Zoology Series sy THE NATURAL HISTORY MUSEUM | VOLUME 62 NUMBER 1 27 JUNE 1996 The Bulletin of The Natural History Museum (formerly: Bulletin of the British Museum (Natural History)), instituted in 1949, is issued in four scientific series, Botany, Entomology, Geology (incorporating Mineralogy) and Zoology. The Zoology Series is edited in the Museum’s Department of Zoology Keeper of Zoology: Dr C.R. Curds Editor of Bulletin: Dr N.R. Merrett Assistant Editor: Dr B.T. Clarke Papers in the Bulletin are primarily the results of research carried out on the unique and ever- growing collections of the Museum, both by the scientific staff and by specialists from elsewhere who make use of the Museum’s resources. Many of the papers are works of reference that will remain indispensable for years to come. All papers submitted for publication are subjected to external peer review for acceptance. A volume contains about 160 pages, made up by two numbers, published in the Spring and Autumn. Subscriptions may be placed for one or more of the series on an annual basis. Individual numbers and back numbers can be purchased and a Bulletin catalogue, by series, is available. Orders and enquiries should be sent to: Intercept Ltd. P.O. Box 716 Andover Hampshire SP10 1YG Telephone: (01264) 334748 Fax: (01264) 334058 Claims for non-receipt of issues of the Bulletin will be met free of charge if received by the Publisher within 6 months for the UK, and 9 months for the rest of the world. World List abbreviation: Bull. nat. Hist. Mus. Lond. (Zool.) © The Natural History Museum, 1996 Zoology Series ISSN 0968-0470 Vol. 62, No. 1, pp. 1-70 The Natural History Museum Cromwell Road | London SW7 5BD Issued 27 June 1996 Typeset by Ann Buchan (Typesetters), Middlesex Printed in Great Britain at The Alden Press, Oxford Bull. nat. Hist. Mus. Lond. (Zool.) 62(1): 1-30 Issued 27 June 1996 Deep-sea conoidean gastropods collected by the John Murray Expedition, 1933-34 ALEXANDER V. SYSOEV Zoological Museum of Moscow State University, Hertzen Street 6, Moscow 103009, Russia Synopsis. Conoidean gastropod molluscs (formerly treated within the family Turridae) from 20 deep-sea (bathyal) stations of the John Murray Expedition (North-Western Indian Ocean) are described. A total of 50 species from 11 families and subfamilies, and 31 genera and subgenera, have been found in the material studied. 17 species are described as new. INTRODUCTION The John Murray Expedition worked aboard the ‘Mabahiss’ in the northwestern part of the Indian Ocean in 1933-34 and collected invaluable biological material from the area which has not been subsequently explored on such a large scale. However, no account of the gastropods collected by that expedition have ever been published. Through the courtesy of Dr John D. Taylor of The Natural History Museum, London I was able to study the conoidean gastropods from that collection. This paper deals with deep-sea samples of that part of the Conoidea (= Toxoglossa) which was usually treated as the family Turridae. The Turridae s.l., unlike the Terebridae and Conus, are very characteristic of the deep-sea molluscan fauna, of which they comprise a considerable part. The material studied was previously loaned to the late A.W.B. Powell who mentioned some of the species in two parts of his revision of Indo-Pacific Turridae (Powell, 1964, 1969). However, none of Powell’s species identifications was found on labels accompanying the samples. MATERIAL AND METHODS The molluscs studied were collected at 20 bathyal (depth 183-2312 m) stations of H.E.M.S. ‘Mabahiss’ in the North-Western Indian Ocean (Table 1). The classification of Conoidea used in the present paper follows that adopted by Taylor et al. (1993). All the material including the type specimens of the new species is stored in the Natural History Museum. ABBREVIATIONS USED IN THE TEXT The Natural History Museum, London | JME John Murray Expedition i} ial shell height 1D shell diameter | Hs spire height | Hb body whorl height | Ha aperture height © The Natural History Museum, 1996 TAXONOMY Family DRILLIIDAE Morrison, 1966. ICZN pending Genus DRILLIA Gray, 1838 Type species: Drillia umbilicata Gray, designation Gray, 1847) 1838 (subsequent Drillia altispira Sysoev, new species Figs 18 & 19 MATERIAL. stn 176, | shell (holotype, No. 1993088). DESCRIPTION. The shell is rather large, claviform, with very high spire exceeding half of the shell height, thick and solid, light-brown, consisting of almost 12 whorls. The protoconch is missing. The whorls are weakly convex and slightly angled; the point of angulation is situated below the periphery in the spire whorls, but shifts upwards on the last whorls. The subsutural slope is concave, and the prominence of concavity increases towards the body whorl. Sutures are clear, straight, and shallow. Axial sculpture consists of oblique, narrowly crested folds terminating on the subsutural slope. Some folds form weak nodules just below the suture. The folds become subobsolete on the last quarter of body whorl, probably as a result of preceding shell damage. There are 14 folds on the penultimate whorl and about the same number on the body whorl. Spiral ribs are numerous, uniform, rounded, moderately strong, with interspaces equal to them in width. The ribs cover the entire shell surface but become narrower, closer, and weaker on the subsutural slope. The shell base forms a weak bend in passing to a moderately developed fasciole. The aperture is rather small, inversely pyriform, with a distinct stromboid notch. The outer lip with a thin edge, projects strongly and forms an alate expansion between the anal sinus and stromboid notch. There is no prominent prelabral varix, only a thin fold curved in correspondence to growth lines is present. The inner lip is covered by thick and wide glossy callus which is mostly free along its outer edge and forms a shallow false umbilical cavity. The parietal callus pad is large and rounded, constricting the entrance to the anal sinus. The anal sinus is deep, U-shaped, with spout-like edge, directed slightly adapically. The canal is short, slightly bent to the right, shallowly notched and obliquely truncated. H = 37.9, Hb = 17.5, Ha = 14.2, D= 11.4 mm. 2 A.V. SYSOEV Table 1 Stations of H.E.M.S. ‘Mabahiss’ where deep-sea conoideans were collected. No. Position Area Date Depth, m Gear 26 12°29'30" N, SUI S0ME: Gulf of Aden 11.10.1933 2312 AT 33 13°41'00" N, 48°17'00" E to Gulf of Aden 15.10.1933 1295 AT 13°40'00" N, 48°18'00" E 34 13°05'36" N, 46°24'42" E Gulf of Aden 16.10.1933 1022 AT 35 13°14'24" N, 46°14'12" Eto Gulf of Aden 16.10.1933 457-549 OT 13°13'24" N, 46°10'00" E 42 17°26'00" N, 55°49'00" E Hadramaut 27.10.1933 1415 TD 62 22°53'30" N, 64°56'10" E to Gulf of Oman 18.11.1933 1893 AT 22°56'30" N, 64°56'30" E 106 05°38'54"S, 39°15'42" E to Zanzibar 12.01.1934 183-194 AT 05°40'18" S, 39°17:36"E 107 05°15'30" S, 39°33'00" E to Zanzibar 12.01.1934 421-457 AT 05°17'14"S, 39°32'48" E 118 04°05'54" S, 41°10'12" Eto Zanzibar 17.01.1934 1789 AT 04°17'00" S, 41°11'48" E 119 06°29'24" S, 39°49'54" E to Zanzibar 19.01.1934 1207-1463 AT 06°32'00" S, B95 S50 0b 122 05°21'24" S, 39°23'00" E to Zanzibar 22.01.1934 732 OT 05°22'36" S, 39°22'18" E 143 05°15'48" S, 73°22'48" E to Maldive Is. 30.03.1934 797 AT 05°13'42"S, 13°23'36E 145 04°58'42" S, 73°16'24" E Maldive Is. 02.04.1932 494 AT 158 04°42'30" S, 72°42'30" E to Maldive Is. 07.04.1934 786-1170 AT 04°36'48" S, 72°48'54" E 176 12°04'06" N, 50°38'36" E Gulf of Aden 02.05.1934 665-732 AT 180 12°03'24" N, 50°40'12" E Gulf of Aden 02.05.1934 397 G 184 14°36'06" N, 51°00'18" E to Gulf of Aden 04.05.1934 1270 AT 14°38'42" N, 50°57'42" E 185 13°48'06" N, 49°16'48" E to Gulf of Aden 05.05.1934 2000 AT 13°48'36" N, 49°16'24 E 188 13°43'18" N, 47°56'54" E to Gulf of Aden 06 05.1934 528 AT 13°46'00" N, 47°50'42 E 193 13°06'12" N, 46°24'30" E to Gulf of Aden 07.05.1934 1061-1080 AT 13°03'00" N, 46°21'42"E AT - Agassiz trawl, OT — otter trawl, TD — triangular dredge, G — grab. The new species resembles