUS (Jap, Jour. Malac.} Hme Vol. S9, No.3 (2000): t81-190

OriginalArticles thg

A NewSpeciesof Volutomitra ) from New(:Caledonia

Philippe BoucHET and Yuri I. KANToR

Mu.yeum national d'llistoire naturelle, 55, rue de BtofPbn, 75005 Paris, France andA.N. Severtzov lnstitute ofProblems qf'Evotution (tfRus.s'ian Academ.y ofSciences, Leninski prosp, 33, Mosc'ow i1707J, R"ssia

Abstract: Volutomitra glabella n. sp., from off New Calcdonia, js the secend representative of the frem the tropieal South-Wcst Pacific, where it has been recorded alive on hard bot- toms in 258-525 m. Its anatomy is cssentially similar to that of othcr boreal, Antarctic and

Austratasian of Volutomitridae. It is sympatric with Lhe V. vaubani species-complex,

frem which it ditifers by its larger adult size (17-25 mm), mure vividly celoured shcll, and larg- er protoconch (,average diameter 14ZLOum vs average I030 ptm in V. vaubani).

Keywords:South Pacific. new species, bathyal, hard bottoms

Introduction

The family Volutomitridae has a predominantly antiboreal distribution, with several genera and species in cold waters in the southern hemisphere, especially South Australia and New Zea}and,

and a few species at high northern latitudes in the Pucific and Atlantic oceans. Volutomitrids

remain exceptional at low latitudes, tmd then occur essentially in deep water. Based on material

dredged off New Caledonia, Cernohorsky (1982) described Vbtutomitra vaubani from a depth of 395 m, Examination of the vast materjal collected since then by expeditions in the New Caledonia

area revealed that, in addition to many samples of that species, a second, larger, and undescribed species of Vblutomitra co-occurs with it. The purpose of the present puper is ro describe and name this new species, which in the last two years has occasionally appeared on the shell market as Vblutomitra va"hani,

[faxonomy

Class Gastropoda Unranked group Caenegastropoda Order Family Volutomitridae Gray, 1854 Genus Vbtutomitra H. & A. Adams, 1853

7ype specieb': Mitra groentandica in MOIIer, 1842 (by subsequent designation, Fischcr 1884). Recent, northern Nerth AtlanticBeck

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Vbtutomitra glabetla n, sp. (Figs. 1, 2 A, C-E, 3-5)

7iype material: Holotype and 4 paratypes in MNHN, 1paratype each inNMNZ,AMS, NSMT, NMP.

7i}7)e locality: of NewCaledonia,Banc Eponge. 24057'S,168021il-],517-559 m [SMIB 1O, sta, DW205].South

Material examined (all in MNHN): North of New Caledonia. LAGON, R/V Atis: sta, 444, Atell de Surprise, 18"15'S, 162059'E, 300-350 m, 1 lv.- Sta, 475, 18e36'S, 163el1'E, 415-460 m, 1 dd juv.- Sta. 1 152, 18"58'S, 163024'E. 335 m, 3 lv (1 paratype NMNZ). MUSORSTOM 4, RfV 1'kuban: sta, DW156, 18054'S, 163019'E, 525 m, 7 dd, 1 lv juv.- Sta. DW164, 18033'S. 163"13'E, 255 m, 3 dd,- Sta, CC174, 19"OO'S, 163"18'E, 365 m, 1 dd.- Sta, DW181, 18"57'S, l63a22'E, 350 m, 2 dd, 3 lv (1 paratype MNHN, Figs IF-G).- Sta. DW]84, 19e04'S, 163e27'E, 260 m, 4 lv.- Sta. DW196, 18055'S, 163024'E, 450 m, 1 lv,

SMIB 6, RIV Atis: sta. DWI15, 19abOO'S. 163e27iE, 280-285 m, 1 ddr Sta. DWI18, 18"58'S, 163026'E, 290-300 m, 2 dd.- Sta. DWI19, ]8e59'S, 163"26'E, 295-305 m, 1 dd.- Sta. DW120,

18058'S, l63026'E, 310-325 m, 2 dd.- Sta. DW121, 18058-S, 163-26'E, 315 m. 4 dd.- Sta. DW122,

18058tS, 163025'E, 325-330 m, 2 dd.