Drillia tasconium Melvill & Standen, 1901 from the Persian Gulf but differs in the high spire, larger and more solid shell, and absence of spiral sulci which deeply furrow the subsutural area in D. tasconium. DISTRIBUTION. Gulf of Aden, 655—732 m. Genus HORAICLAVUS Oyama in Taki & Oyama, 1954 Type species: Mangelia splendida A.Adams, 1867 (original designation) Horaiclavus splendidus (A.Adams, 1867) Figs 20 & 21 Mangelia splendida A.Adams, 1867, p. 309, pl. 19, fig. 24. Horaiclavus splendidus (A.Adams) — Shuto, 1965, p. 154-155, pl. 29, figs 13-15, text-figs 3, 5; Powell, 1966, p. 142, pl. 23, fig. 13; Habe, 1970, p. 120, pl. 38, fig. 13; Kuroda et al., 1971, p. 212-213, pl. 55, fig. 4; Shuto, 1975, p. 166, pl. 6, fig. 17. TYPELOCALITY. Goto Islands, Japan. MATERIAL. stn 176, 3 shells; 188, 1 shell. All the JME shells, unlike the specimens illustrated elsewhere, are uniformly coloured; the colour has probably faded since the time of collection. The protoconch consists of about 2.5 whorls, with very small and adpressed initial volution. This is more than in the holotype (Shuto, 1975, pl. 6, fig. 17; though it is impossible to evaluate the exact number of volutions from the lateral side of the illustrated shell) and in Pleistocene shells (1 2/3 whorls, Shuto, 1965). The JME shells are more similar to Japanese shells than to the geographically closer H. madurensis (Schepman, 1913). The latter species has somewhat broader and much smaller shell (holotype is 14.8 mm in height at 8 teleoconch whorls, according to Shuto, 1970, vs. 27.8-32.8 mm at 8-9 whorls in the JME shells) with shorter canal and weaker ribs. However in fact these species differ only slightly from each other, and H. madurensis seems to have no more than subspecific status. The familial position of Horaiclavus is still uncertain. Its radula was never figured, though Oyama (Taki & Oyama, 1954) mentioned that it is similar to that of Comitas and Inquisitor. Shuto (1983) described the radula of Horaiclavus as ‘true toxoglossate according to Kuroda, Habe and Oyama (1971, p. 327). However, the cited page contains no information about this genus. Until the examination of the radula, I have to follow Shuto (1975, 1983) in assigning Horaiclavus to Drilliidae. DISTRIBUTION. Japan, 50-200 m, and Gulf of Aden, 528-732 m. The present record is a considerable geographic as well as bathymetric range extension for the species. DEEP-SEA CONOIDEAN GASTROPODS 3 a Figs 1-5 Radulae. 1 — Splendrillia zanzibarica Sysoev, new species, holotype; 2 — Inquisitor adenicus Sysoev, new species, paratype, stn 33, H = 34.4 mm; 3— /. indistinctus Sysoev, new species, paratype, stn 145, H = 27.4 mm; 4— Typhlomangelia adenica Sysoev, new species, paratype, stn 185, H = 7.7mm; 5-— Borsonia ( Cordieria) symbiophora Sysoev, new species, paratype, stn 185, H = 21.8 mm. Scale-line = 0.1 mm. 4 A.V. SYSOEV 10 11 12 13 Figs 6-13 Radulae (6-9) and opercula (10-13). 6 — Typhlomangelia maldivica Sysoev, new species, paratype, stn 143, H = 27.8 mm; 7 — Bathytoma (Parabathytoma) oldhami (E.A. Smith, 1899), stn 145, H = 41.0 mm; 8— B. (P_) regnans Melvill, 1918, stn 34, H = 26.6 mm; 9- B. (P.) fissa (von Martens, 1901), stn 176, H = 35.3 mm; 10— Splendrillia zanzibarica Sysoev, new species, holotype; 11 — Inquisitor indistinctus Sysoev, new species, paratype, stn 145, H = 27.4 mm; 12, 13 — Borsonia (Cordieria) symbiophora Sysoev, new species, paratypes, stn 185, H = 21.8 mm (12) and stn 26, H = 20.1 mm (13). Scale-lines 0.1 mm (6-9) and 1 mm (10-13). DEEP-SEA CONOIDEAN GASTROPODS 15 Figs 14-17 Opercula. 14— Typhlomangelia adenica Sysoev, new species, paratype, stn 185, H = 7.7 mm; 15— T. maldivica Sysoev, new species, para- type, stn 143, H = 27.8 mm; 16— Bathytoma ( Parabathytoma) regnans Melvill, 1918, stn 34, H = 26.6 mm; 17— B. (P.) fissa (von Martens, 1901), stn 176, H = 35.3 mm. Scale—line = 0.1 mm. Genus SPLENDRILLIA Hedley, 1922 Type species: Drillia woodsi Beddome, 1883 designation) (original Splendrillia zanzibarica Sysoev, new species Figs 1, 10 & 22 MATERIAL. stn 119, 1 specimen (holotype, No. 1993089) and 1 shell (paratype, No. 1993090). DESCRIPTION OF HOLOTYPE. The shell is of medium size for the genus, slender, angularly claviform, grayish-white with dull surface, rather thin, consisting of 5.5 remained whorls. The protoconch is missing; the bluntly closed beginning of initial teleoconch whorls is seen in the shell apex. The whorls are angled slightly above the periphery, concave above the angulation and almost flat below it. Sutures are clear, shallow, slightly wavy. The spire is high, occupying about 0.4 of the shell height. Axial sculpture consists of strong oblique folds, rapidly disappearing on subsutural slope and forming pointed tubercles at the whorl periphery. Intervals between folds are narrower than the folds themselves. The folds reach the lower part of the shell base but weaken greatly on the last third of the body whorl. There are about 15 folds on the body whorl and 12 on the penultimate. Spiral sculpture is absent except for very indistinct striation seen only on the shell base near aperture. The aperture is oval and gradually narrows towards the canal. The inner lip is covered by a longitudinally rugose callus. The anal sinus is rather shallow, broadly open, with the edge somewhat spout-like. The canal is moderately long and wide. H = 19.3, Hb = 11.7, Ha = 9.3, D= 8.1mm. The operculum is oblanceolate, with a terminal nucleus. Radula is typical for the genus, with small central and comb-like lateral teeth. Each marginal tooth is accompanied by a transparent, usually more or less triangular plate at its base (better seen in detached teeth) which is probably a part of radular membrane serving as a tooth ligament. Mean length of marginal teeth is 0.26 mm. The shell of the paratype (H = 19.4 mm) is badly worn and does not differ from the holotype in essential characters. The species is characterized by strongly angled whorls, oblique axials, and, especially, long canal. It resembles the East African Drillia indra Thiele, 1925 while differing in having almost 3 times larger shell with long axials and without a distinct angulation at the shell base. DISTRIBUTION. Zanzibar, 1207—1463 m. Family TURRIDAE H. & A.Adams, 1853 Subfamily CLAVATULINAE Genus TURRICULA Schumacher, 1817 Type species: Turricula flammea Schumacher, 1817 (monotypy) (= Murex tornatus Dillwyn, 1817) Turricula new species Figs 26 & 30 MATERIAL. stn 119, | shell. A single young shell (H = 21.0 mm) apparently represents a new species. It resembles most closely the shallow-water species Turricula catena (Reeve, 1843) distributed in the Red Sea and Persian Gulf, but clearly differs in the absence of the subsutural fold sculptured with axial ribs and in the absence of spiral sculpture on the shell base. In general shell outline it also resembles Surcula fulminata var. gloriosa Melvill, 1917, which Powell (1969) dismissed as an abnormal variant of Turricula tornata fulminata (Kiener, 1839-40), but is distinguished in its heavy nodulose peripheral carina and almost complete absence of spiral ribs except for those on the canal. However it seems A.V. SYSOEV DEEP-SEA CONOIDEAN GASTROPODS reasonable to refrain from formal description of a new species based on a single young shell. DISTRIBUTION. Zanzibar, 1207—1463 m. Subfamily COCHLESPIRINAE Powell, 1942 Genus COCHLESPIRA Conrad, 1865 Type species: Pleurotoma cristata Conrad, 1847 (monotypy) Cochlespira travancorica (E.A.Smith, 1896) Fig. 25 Pleurotoma (Ancistrosyrinx) travancorica E.A.Smith, 1896, p. 368; 1906, p. 163; Alcock & Anderson, 1898, pl. 7, figs 1, la. Pleurotoma (Ancistrosyrinx) travancorica var. granulata E.A.Smith, 1904, p. 459. Ancistrosyrinx travancorica vat. Schepman, 1913, p. 420. Cochlespira travancorica travancorica (E.A.Smith) & C. travancorica travancorica forma granulata (E.A.Smith) — Powell, 1969, p. 396-397, pl. 307. granulata (E.A.Smith) — TYPE LOCALITY. off Travancore (India), 460 fms (travancorica), ‘Investigator’, stn 229, 360 fms (var. granulata). MATERIAL. stn 107, | shell. The JME specimen was determined and described by Powell (1969: 396-397) as C. travancorica travancorica forma granulata. However this specimen differs from the latter variety (in accordance with both original and Schepman’s (1913: 420) descriptions) in the absence of granules on the spiral ribs of the shell base and subsutural slope. The only character in common with the variety granulata, is the presence of weak spiral ribs on the subsutural slope. At the same time, these differences are taxonomically insignificant since the variety granulata, as it was mentioned by Powell, apparently does not warrant even a subspecific status and represents a form of intraspecific variability (this, in particular, makes superfluous the recognition of the nominotypical subspecies by Powell (1969: 396)). DISTRIBUTION. East Africa, India, Indonesia, 338—743 m. Cochlespira zanzibarica Sysoev, new species Figs 23 & 24 MATERIAL. stn 119, | shell (holotype, No. 1993091). DESCRIPTION. The shell is small, light-brown, with glossy surface, rather solid, fusiform, consisting of 6.5 preserved whorls. The protoconch is missing and the upper whorls are heavily eroded. The whorls are angled below the periphery and concave upper and below the angulation; the position of the angulation on the spire whorls shifts upward towards the body whorl. The sutures are straight and very shallowly impressed. The sculpture consists only of a strong median keel with rounded, pointed tubercles (16 on the body and penultimate V/ whorls) and a low ridge on the upper part of the shell base. The ridge is hard to trace above the suture on last spire whorls. The growth lines are very thin. The shell base is angled in its upper part, where the ridge is situated, and slightly and evenly concave below the angulation; it smoothly passes into the canal. The aperture is narrow, with the inner lip weakly and evenly curved. The anal sinus is broad and moderately deep, with the apex situated in the middle of subsutural slope. The canal is straight and rather short (the end is apparently slightly broken). H = 16.9, Hb = 10.2, Ha = 7.6, D=6.9 mm. The new species obviously belongs to the ‘semiplana group’ sensu Powell, 1969 which includes two fossil and two Recent deep-water species and is characterized by the presence of a strong basal keel. The new species differs from all known species of the group by the complete absence of spiral sculpture on the shell base, except for low ridge and in its more stout shell with rather short canal. It is also similar to species of the genus Chesasyrinx Petuch, 1988 known from Miocene of Maryland, USA. Although Petuch (1988: p. 38-39) did not compare Chesasyrinx with Cochlespira in the original description, the striking similarity of shells of Chesasyrinx and ‘semiplana group’ of Cochlespira may be reason for synonymizing these genera. DISTRIBUTION. Zanzibar area, 1207-1463 m. Genus COMITAS Finlay, 1926 Type species: Drillia fusiformis Hutton, 1877 (= Surcula huttoni Suter, 1914) (original designation) Comitas subsuturalis (von Martens, 1902) Figs 31-40 Pleurotoma (Brachytoma) subsuturalis von Martens, 1902, p. 239. Brachytoma subsuturalis (von Martens) — von Martens, 1903 [1904], p. 85, pl. 1, fig. 7. Comitas subsuturalis (von Martens) — Powell, 1969, p. 285, pl. 226, figs 3-4. TYPE LOCALITY. ‘Valdivia’, stn 256, off Somali, 1134 m. MATERIAL. stn 33, 3 specimens and 4 shells; stn 34, 1 specimen and 8 shells; stn 118, 1 specimen and 1 shell; stn 143, 9 specimens; stn 145, 2 specimens; stn 184, 7 specimens and | shell; stn 193, 3 shells. Examination of the large series of JME specimens revealed very a high variability of C. subsuturalis in sculpture and shell proportions. Some specimens are very similar to the original illustration of von Martens (e.g. Figs 37, 38) whilst others, often from the same station, may differ in narrower (or, conversely, broader) shells with more or less high position of peripheral keel, variously differentiated spiral ribs on the shell base, and more or less prominent and numerous tubercles on subsutural fold. The largest JME shell is 32.4 mm in height and 12.0 mm in width. Figs 18-30 Clavusidae, Clavatulinae and Cochlespirinae. 18, 19 — Drillia altispira Sysoev, new species, holotype; 20, 21 — Horaiclavus splendidus (A. Adams, 1867), stn 188 (20) and 176 (21), H = 32.4 (20) and 32.8 (21) mm; 22 — Splendrillia zanzibarica Sysoev, new species, holotype; 23, 24 — Coch- lespira zanzibarica Sysoev, new species, holotype; 25 — Cochlespira travancorica (E.A. Smith, 1896), stn 107, H = 19.1 mm; 26, 30— Turricula new species, stn 119, H = 21.0 mm; 27 — Comitas elegans Sysoev, new species, holotype; 28 — Leucosyrinx claviforma (Kosuge, 1992), stn 158, H = 28.0 mm; 29— Comitas curviplicata Sysoev, new species, holotype. Powell (1969, p. 285) mentioned ‘a related new species from the Gulf of Aden in 1270 metres’, i.e. from stn 184, but did not give a formal description nor reasons for this. However, extreme variants of C. subsuturalis from stn 184 are connected by intermediate forms and can be therefore identified as that species. Specimens from two stations off Maldive Islands comprise a distinct group differing from East African shells in smaller size (15.5—-21.0 mm at 6-7 teleoconch whorls vs. 21.8—28.7 in typical C. subsuturalis) and more slender shell proportions (H/D ratio is 2.82—3.12, mean 2.95 (n = 11) vs. 2.10—2.94 (2.10—2.70 in 19 out of 20 shells measured), mean 2.34). These differences are probably connected with geographical isolation of the Maldive Islands resulting in formation of morphologically isolated population of the species. If the above-mentioned metric differences will be confirmed in additional samples, the population of C. subsuturalis from Maldive Islands should be considered as a distinct subspecies. This population represents a transition (both geographical and conchological) to C. exstructa von Martens, 1903, described from Nicobar Islands. The latter species is distinguished only by an even narrower shell (H/D ratio is 3.43 in the holotype) with longer axial folds (as far as it can be judged from von Martens’ figure). Examination of type material may however reveal that C. exstructais a synonym of C. subsuturalis. A similar statement is probably true for C. obtusigemmata Schepman, 1913, which does not differ from C. subsuturalis in essential conchological characters. DISTRIBUTION. East Africa from the Gulf of Aden to Zanzibar, and Maldive Islands, 494-1789 m. Comitas erica (Thiele, 1925) Fig. 41 Leucosyrinx erica Thiele, 1925, p. 236, pl. 36(24), fig. 25. Comitas erica (Thiele) — Powell, 1969, p. 284, p. 226, fig. 2. TYPE LOCALITY. ‘Valdivia’, stn 191, off Siberut Id. (Sumatra), 750 m. MATERIAL. stn 143, 2 specimens. Also mentioned by Powell (1969) from stn 108 (Zanzibar area, SE of Pemba Island, 786 m). Both specimens are very similar to the original figure and the species variability thus seems to be rather low. One of JME specimens is larger than Thiele’s holotype (20.9 vs. 16.5 mm), but has the appearance of an immature shell. The species was found in the sample also containing C. subsuturalis, but it can be easily distinguished from the latter, by the grayish-white shell with weaker subsutural tubercles, closer-spaced spiral ribs, two of which on the upper shell base are more prominent, and larger protoconch (0.95 mm in diameter vs. 0.7-0.8 mm in C. subsuturalis from the same sample). DISTRIBUTION. Sumatra, 750-797 m. Zanzibar and Maldive Islands, Figs 31-41 A.V. SYSOEV Comitas paupera (Watson, 1881) Figs 42-48 Pleurotoma ( Drillia) paupera Watson, 1881, p. 411. Pleurotoma ( Typhlomangelia) paupera Watson — Watson, 1886, p. 317-319, pl. 25, fig. 3. Turricula paupera Watson — Powell, 1969, p. 244, pl. 202. TYPE LOCALITY. ‘Challenger’, stn 191, off the Arrou Islands (Arafura Sea, Indonesia), 800 fms. MATERIAL. stn 62, 2 specimens; stn 185, 3 specimens and 4 shells. Proper determination of this species is rather intricate due to the very heterogeneous