BATHUS 4, RIV Atis: sta. DW903, 190oo'S, 163014'E, 386-400 m, 2 dd.- Sta. DW906, 1900I'S, 163015'E, 339-350 m. 1 dd.- Sta. DW908, 18th58'S, 163"11'E, 502-527 m, 1 dd.- Sta. DW91g, 18049'S. 163016'E, 613-647 m, 2 dd.- Sta. DW923, 18"52'S, 163e24"E, 470-502 m, 1 lv, 4 dd (1 paratype MNHN Fig. ID-E; Fig. 2A),- Sta, DW924, 18n55'S, 163024'E, 344-360 m, 13 dd.- Sta. DW 92S, 18055'S, 163"24'E, 370-405 m, 4 lv,- Sta. DW926, 18e57'S, 163025'E, 325-330 m, s dd.-

Sta. DW929, 18e52'S, 163023'E. 502-516 m, 1 dd,- Sta. DW931, 18055"S, 163D24'E, 360-377 m, 7

dd.- Sta. DW932, 19008'S, 163e29"E, l70-190 m, 1 ddi Sta. DW936, 19004'S, 163028'E, 252-258 m, 2 lv (1 paratype NSMT).- Sta. DW939, 18052'S, 163"23'E, 502-516 m, 1 dd.- Sta, DW940, 19000'S, 163026'E, 305 m, 4 dd (1 paratype NMP).- Sta. DW941, 19"02'S, 163D27'E, 270 m, 5 lv, 3 dd (2 paratypes, Figs IC, H),- Sta, DW942, 19"04'S, 163"27'E. 264-270 m, 9 dd (1 paratype AMS). Seuth of New Caledonia. SMIB 10, RfV Atis: sta. DW205, 24"57'S, 168e21'E, 517-559 m, 1 dd (holotype).- Sta. DW207, 24"57'S, 168"21'E, 508-553 m, 1 ddjuv.

Distrihution: North Southof New Caledonia, on hard bottoms,alive in258-525 m, shells in 190-613 m (Fig. 4).and

Desct'iption of the hototype: Shell solid, glossy, broadly ovoid, width 5e% of height, consisting ol' ca. 2,O protoconch and 5.5 teleoconch whorls. Protoconch eroded in holotype, in a specimen from BATHUS 4 sta. DW923 it is large, bulbous, diameter 1400 "in, exposed height 1200 "m, 11'rst whorl convex, smooth, second whorl with flat sides, a few axial ribs before indistinct protoconch- teleoconch beundary (Fig, 2A), Teleoconch wherls moderately convex with tightly impressed suturc, ramp slightly concave, exposed part below periphery rnore convex. On early teleoconch whorls sculpture consisting of strong straight ribs, crossed by spiral cords, intersection grossly can- cellate; 21 ribs on first whorl, 24 on second, 34 on third; axial ribs gradually becoming obsoletc on last twe whorls, last whorl only with indistinct undulation$ at shouldcr; 3 spiral cords, rather prumi- nent on first two whorls, inconspicuous and restricted to subsutural ramp on subsequent whorls.

Last wborl smooth and glossy, canal with 19 strong, closely spaced cords. Aperture height 70% of

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Fig. 1. Shells ef Vblutomitra gtabeXla n, sp. (A-H). A-B. Holotype. C. BATHUS 4, sta. DW 941. D-E. BATHUS 4, sta. DW 923. F-G. White form MUSORSTOM 4, sta. DW 181. H. BATHUS 4, sta. DW 941. Velutomitra vaubani Cernohorsky, 1982, (I & J) South of New Caledonia. I. SMIB2, sta. DW8. 22e54'S, 167"13'E, 435-447 m. J. Holotype, RtV Vtiuban 1978-79, sta. 15, 22=49'S, 167012'E, 395 m, Scale bar = 1 cm.