type material. Powell (1969) erroneously described Watson’s syntypes as consisting of two shells with rounded lower whorls and obsolescent sculpture and one shell angulate with strong axially costae. He illustrated the latter specimen and designated it as the holotype (correctly named lectotype in the ‘Measurement’ and “Types’ paragraphs). Actually, one of the two paralectotypes (H = 33.3 mm) has a rounded body whorl with obsolete axials while the axial sculpture on spire whorls is almost the same as in the lectotype (Figs 42 & 43). The second paralectotype (Fig. 44) is represented by a small (H = 18.5 mm) and quite dissimilar shell, characterized by strongly angled whorls with axial sculpture consisting of strong and rounded tubercles at the place of whorl angulation. This specimen is so distinct that one could easily assign it to a separate species if it was not found in the same sample. Two at first glance rather different shells from the JME stn 62 fit however in the range of variability described above. The larger shell (H = 23.4 mm, Fig. 45) is comparable to the lectotype, but has less convex whorls with a less pronounced subsutural slope. The smaller shell (H = 17.5 mm, Fig. 46) is very similar to the smaller paralectotype, but has even stronger peripheral nodules. Small specimens (14-15 mm in height) from stn 185 (Figs 47 & 48) have shells and sculpture intermediate between the extreme variants. In some of the latter specimens, the spiral ribs are more widely spaced on the shell base and reduced in number on the subsutural slope. Thus, C. paupera appears to be very variable species with extreme variants being quite dissimilar to each other but connected by intermediate forms. The species was included by Powell (1969) into the genus Turricula Schumacher, 1817. However its operculum has a terminal nucleus characteristic of the genus Comitas. Surcula suratensis Thiele, 1925 (= Surcula coreanica of von Martens, 1903, not of Adams & Reeve, 1850) is similar to some forms of C. paupera and after examination of type material it may appear to be a synonym of the latter. DISTRIBUTION. Indonesia, Gulf of Aden, and Gulf of Oman, 1463-2000 m. Comitas subsuturalis (von Martens, 1902) (31-40) and C. erica (Thiele, 1925) (41). 31, 32 — stn 184, H = 37.3 mm; 33—stn 184, H = 25.5 mm; 34—stn 184, H = 24.0 mm; 35 - stn 33, H = 21.0 mm; 36 — stn 184, H = 28.6 mm; 37 — stn 34, H = 27.0 mm; 38 — stn 184, H = 21.7 mm; 39—stn 143, H = 21.0 mm; 40 —stn 145, H = 26.2 mm; 41 — stn 143, H = 20.9 mm. DEEP-SEA CONOIDEAN GASTROPODS A.V. SYSOEV Figs 42-48 Comitas paupera (Watson, 1881). 41 —lectotype, H = 40.0 mm; 43, 44 — paralectotypes, H = 33.3 (43) and 18.5 (44) mm, BM(NH) 1887.2.9.1009-11; 45, 46 — stn 62, H = 23.4 (45) and 17.5 (46) mm; 47, 48 — stn 185, H = 14.0 (47) and 14.9 (48) mm. Comitas curviplicata Sysoev, new species Fig. 29 MATERIAL. stn 184, 2 specimens (holotype No. 1993092 and paratype No. 1993093). DESCRIPTION OF HOLOTYPE. The shell medium size, fusiform, rather thin, covered with light-brown periostracum, consists of 7 whorls. The protoconch is missing. The whorls are angled at the periphery or below it (on upper whorls). The subsutural slope is broad and weakly concave. Sutures are slightly wavy, shallow. Axial folds are low, reversed-sigmoid, they form strong rounded nodules below the subsutural slope and then rapidly weaken and become narrowly crested, but extend to the lower suture on the spire whorl and to lower part of the shell base. There are 15 axial folds on the body whorl and 13 on the penultimate. The subsutural slope is devoid of axial sculpture. Growth lines are mostly indistinct except those on the subsutural slope, but some are rough and raised. Spiral ribs are unequal in width and prominence, generally they are most closely spaced in intervals between peripheral nodules and further apart on the lower shell base and the canal. Two ribs on the upper shell base are raised and most prominent. The subsutural slope is covered with subobsolete, closely set ribs which are obsolete on some parts of the shell; there are also two shallow grooves in the middle of subsutural slope. The body whorl is large, occupying about 0.66 of the shell height. The shell base is almost flat, evenly curved in transition to the canal. The aperture is oval, with the inner lip evenly curved and covered by a wide callus which bears oblique folds being the continuation of spiral ribs. The anal sinus is moderately deep, broadly V-shaped, with its apex being in the middle of subsutural slope. The canal is moderately long, slightly turned to left. The operculum is large, leaf-shaped, with terminal nucleus. H = 32.4, Hb = 21.6, Ha = 17.6, D = 13.0 mm. The paratype (H = 29.2, D = 11.8 mm) is similar to the holotype, but has rather worn surface. The new species is similar to C. arcana (E.A.Smith, 1899) from Andaman Islands and Southern India (338-658 m) but differs in | DEEP-SEA CONOIDEAN GASTROPODS having a slightly larger shell with shorter and broader canal, shorter spire (its height is less than that of aperture plus canal), and long axials with characteristic reversed-sigmoid curvature and less strong and more numerous nodules. DISTRIBUTION. Gulf of Aden, 1270 m. Comitas elegans Sysoev, new species Fig. 27 MATERIAL. stn 176, | shell (holotype, No. 1993094). DESCRIPTION. The shell is medium size, fusiform, slender, with a high turreted spire, solid, white under a grayish-brown periostracum, and consists of 8 whorls. The protoconch is missing. Whorls are roundly angled at the shoulder, with conspicuous subsutural fold, slightly concave subsutural slope, and almost vertical lateral sides. Sutures are shallow, wavy, and slightly channelled. Axial folds (12 on the body whorl and 11 on the penultimate one) are strong, oblique, broad, and rounded. They rapidly disappear on subsutural slope and slightly weaken towards the lower suture. On the body whorl, the folds are present on the whorl periphery and fade on the upper part of the shell base. Intervals between folds are narrower than the folds themselves. Spiral ribs are low, broad, rounded and divided by narrow interstices in the upper part of whorls below the subsutural slope. Towards the lower suture and on the shell base they become narrower, thread-like, and more widely spaced. Subsutural slope is smooth except for indistinct spiral grooves in the middle. Growth lines are numerous, thin, prominent on the subsutural slope. The shell base is almost flat; weakly curving, it passes smoothly into a long and straight canal. The aperture is narrow, elongate-oval and not differentiated from the canal. The inner lip is almost straight along most of its length, covered by wide but thin callus. The anal sinus, judging from growth lines, is rather deep, V-shaped, with the apex situated just below the middle of subsutural slope. H = 38.3, Hb = 24.7, Ha = 19.0, D= 11.6mm. The new species is very similar to Drillia investigatoris E.A.Smith, 1899 and differs in-having a much smaller shell (65 mm in the unique holotype of D. investigatoris) with subsutural slope devoid of spiral sculpture. DISTRIBUTION. Gulf of Aden, 655—732 m. Comitas sp. | MATERIAL. stn 184, | shell. | A single broken shell (H = 24.4, upper spire whorls missing) is rather similar to C. trailli(Hutton, 1873) from New Zealand, but the worn condition of the shell hampers its proper identification. | Genus LEUCOSYRINX Dall, 1889 TYPE SPECIES. Pleurotoma verrilli Dall, 1881 designation) (original | Leucosyrinx claviforma (Kosuge, 1992) Fig. 28 Comitas claviforma Kosuge, 1992, p. 165-166, pl. 58, figs 7-8. TYPE LOCALITY. off Port Hedland, Western Australia, 376 m. | MATERIAL. stn 158, 1 shell. 11 The JME shell differs from the unique holotype in having slightly less angled whorls, lower spire (though the shell is smaller: H = 28.0 mm vs. 38.1 mm in the holotype), and reddish brown colour (ashy white in the holotype). However, all other conchological characters agree with original description and illustrations. The species was described as Comitas, but the narrow, turreted shell with angled whorls and the anal sinus scars indicating a deep sinus with greatly projected outer lip are more typical of the genus Leucosyrinx. DISTRIBUTION. Western Australia and Maldive Islands, 376-1170 m. Subfamily CRASSISPIRINAE Morrison, 1966 Genus INQUISITOR Hedley, 1918 Type species: Pleurotoma sterrha Watson, designation) 1881 (original Inquisitor nodicostatus Kilburn, 1988 Fig. 49 Crassispira aesopus (non Schepman, 1913) — Kilburn, 1973, p. 572, fig. 13a. Inquisitor nodicostatus Kilburn, 1988, p. 259-261, figs 36, 42, 213-214. TYPE LOCALITY. Natal, 164-169 fms. South Africa (29°43'S, 31°05'E), MATERIAL. stn 106, | shell. DISTRIBUTION. South Africa and Zanzibar, 183—310 m. Inquisitor adenicus Sysoev, new species Figs 2 & 50-53 MATERIAL. stn 33, 1 paratype No. 1993096/1; stn 34, 7 paratypes No. 1993096/2-8; stn 193, holotype No. 1993095 and 4 paratypes No. 1993096/9-12. DESCRIPTION OF HOLOTYPE. The shell is medium sized for the genus, claviform, thin but solid, with a high spire, covered with olivaceous-brown periostracum, and consisting of 8 whorls. The protoconch is missing and the upper whorls are eroded. Whorls are obtusely angled at the shoulder and moderately convex. Subsutural slope is broad and concave and the subsutural fold is weak and indistinct. Sutures are shallow and wavy. Axial folds are strong and rounded, slightly oblique, with interstices narrower than folds. Folds extend from the lower suture to the lower part of subsutural slope where they rapidly disappear. There are ten folds on each of two last whorls. Growth lines are thin, prominent on the subsutural slope. Spiral cords are strong, rounded, almost equally developed on axial folds and in interstices; they are much narrower than the interspaces between them. The cords are absent on subsutural slope and become narrower, closer and much weaker on lower part of the shell base towards the canal end. There are four cords on the spire whorls (five on the penultimate one, the lowest cord submargins the suture), five on the body whorl periphery, and, below a wider interval, about 12 on the shell base and canal. The aperture is elongate-oval, rather narrow, becoming slightly narrower as it passing into the canal. Outer lip with thin edge; inner lip 12 concave, covered with wide, longitudinally rugose callus, which does not form a pad at the sinus entrance. The anal sinus is deep, narrows markedly towards the apex, U-shaped (type (c) of Kilburn, 1988), and its deepest point is situated slightly below the middle of subsutural slope. The canal is moderately long and straight. H = 33.0, Hb = 18.4, Ha = 13.8, D= 10.3 mm. The paratypes vary slightly in the character of spiral sculpture: the cords may be unevenly spaced, rarely with thinner additional cords in some intervals. The subsutural fold is variously developed, sometimes it is distinct. The canal is narrower and sometimes curved in smaller shells and broader and therefore visually shorter in larger paratypes. The largest paratype is 34.3 mm in height. In the only paratype with an intact protoconch, it consists of 1.5 rapidly increasing globose whorls (1.0 mm in diameter) with a smooth surface. An operculum was not found; it might be lost in the dried animal. The radula (Fig. 2) is typical of the genus, teeth with tapering distal end of the shaft, poorly differentiated cutting edge, and large and broad accessory limb. The tooth length is 0.17 mm (in paratype from stn 33, H = 34.3 mm). The new species resembles Funa laterculoides (Barnard, 1958) in general outlines but clearly differs in its protoconch, colour, radula, and details of sculpture. DISTRIBUTION. Gulf of Aden, 1022-1295 m. Inquisitor indistinctus Sysoev, new species Figs 3, 11 & 54-55 MATERIAL. stn 145, 3 specimens and | shell (holotype No. 1993097 and 3 paratypes No. 1993098). DESCRIPTION OF HOLOTYPE. The shell is medium size, claviform, slender, with rather high spire, thin but solid, covered with solid olivaceous periostracum, and consists of protoconch and 9 teleoconch whorls. The protoconch is small (0.95 mm in diameter) consisting of 1.5 smooth glossy whorls. Definitive whorls weakly convex, with a slight angulation at the shoulder. Subsutural slope concave except for weak subsutural fold. Sutures straight, moderately deep, become slightly channelled on the body whorl. The axial sculpture is represented by strong oblique rounded folds, gradually disappearing on subsutural slope and reaching the lower suture or, on the body whorl, the shell base. The folds tend to weaken on the last half of the body whorl. There are 14 folds on the body and penultimate whorls. Spiral cords override the axial folds, they are low, wide and rounded, the intervals are approximately equal to cords in width or somewhat wider. There are 16 cords on the body whorl and 7-8 on the penultimate. The shell base is weakly convex, and smoothly passes into the canal. The aperture is rather narrow, elongate-oval, and not differentiated from the canal. The latter is short and wide, somewhat expanded and shallowly notched at the end. The anal sinus is moderately deep, rounded, symmetrical, and occupies the entire subsutural slope. The inner lip is covered with thick white callus with a free edge in the lower part. Columella almost straight. H = 31.3, Hb = 16.5, Ha = 12.4, D=8.3 mm. The paratypes are smaller (H no more than 27.4 mm) and vary slightly in the prominence of the axial and spiral sculpture A.V. SYSOEV including the subsutural fold. In one paratype, there is a weak but distinct callus pad at the entrance to anal sinus. The operculum is oblanceolate, with terminal nucleus. Marginal teeth of the radula possess broad leaf-shaped accessory limb, which does not reach the distal end of the shaft. The mean tooth length is 0.14 mm (in paratype with H = 27.4 mm). DISTRIBUTION. Maldive Islands, 494 m. Inquisitor stenos Sysoev, new species Figs 56 & 57 MATERIAL. stn 176, | shell (holotype, No. 1993099). DESCRIPTION. The shell is rather small, slender, with high spire comprising about 0.5 of the shell height, yellowish-white, and consisting of 9 whorls. The protoconch is missing. Whorls are strongly angled at the periphery, and the whorl profile is very weakly concave above the angulation and almost flat below it. There is a narrow and weak subsutural fold. Sutures are straight and moderately deep. The axial sculpture consists of slightly oblique and widely spaced folds (9 on each of two last whorls) forming longitudinally elongated and sometimes pointed strong tubercles in the middle of the whorl. On early spire whorls, the folds are obsolete but visible on the lower part of subsutural slope and reach the lower suture. Towards the body whorl, they become obsolete near the lower suture and over most part of subsutural slope, but again become longer and extend over entire shell base in the last half of the body whorl. The last fold situated behind the aperture edge is much wider and stronger than other. Spiral sculpture consists of indistinct, rather broad ribs which are obsolete or subobsolete in interstices between axial folds and absent on the subsutural slope. The aperture is narrow, elongate-oval, with a thick labial callus and straight columella. The labrum has a thin edge and low and narrow fold-like varix behind the edge. The anal sinus is moderately deep, U-shaped, with slightly constricted entrance (type (b) of Kilburn, 1988). There is a moderately large, pointed, and outwardly projecting parietal tubercle. H = 21.3, Hb = 10.7, Ha = 18.4, D= 5.7 mm. The new species 1s distinguished by its small narrow shell with high spire, broad subsutural slope, low but strongly tuberculated at the periphery axial folds, and obscure spiral ribs. DISTRIBUTION. Gulf of Aden, 655-732 m. Inquisitor angustiliratus Sysoev, new species Fig. 58 MATERIAL. stn 188, | shell (holotype, No. 1993100). DESCRIPTION. The shell is moderately large, with a rather high spire, strong, yellowish-gray, and consisting of 8 whorls. The protoconch is missing. Whorls are moderately convex, slightly angled at the shoulder, with indistinct subsutural fold and moderately wide (about 1/3 of the whorl height), concave subsutural slope. Sutures are wavy, and slightly channelled. Axial folds (12 on the body whorl and 11 on the penultimate one) are narrow, slightly oblique, with sharpened crests, and long (they reach the lower suture and extend over the entire shell base). The folds begin in the lower part of the subsutural slope Figs 49-62 Crassispirinae. 