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Fig. 2. Vbl"tomitra gtabeUa n. sp. (A, C, D, E) and V vaubani (B). A, B. Protoconchs. A, C. Juvenile. BATHUS 4, sta. DW923. B. sta, DW46, 22'53"S,

167'17'E, 570-610 m. D. Radula. MUSORSTOM 4, sta. DW184,BIOCAL,shell height 20.4 mm. E. Jaw, (Same specimen as D).

shell height, narrowly-elongate, smooth inside, outer lip simple. Columella without a callus, with 4

widely spuced p]aits, the most abapicaL smallest. Siphonal canal shert, straight. Colour ef the shell

pinkish brown, with irregularly spaced subtriangular lighter spots, some of which coalesce and

form irregular axial stripes near shoulder. Din:ensions: shell height 24.0 mm, diameter 12,O mm, last whorl height 19,1 mm, aperture height 16.5 mm, Dimensions of the largest specimen (BATHUS 4 sta. DW925): shell height 25.0 mm, diameter 13.0 mm, last whorl height 20.3 mm, aperture height 18.3 mm.

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Bouche{ & KanLor: New Species of Vblutomitra 185

ilthzl p]1

,tt'SE'

ocm

D eeme

C ...... / F

Fig. 3. Anatomy of Vblutomitra gtabeUa n. sp. A-E. MUSORSTOM 4, sta. DWI84. F. BATHUS 4, sta. DW923, A, B. Body removed from shell. C. Mantle. D. Anterior digestive system, oesephagus uncoiled, pro- boscis sheath epened to show the proboscis. E. Stomach. F. Operculum ofjuvenile. Abbreviations: aldg, anterior lobe of the digestive gland; cae, caecum of thc stomach; cme, cut man- tlc edge; ct, ctenidium; ddg, ducts of the digestive gland; gl., gland ef Leiblein; hg, hypobranchial gland; mpo, muscular posterior part of the mid-oesophagus; mr, muscu]ar rod embracing the radu- 1ar sac; ne, nephridium; ng, nephridial gland; nr, nerve ring; os, osphradium; per, pericardium; pg, pallial gonoduct; pldg, posterior lobe of the digestive gland; pr, proboscis; prp, propodium; re, rec- tum; s, siphon; sg, salivary gland; t, cephalic tentac}e; vc, ventral channel of thc valve of Leiblein; vL, valve of Lciblcin; vpr, vcntral proboscis retractor.

Anatomy: A female specimen from MUSORSTOM 4 sta. DW184 has been studied anatomical- ly. It had a shell height of 20,4 mm, diameter 1O.3 mm, last whorl height 16.3 mm, aperture height

14.1 mrn.

Externat anatomy (Figs. 3 A-B): The body consists of 3.5 whorls, the mantle spans O.8 whorl, the nephridium nearly O.5 whorl, and the digestive gland 2.25 whorl. The body is yellowish, it lacks specific pigmentation except small ponions along the mantle edge and near the siphon base, as well

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as the propodium of the foot, which bear very small dark-purple spots, Operculum absent, although there is a smal1 callus on the upper surface of the foot at the place of the operculum. The fbot has an ovate solc, folded lengthwise. The siphon is short, simple, slightly protruding beyond the man- tle edge. The columellar muscle is thick anteriorly, consisting of 1.5 whorls, with four deep

grooves, corresponding to the columellar plaits. In its posterior part, in the place of the attachment

to the columella, the musc]e is splitted into separate branches. each insening between the columel-

lar plaits. The mantle edge is rather thick, while thc central part of thc mantlc is extremely thin and transparent, the mantle organs poorly visible through it. The mantie covers the head base. The head

is narrow and small, with very short conical tentacles and large eyes.

In a juvenile with a 5.6 mm shell height (BATHUS 4, sta, DW923), a very smull operculum was present (Fig, 3F), length O.85 mm, or 23% of aperture height. It is narrow oval, transparcnt, its nucleus already corroded, with very few growth lines and long and narrow attachment of the co]- umellar muscle (dottcd on the figure), In shape the operculum is similar to that illustrated by Cernohorsky (1982) for V vaubani.