49 — Inquisitor nodicostatus Kilburn, 1988, stn 106, H = 36.8 mm; 50-53 — I. adenicus Sysoev, new species, holotype (50) and paratypes, stn 33 (51) and 34 (52-53), H = 34.3 (51), 30.9 (52) and 26.3 (53) mm; 54, 55 — I. indistinctus Sysoev, new species, holotype (54) and paratype, stn 145, H = 26.0 mm (55); 56, 57 — 1. stenos Sysoev, new species, holotype; 58 — 1. angustilirata Sysoev, new species, holotype; 59, 60 — Paradrillia agalma (E.A. Smith, = 1906), stn 176, H = 15.3 mm; 61 — P. agalma?, stn 180, H = 14.4 mm; 62 — Ceritoturris sp., stn 42, H = 11.8 mm. DEEP-SEA CONOIDEAN GASTROPODS 14 and are most prominent in the whorl periphery. Spiral ribs are strong, cord-like, narrow, widely spaced (but become progressively closer to each other and lower on the shell base and canal); they override the axial folds and form nodules at intersections. Interspaces between ribs are very finely spirally striate and sometimes bear a thin thread. The number of ribs increases from 3 to 5 in successive spire whorls. The subsutural fold is sculptured by two thin spiral riblets. Subsutural slope is smooth except for very fine spiral striations and 3-4 thin threads in the lower half. Growth lines are numerous, thin, and clear, on the body whorl some of them are rough and raised. The shell base is almost flat and not differentiated from the canal. The aperture seems to be rather wide (the outer lip is broken). The inner lip is weakly and evenly curved, and covered by thick callus. The parietal callus pad is very weak. The anal sinus (judging from its scars) is moderately deep, openly U-shaped (type (c) of Kilburn, 1988). The canal is broad and widely open. H = 39.5, Hb = 21.4, Ha = 16.5, D= 10.8 mm. The new species resembles [ coxi (Angas, 1867) from south-eastern Australia in whorl outline and the character of sculpture but clearly differs in the shell proportions. It also looks somewhat like an extremely stretched out specimen of /. crassa (E.A.Smith, 1888). DISTRIBUTION. Gulf of Aden, 528 m. Genus PARADRILLIA Makiyama, 1940 Type species: Drillia dainichiensis Yokoyama, 1923 (original designation) Paradrillia agalma (E.A.Smith, 1906) Figs 59-61 Pleurotoma (Surcula) agalma E.A.Smith, 1906, p. 162-163; Annandale & Stewart, 1909, pl. 21, figs 4, 4a. Paradrillia agalma (E.A.Smith) — Powell, 1969, p. 317, pl. 246, figs 1, 2. TYPE LOCALITY. ‘Investigator’, stn 269, West of Cape Comorin (SE India), 464 fms. MATERIAL. stn 176, | shell; stn 180, 2 shells. The shell from stn 176 is quite typical. It consists of protoconch and 7 teleoconch whorls. The previously undescribed protoconch is basically similar to that of P melvilli figured by Powell (1969, pl. 242, fig. 2). It consists of 2.5 whorls with smooth glossy surface, the tip is small and papillate, and a thin low-set keel is developed on the last whorl. There are 17 axial ribs on the body whorl and 15 on the penultimate one. The shells from stn 180 (Fig. 61) were referred to P agalma with some doubts. They differ in more numerous peripheral tubercles, weaker axial and spiral sculpture, and in the presence of two spiral threads on the subsutural fold and two riblets below the periphery on the spire whorls. They seem to be a transition to P. melvilli Powell, 1969 in sculpture, but that species is almost half the size, with a proportionally higher spire and truncated anterior end. However, the general pattern of sculpture, shell outline and the character of protoconch are similar in all JME shells and those from stn 180 are provisionally assigned to P agalma pending examination of additional material. DISTRIBUTION. Ceylon and Gulf of Aden, 655-848 m. A.V. SYSOEV Genus CERITOTURRIS Dall, 1924 Type species: Ceritoturris bittium Dall, designation) 1924 (original ?Ceritoturris sp. Fig. 62 MATERIAL. stn 42, | shell. A single heavily damaged shell (H = 11.8 mm) from stn 42 can possibly be referred to Ceritoturris on the basis of its resemblance to both type species and C. nataliae Kilburn, 1988. The very bad condition of the shell renders more precise identification impossible. DISTRIBUTION. West Arabian Sea, 1415 m. Genus PTYCHOBELA Thiele, 1925 Type species: Clavatula crenularis Lamarck, 1816 (= Murex nodulosus Gmelin, 1791) (original designation) Ptychobela cf. suturalis (Gray, 1838) Figs 79 & 80 Drillia suturalis Gray, 1838, p. 29. Ptychobela suturalis (Gray) — Kilburn, 1989, p. 190, figs 5-6 (holotype) & 7-8. TYPE LOCALITY. unknown. MATERIAL. stn 35, | shell; stn 188, 2 shells. These shells are difficult to determine primarily due to the existence of numerous species of uncertain status which have been described within the genus Drillia and, as far as it can be judged from drawings (when present) and rather brief descriptions, are similar to the JME material (e.g. Drillia incerta E.A.Smith, 1877, D. atkinsoni E.A.Smith, 1877, D. variabilis E.A.Smith, 1877, etc.). The question cannot be resolved without comparative examination of type material. Nevertheless, these shells are quite comparable with the holotype of P suturalis illustrated by Kilburn (1989). The latter species is however characterized by usually very short axial folds with strong peripheral nodules (but the folds in the holotype are rather long — see Kilburn, 1989, fig. 6). The smaller of the JME shells (stn 188, H = 22.7, D = 7.6 mm, Fig. 80) differs also in complete absence of additional spiral threads in the interspaces between main cords. The other shell (stn 188, H = 22.9, Hb = 15.6 mm, upper spire whorls are broken off and only 3.5 last whorls are intact) also lacks additional threads on upper spire whorls, but they appear on penultimate whorl and become rather strong on the body whorl. The anal sinus is similar in shape to that of P suturalis, but some of the sinus scars indicate that during the shell growth the sinus may be quite different: very deep and narrow, asymmetrical, with the upper edge of the slit almost parallel to the suture. The rather large shell from stn 35 (H = 31.1, D= 11.0 mm, Fig. 79) is characterized by fairly long axials reaching the lower suture and a peculiar spiral sculpture of ribs with the upper (directed adapically) slope being much steeper than the lower. This produces a somewhat tiled pattern and, when illuminated from the shell apex, the sculpture seems to consist of very wide flattened ribs. | excavated DEEP-SEA CONOIDEAN GASTROPODS DISTRIBUTION. According to Kilburn (1989), P suturalis is a central West Pacific species (from Taiwan and Singapore to Queensland and Western Australia). Ptychobela cf. nodulosa (Gmelin, 1791) Fig. 84 Murex nodulosus Gmelin, 1791, p. 3562. Clavatula crenularis Lamarck, 1816, p. 9, pl. 440, figs 3a,b. Ptychobela nodulosa (Gmelin) — Kilburn, 1989, p. 187-190, figs 1-2) & 3-4 (neotype). TYPE LOCALITY. unknown. MATERIAL. stn 188, | shell. The situation with this species is the same as in the preceding case. The JME shell resembles rather closely the neotype of P nodulosa designated and illustrated by Kilburn (1989) in the shell outline and the character of sculpture. However, it lacks the characteristic colour pattern of P nodulosa being of uniform light-brown colour (the shell was dead collected and probably faded), and has a slightly higher spire (Hs/H = 0.49 vs. 0.41 in the neotype of P nodulosa) and more convex whorls. A characteristic feature of the shell considered is that axial folds extend from suture to suture on the uppermost whorls and only on 8th teleoconch whorl the typical subsutural slope without axial sculpture is developed. Subfamily TURRINAE H. & A.Adams, 1953 Genus GEMMULA Weinkauff, 1875 Type species: Pleurotoma gemmata Reeve, 1843 (subsequent designation Cossmann, 1896) (= Gemmula hindsiana Berry, 1958) Gemmula (Gemmula) vagata (E.A.Smith, 1895) _ Figs 63-65 Pleurotoma vagata E.A.Smith, 1895, p. 3, pl. 1, fig. 3; 1904, p. 456; Alcock et al., 1907, pl. 14, figs 3, 3a. | Gemmula vagata (E.A.Smith) — Powell, 1964, p. 258-259, pl. 196, fig. 10. | TYPE LOCALITY. ‘Investigator’, stn 172, off Trincomalee, | Ceylon, 200-350 fms. | MATERIAL. stn 176, 8 shells, stn 188, 3 shells. The largest shell is 49.8 mm in height (apex slightly broken). Examination of a growth series showed that its characteristic features (i.e. almost vertical sides of whorls, very strongly subsutural fold, and channelled sutures) are developed only when the shell reaches a certain size _ (approximately 35 mm in height and more than 10 teleoconch whorls). Young shells can be determined only by comparison with larger specimens. A peculiar and previously undescribed feature of the species is the presence of spiral lirae inside the aperture in large individuals. DISTRIBUTION. Gulf of Aden to Andaman Islands, 338-1061 m. Gemmula (Gemmula) amabilis (Jickeli in Weinkauff, 1875) Figs 66, 67 & 71 Pleurotoma amabilis Jickeli in Weinkauff, 1875, p. 29, pl. 6, figs 4, 6. Pleurotoma (Gemmula) amabilis Weinkauff — Sturany, 1903, pl. 3, figs 3a-c. Gemmula amabilis (Weinkauff) — Powell, 1964, p. 261-262, pl. 200, fig. 1, pl. 201, figs 3-7. ?Gemmula amabilis (Weinkauff) — Kosuge, 1990, p. 153-154, pl. 55, fig. 13, text-fig. 6; Kosuge, 1992, p. 163, pl. 58, fig. 1, text-figs 7, 11, 12-14. TYPE LOCALITY. Massawa (Ethiopia), Red Sea. MATERIAL. stn 34, 18 specimens and shells; stn 119, 1 shell; stn 145, 1 specimen and 2 shells; stn 176, 2 shells; stn 185, 1 shell; stn 188, 2 shells; stn 193, 3 specimens. The taxonomy of this species is rather confused. The name amabilis had long been unused until Powell (1964) applied it to shells from the JME material (the specimen from the JME stn 176 figured by him (Powell, 1964, p. 200, fig. 1) was not found). However, Powell expressed some doubts on the identification of JME specimens as G amabilis, having indicated certain differences from the sketchy and rather inadequate original figures. Kilburn (1983) reported that the types of Pleurotoma amabilis were probably lost and therefore the name amabilis must remain a nomen dubium. He also compared the specimen figured by Powell with G. pulchella Shuto, 1961 from the Pliocene of Japan. The latter species is characterized by smaller and much more numerous gemmules (29 on the body whorl of 10.2 mm holotype and up to 34 at the shell height of 26.57 mm — Shuto, 1965). Later on, Kosuge (1990, 1992) used the name amabilis for North-Western Australian shells and at the same time mentioned that Powell’s specimen differs from the original drawings and, as far as it can be judged from the text (Kosuge, 1990, p. 154), may not be conspecific with amabilis. The shells figured by Kosuge as G amabilis, actually resemble G. gemmulina (von Martens, 1902) sensu Powell, 1964 more in the shell outlines and the character of spiral sculpture of alternating primary cords and intermediate threads. The latter species is distributed from Taiwan to Indonesia (Powell, 1964), which is far closer to North-Western Australia than the Red Sea and the Gulf of Aden. Despite the obviously ambiguos status of the name amabilis, it seems reasonable to conserve its application to the Gulf of Aden shells similar to those described by Powell, because no other available name for them exists. Perhaps, the examination of a large series of the Red Sea Gemmula, especially from the type locality of amabilis, would clear up the question of proper application of the name. The specimens of G amabilis are characterized by a small slender shell (up to 25.9 mm, usually 22-25 mm) with more or less channelled sutures, a moderately developed subsutural fold covered by 1-3 riblets, 2-3 prominent and widely spaced cords on the upper shell base without intermediate threads, few (usually two) thin threads on the subsutural slope, and 16—24 peripheral gemmae on the body whorl (usually 22, mean 20.7 at the shell height 18.2-25.9 mm; the number of gemmae does not show a strict correlation with the shell height). This deep-water species, due to its small size, can be confused with young specimens of other species of Gemmula. However, the presence of the ‘tertiary’ apertural DEEP-SEA CONOIDEAN GASTROPODS notch (characteristic for mature individuals of Gemmula — see Kantor & Sysoev, 1991) in one of the shells (H = 20.2) indicates that the mentioned shell size characterizes adult specimens of G; amabilis. DISTRIBUTION. North-western Indian Ocean, 494-2000 m; also probably from north-western Australia, 300-496 m. Gemmula (Gemmula) cf. congener (E.A.Smith, 1894) Pleurotoma congener E.A.Smith, 1894, p. 160-161, pl. 3, figs 4, 5. Gemmula congener subspecies congener (E.A.Smith) — Powell, 1964, p. 251-252, p. 191, figs 1-4; Cernohorsky, 1987, p. 123-124, figs 1, 2-3 (holotype), 4—S. TYPE LOCALITY. Bay of Bengal, 128 m. MATERIAL. stn 176, | shell. A single broken and heavily worn shell (H = 39.6 mm) can probably be referred to G congener by its very strong gemmulated subsutural fold. DISTRIBUTION. Indian Ocean, 198—732 m. Subgenus UVEDOGEMMULA MacNeil, 1960 Type species: Pleurotoma unedo Kiener, 1839-40 (original designation) , Gemmula (Unedogemmula) unedo (Kiener, 1839-1840) Fig. 75 | Pleurotoma unedo Kiener, 1839-1840, p. 19, pl. 14, fig. 1. , Gemmula (Unedogemmula) unedo (Kiener) — Powell, 1964, p. 269-270, pl. 175, figs 1, 6, pl. 208, figs 1, 2; Kosuge, 1988, p. 121-122, text-figs 4, 13-15, pl. 47, figs 9, 10. Pleurotoma invicta Melvill, 1910, p. 15, pl. 2, fig. 27. | TYPE LOCALITY. ‘Mers de Il’Inde’ (unedo), Persian Gulf (invicta). MATERIAL. stn 145, | specimen. The shell from stn 145 is peculiar in the complete absence of spiral sculpture on the body whorl. The sculpture is represented only by very rough growth lines which is probably a senile abnormality. DISTRIBUTION. Persian Gulf to Japan, 73-503 m. Subgenus PTYCHOSYRINX Thiele, 1925 Type species: Pleurotoma (Subulata) bisinuata von Martens, 1901 (original designation) Figs 63-75 17 Gemmula (Ptychosyrinx ) bisinuata (von Martens, 1901) Figs 72-74 Pleurotoma (Subulata) bisinuata von Martens, 1901, p. 17. Drillia (Subulata) bisinuata (von Martens) — von Martens, 1903 [1904], p. 82, pl. 1, fig. 8. Ptychosyrinx bisinuata (von Martens) — Thiele, 1925, p. 176(210), text-fig. 28 (rad.), pl. 46(34), fig. 28 (operc.); Powell, 1964, p. 289-290, pl. 223, figs 1,2. Gemmula_ (Ptychosyrinx) bisinuata Cernohorsky, 1987, p. 130, figs 15-17. (von Martens) — TYPE LOCALITY. ‘Valdivia’, stn 264, near the coast of Somalia, 1079 m. MATERIAL. stn 119, 1 specimen and | shell; stn 184, 1 specimen. The species is rather similar to the closely related G teschi (Powell) (see below). The main differences are that, in G bisinuata, the subsutural rib is clear, thin, straight or slightly wavy, without nodules; the sutures are poorly seen and very shallowly impressed. In G. teschi, the subsutural rib is very weakly developed or absent (or there is a weak to moderate subsutural fold), covered with nodules which are the continuation of axial folds; the sutures are clear, more or less channelled. Additionally, the spiral ribs on the shell base in G. bisinuata are clear and prominent; two lower ribs out of three upper ones are much stronger than the other. In G. teschi, the ribs are thinner, more uniform, more numerous and closely spaced; the shell base is evenly convex. DISTRIBUTION. East Africa from the Gulf of Aden to Malagasy, 818-1463 m. Gemmula (Ptychosyrinx) teschi (Powell, 1964) Figs 68-70 Ptychosyrinx timorensis teschi Powell, 1964, p. 291-292, pl. 223, figs 5, 6; Abbott & Dance, 1990, p. 238, fig. (holotype). TYPE LOCALITY. ‘Albatross’, stn 5587, NW of Sipadan Id., Borneo, 415 fms. MATERIAL. stn 118, 6 specimens. Although having been described as a subspecies, G teschi