Mantle (Fig. 3C): The ctenidium is very long and nearly occupies the entire mantle length, it is curved and clcarly separatcd into two parts. Anterior part (close to the mantle edge) is narrow and

the lamellae are digitifbrm with narrow base, hanging freely into the cavity. Posteriorly the ctenid-

ium becomes much wider and consists of normal, high triangular lamellae. The osphradium is

large, slightly broader than the ctenidium, asymmetrical, with the right side at leasr 1 .5 times broad-

er. The hypobranchial gland is well developed, and covers purt]y the rectum und the pallial gonod- uct. The rcctum is very narrow and runs along the pallial gonoduct. The anal gland was not seen

during dissection.

Digestive s.ystem: The proboscis in retracted position is conical, short, about 1.6 mm, or 11% ol'

aperture height, and occupies less than half the length of the rhynchocoel. Mouth opening circular.

The large unpaired ventral proboscis retractor (Fig. 3D, vpr) is attached to the anterior part of the rhynchodaeum (proboscis sheath) and to the bottom of the cephalic haemocoe]. The small $erni- closed funncl-shaped chitinous jaw (Fig, 2F) lines the anterk)r surface of the buccal cavity. The radula is 3,4 mm long, er 25% of aperture length. Its morphology is typical for Volutomitridae (Fig. 2D). The wishbone-shaped central teeth have a rather long central cusp, with a deep dorsal groove that interlocks which the cusp of the preceding row. The fbrming part is long and comprises 20% of the whole radular length. Thc part of the radula in the sublingual pouch is remarkably long and comprises 113 of the whole membrane length. In its posterior part, the radular sac is embraced by a muscular convoluted rod, which protrudes beyQnd the rear of the proboscis (Fig. 3D, mr), The merphology of the ocsophagus is very similar to that described fOr ether Volutomitridae (Ponder, 1972; Arnaud & van Mol, 1979; Kantor & Harasewych, 1992). The anterior oesophagus (betwcen proboscis and valve of Leiblein) is relatively short and thin. The valve of Leiblein (Fig. 3D, vL) is poorly demarcated and has nearly the same diameter as the oesophagus, but it is distinguished by a more whitish co]our. Thc torsion of the posterior part of the valve of Leiblein is made visible by the rotation of the ventral channel (Fig. 3D, vc), which is seen as a dark strip through the valve wall. Soon after the valve, the mid-oesophagus widens considerably and forrns two loops before passing through the circum-oesophageal nerve ring. The posterior part of the mid-oesophagus (Fig.

3D, mpo) becomcs cxtremely thick and convoluted unti] the entrance of the yery small and tubular

gland of Leiblein (Fig. 3D, gL). The posterior oesophagus (after the opening of the gland ot'

Leiblein) becomes very narrow and thin-walled, and fbllows to the stomach. The stomach is reLa-

tively small and adjoins the nephridium. The stomach itself has a smal1 caecum (Fig. 3E) and is simiLar to that of Pecu.tator hedteyi (Ponder, 1972). Thc digestive gland is clcarly bilobate. the

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Bouchet & Kantor: New Species of V71uromitra 187

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,;.r.f,,. ･-Elltz-. . : No. of spec. × 4

'.tt. /t.. t eq t ', lhlutomitra .･･''' glabella ' '' '-.. ',.. type locality " ' ''': ', iir examinedmaterial ' tt. tt/ 'ttttttt tt ' ' 1:t ttt .･. tt tt.tt i,'' ,.er

Fig. 4. Geographical and bathymctrical distribution of Volutomitra glabelta n. sp. Each column represents 4 specimens Clv or dd).

small antcrior (righO lobe (Fig. 3B, aldg) lies in frent of the stomach, while most of the gland is formed by the posterior (left) lobe (Fig. 3B, pldg). The ducts of the digestive gland are paired, very closely spaced at the entrance of the oesophagus into the stomach (Fig. 3E, ddg). The salivary glands are fused, but with rather thick paired ducts which, ttfter leaving the gland, fbllow along both sides of the oesophagus and enter its wall in front of the valve Df LeibLcin. The accessory sali- vary gland, if present, was not seen during the dissection, probably due to its very minute size, as in other Volutomitridae.

Remarks: Vblutomitra glabelta is rather constant in shell shape, with adult sizes ranging from 17.0 to 26.5 mm and a diamctcrtheight ratio ofO.49-O,54 (average O.51, a= O.08). It is more vari-

able in terms of degree of sculpture and colour. In some specimens, the spiral cords are well dcfined on the last wherl as well as on the spire whorls, but even then arc always less pronounced on the periphery. There may be up te 9 cords on the penultimate whorl and 6 well defined subsu- tural cords on the last whorl. The colour ranges from clearly spotted to axially stripped, but therc

are also completely white specimens.

Etymology: The specific name is that of the West African marginellid A4arginetla glabetla (L,, 1758), which bears a general superficiaL resemblance to the present new Vbkttomitra. It is uscd as

a noun ln apposMon.

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Discussion

Conif)ari.son "'ith other species: The distribution of Votutomittzi glabelta appears to be diajunct, with a number of stations clus- tcrcd in an area of 80 kin of maximal extension near Grand Passage, North ef New Caledonia, and

isolated occurences on Banc Eponges, on Norfblk Ridge, South of Ncw Caledonia. It is not known

whether its absence off New Caledonia itself is apparent or real, About l50 hauls havc been made

off the coasts of New Ca]edonia. but thcsc have mostly hit the sediment cone ut the outlet of thc reet' passcs, whereas U gtabella appears to prefer hard bettoms. Both in the North and Suuth of New Caledonia, V glabella is sympatric with other volutomitrids that superficially resemble V

vaubani. Several species appear to be involved and we refer to them as the Vbtutomitra vaubani species-complex. VL vaubani s.l. outnumbers the ncw species in the South, while V glabella out- numbers U vaubani s.l, in the North. In the North they co-occur syntopically at several stations

(MUSORSTOM 4 sta, DWI56, DW164; BATHUS 4 sta. DW929), and in the South they are also

fbund microsympatrically on Banc Epongc, although not in the same haul.

Vblutomitra glahella differs from the U vaubani species-complex by its broader shetl with lower spire, and usuully much larger size, although the size of the largest V, vaubani and smallest adult gtahella stight]y overlap (17.0 mm). Even at comparable sizes, young or small V glabeUa dif- I'er from V vauhani by their significantly larger protoconch. In V glabella the protoconch has an "vaubani" average diameter of 1440 "m (range 1350-1650 um, a= O.1 l, n = 8), while in complex it ls in average 1e30 ptm (runge 900-1200 ptm. a= O.08, n = 1O). Moreover in gtabella protoconch is slightly more e]cvated (average ratio ot' exposed heightidiameter is O.86, a= O.08 for gtabella and 0.80, a= O,08 for vauhani). The new species is clearly scparated on a scattered diagram (Fig. 5). V gtabeUa has also mere numerous (42-52 on the penultimate whorl vs 22-35) and weaker axial ribs.

1.5 mm =o==oo :.tdeRi.n.

o

1.0 mm

O.5 mm 1.0 mm 1.5 mm 2.0 mm Protoconch diameter

Fig. 5. Seattered diagrum of protoconch measurements of Vblutomittu glabella and Vbtutomitra va"bani.

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Bouchet & Kantor: Ncw Species of W)lutomitra 189

Generic position: Conchological and anatomical characters point to a generic allocation of the new species to Volutotnitra, but it also has resemblance to spccies earlier classified in the nominal genus Waimatea Finlay, 1927 (type-species Mi.tra inconspictta Hutton. 1885, upper Eocene of New Zealand). Cernohorsky (1970) had considered Wtiimatea a valid genus, then including 1O fossil and one Recent species, but when he described V?)lutomitra vaubani from New Caledonia, he treated

WZiimatea as a subgenus of IV?)lutomitra, based on similarities in opercular and radular characters

(Cemohorsky 1982). Mitra incenspictia is conchologically most similar to Vblutomitra banksii (Dell, 1951) from the Chatham Rise, Ncw Zealand, and to VL groenlandica, the type-species of Vbtutomitra, and we follow Maxwell (1992) who synonymized Waimatea with Vblutomitra.

"VVaimated' Ponder (l998) emphasised that the generic position of obscura (Hutton. I873) is unccrtain, and species earlier classified in Vtiaimatea may not all belong to Vblutoinitra, The anatomy of the digestive system of Vbl"tonzitra glabeUa agrees with that of all othcr yolu- tomitrids examined so fai': Peettlator hedlevi (Murdoch, 1905) and Microvotuta marginata (Hutton, 1855) (Ponder, 1972); Vblutomitra .fhagitlima (Watson, 1882), V ct{rta (Strebel, 1908) (Arnaud & van Mol, 1979). and U ala,s'kana Dall, 1902 (Kantor & Harasewych, 1992), There are only minor differences in the proportions of the length ef different parts of the oesophagus, as well as in the degree of fusion of the salivary glands and the presence or absence of cilia in the valve of Leiblein. Kanter & Harasewyeh (1992) speculated on how thejaw and radula of Vbtutornitra alaskana might be used and their discussion also applies here. Although the jaw in Vblutomitra glabeUa has a somewhat different shape (e,g,, it does not overlap ventrally, as in V/ ataskana). the functioning of the whele system is probably the same. We are not certuin whether a chitinous shield is present. since we did not preparc and cxamine seetions of the proboscis, but it probably is not present: dur- ing the preparation of the radula andjaw of V. ala,gkana, the shield usually remains once the tissues of the proboscis have been dissolved, while nothing remained once the tissues of V glabetta had been dissolved. The operculum appeurs to be vestigial in the Votutomitridae and this may account for appar- ent]y contradicting statements in the Literature. An operculum was recorded to be present in Vblutomitra curta and absent in U .fragillima (Arnaud & van Mol, 1979). Cernohorsky (1970: plate 13, figs 3-4) illustrated a rather large (ca. 6.3 × 4.7 mm). oval opercu]um in V alaskana, but Kantor & Harasewych (1992) did not find an operculum in the 24 specimens of that species they examined; however, an opercu]um was present in embryos. Pender (1972) found a vestigial oper- culum in Peculator hedle.v. i. In Vblutomitra glabella an operculum is present in the juveniles and absent in adults. The place of' attachment to the foot is very small and the larger part of the oper- culum is free: considering the small size of the operculum. it appears that it can casily be lost.

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ff 1440 ptm,1ttt/ nt t･l 1030 um) .

References

Arnaud. P. M. & van Mol, J.-J. 1979. Anatomy, ccology and distribution of the Volutidae and Volutomitridae (Gastrepoda: Prosobranchia) of the southern lndian Occan. 71he Vkliger 22(1): 19-31. Ccrnohorsky, W. O, 1970. Systematics of the fainilies Mitridae and Vo]utomitridae. ButL Attckland bxvt. Mus. g: 1-lgo. Ccrnohorsky. W. O. 1982 [`"1981"1. 0n a cullection of buecinacean and mitracean gastropuds (, Neogastropoda) from the Mozambique Channcl and Ntw Caledonia. Btiti. Mt{s. nat. Hist. nat. Paris. ser, 4, 3(4), section A: 985-1O09. Kantor, Yu. I. & Harasewych, M. G. 1992. Morphology of the digestive systcm c)i' Vblutomitra alaskana Dall, 1902 (Gastropoda, Pcctinibranchia, Volutomitridae), with notes on a possible mechanism uf feeding. Ruthenica 2(1): 4S-S3. Maxwell, P, A, 1992, Eocene Mollusca from Lhe vicinity of the McCulloch's bridge, Waihao River, South Canterbury, New Zealand: palcoccology and systematics. NZ GeoL Surv. Pateont. Bull, 65: 1-280. Ponder, W. F. 1972. The morpho]ogy of some miniform gastropods with special reference to their alimen- tary and reproductive systems (NeogaKtropoda). Matacotogia 11(2): 295-342. Ponder, W. F. 1988. Family Volutomitridue. Pp. 842-843 in Beesley, P, L,, Ross, G. J. B. and Wells. A. (cds), Mollusca: T7te southern s.ynthesis. futtna qfAustralia. Vbl. 5B. CSIRO Publishing: Mclbourne. i-viii. 565-1234.

[Received: March 16, 2000; Accepted: June 30. 2000]

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