obviously warrants specific rank from the fossil G timorensis (Tesch, 1915) in having a much broader fusiform shell. The specimens from the JME material vary in the prominence of spiral ribs on the shell base and the number of peripheral tubercles (17—22 on the body whorl, mean 19). In no specimen is the subsutural fold as strong and regularly gemmate as in the holotype. DISTRIBUTION. Indonesia and Zanzibar, 635-1789 m. Gemmula spp. 63-65 — Gemmula (Gemmula) vagata (E.A. Smith, 1895), stn 176 (63, 64) and 188 (65), H = 48.9 (63), 33.3 (64) and 36.3 (65) mm; 66, 67, 71— G. (G.) amabilis (Jickeli in Weinkauff, 1875), stn 193 (66), 34 (67) and 145 (71), H = 25.8 (66), 23.3 (67) and 21.3 (71) mm; | 68-70— Gemmula (Ptychosyrinx ) teschi (Powell, 1964), stn 118, H = 28.2 (68), 27.5 (69) and 24.7 (70) mm; 72-74 — G. (P.) bisinuata (von Martens, 1901), stn 119 (72) and 184 (73, 74), H = 18.8 (72) and 30.5 (73, 74) mm; 75 — Gemmula ( Unedogemmula) unedo (Kiener, 1839-40), stn 145, H = 95.5 mm. A.V. SYSOEV i : f DEEP-SEA CONOIDEAN GASTROPODS Genus LUCERAPEX Iredale, 1936 Type species: Pleurotoma casearia Hedley & Petterd, 1906 (original designation) Lucerapex adenica Powell, 1964 Fig. 76 Lucerapex adenica Powell, 1964, p. 286-287, pl. 221, fig. 3. TYPE LOCALITY. ‘Mabahiss’ (John Murray Expedition), stn 34, Gulf of Aden, 1022 m. MATERIAL. stn 193, 1 shell; also described from stn 34 and 191 by Powell (1964). The specimen figured (H = 31.0 mm which is slightly more than in the type specimens) was probably omitted by Powell whose original decription of the species was based on the JME material. DISTRIBUTION. Gulf of Aden, 274-1080 m. Lucerapex molengraaffi (Tesch, 1915) Figs 77 & 78 Pleurotoma (s.str.) molengraaffi Tesch, 1915, p. 28, pl. 77, figs 54-56. Lucerapex molengraaffi (Tesch) — Powell, 1964, p. 287-288, pl. 220, figs 3, 4, pl. 221, figs 1, 2. TYPE LOCALITY. Timor, Pliocene. MATERIAL. stn 145, 1 shell. DISTRIBUTION. Maldive Islands, Borneo, Celebes, Phillipines, 464-1022 m. Family CONIDAE Fleming, 1822 Subfamily CLATHURELLINAE H. & A.Adams, 1858 Genus BORSONIA Bellardi, 1839 Type species: Borsonia prima Bellardi, 1839 (monotypy) | Borsonia ochracea Thiele, 1925 | Figs 82 & 83 Borsonia ochracea Thiele, 1925, p. 183-184 (217-218), pl. 38(26), figs 1-3, text fig. 26. TYPE LOCALITY. ‘Valdivia’, stn 257, (off Somalia), 1644 m. | MATERIAL. stn 122, 1 specimen; stn 184, | shell. The JME material agrees well with Thiele’s original figures | differing only in having somewhat narrower and more widely spaced spiral ribs (though this may result from rather | schematized appearance of spiral sculpture in many Thiele’s | figures). The columellar pleat is very weak and can be seen only 19 if the aperture is broken. The radular teeth are also very similar to those figured by Thiele; the operculum is absent as in the type specimens. DISTRIBUTION. East Africa from Zanzibar to the Gulf of Aden, 693-1644 m. Subgenus CORDIERIA Rouault, 1848 Type species: Cordieria iberica Rouault, 1850 (subsequent designation Cossmann, 1896) Borsonia (Cordieria) symbiophora Sysoev, new species Figs 5, 12, 13, 81 & 85 MATERIAL. stn 26, 4 specimens (paratypes No. 1993103); stn 118, 6 specimens (holotype No. 1993101 and 5 paratypes No. 1993102). DESCRIPTION OF HOLOTYPE. The shell is of medium size for the genus, broadly fusiform, rather stout, strong, white under olivaceous periostracum and light-brown inside the aperture, and consisting of 7 whorls. The protoconch is missing, and the upper whorls are eroded. Definitive whorls are obtusely angled at the periphery, slightly concave at the subsutural slope and weakly convex below. The body whorl is rather large (0.65 of the shell height), the shell base is weakly convex, and passes smoothly into the canal. The sutures are wavy and channelled. The axial sculpture consists of wide, rounded, short folds which are obsolete in the middle part of the subsutural slope (they can be traced as very low tubercles just below the suture) and do not reach the shell base. The folds are most prominent just below the subsutural slope at the whorl periphery. There are 12 folds on the body whorl and 11 on the penultimate. Spiral sculpture is represented by wide, rounded, low ribs separated by narrow grooves and covering the entire shell surface. On the shell base, the ribs become subobsolete, with wider interspaces. The growth lines override the ribs making their surface rugose. The aperture is rather wide and not differentiated from the wide and short canal. The inner lip is smooth, covered by glossy callus. The anal sinus is symmetrical, wide and shallow, its deepest part is situated in the middle of the subsutural slope. H = 27.4, Hb = 17.8, Ha = 13.9, D= 11.5 mm. The paratypes are very variable in the character of spiral sculpture which may be either well developed or subobsolete to obsolete. There is no correlation between the prominence of spiral ribs on the subsutural slope and on the rest part of the whorl. Paratypes of smaller size have more biconic shells with narrower canals. The protoconch is broken in almost all specimens. In the only specimen with an intact but eroded protoconch, it seems to consist of 1.5 rapidly increasing whorls. The radular teeth are typical of borsoniid group of genera, small (0.24 mm in paratype from stn 185, H = 21.8 mm), straight, rather short, with a cusp at the tooth base. The operculum is small, leaf-shaped, vestigial, with a terminal nucleus. In one paratype from stn 185 the operculum, probably | Figs 76-89 Turrinae, Zonulispirinae and Clathurellinae. 76 — Lucerapex adenica Powell, 1964, stn 193, H = 31.0 mm; 77, 78 — L. molengraaffi | (Tesch, 1915), stn 145, H = 25.1 mm; 79, 80 — Ptychobela cf. suturalis (Gray, 1838), stn 35 (79) and 188 (80), H = 31.1 (79) and 22.7 (80) mm; 81, 85 — Borsonia ( Cordieria) symbiophora Sysoev, new species, holotype (81) and paratype, stn 118, H = 23.8 mm (85); 82, 83 — Borsonia ( Borsonia) ochra- cea Thiele, 1925, stn 122, H = 37.0 mm; 84 — Ptychobela cf. nodulosa (Gmelin, 1791), stn 188, H = 29.6 mm; 86-88 — Typhlomangelia maldivica Sysoev, new species, holotype (86) and paratype, stn 143, H = 32.0 mm (87, 88); 89 — T. adenica Sysoev, new species, holotype. 20 as a result of damage and subsequent repair, is very small, subquadrate, with central nucleus (Fig. 12). All specimens of B. symbiophora bear actinians on their shells and are often entirely covered with them. The new species differs from all known Recent species of Borsonia, in its stout shell with typically uniform spiral sculpture and smooth columella. The species is quite comparable with species of Borsonia and, especially of the Cordieria subgenus primarily differing in the absence of columellar plicae and the presence of operculum. However, the prominence (and even presence) of columellar plicae can vary greatly among species of the same genus and sometimes among shells of the same species. The presence of an operculum also cannot be considered as a diagnostic character because it is very patchily distributed in the subfamily, where repeated and independent reduction and loss of operculum undoubtedly occurred, and many faunas demonstrate a full range of species with well developed, vestigial or missing operculum (e.g. Eastern Pacific — see McLean, 1971). B. symbiophora is also similar to species of the subgenus Borsonellopsis McLean, 1971 of the genus Borsonella Dall, 1890. The type species of Borsonellopsis, Leucosyrinx erosina Dall, 1908, possesses similar sculpture and shell outlines as well as a vestigial operculum, and lacks columellar plicae. On the other hand, it differs considerably from Borsonella s.str. and may not be congeneric. DISTRIBUTION. Gulf of Aden and off Mombasa (Kenya), 1789-2312 m. Genus BATH YTOMA Harris & Burrows, 1891 Type species: Murex cataphractus Brocchi, 1814 (monotypy) Subgenus PARA BATHYTOMA Shuto, 1961 Type species: Pleurotoma striatotuberculata Yokoyama, 1928 (original designation) The differences between the generally accepted subgenera of Bathytoma (see Powell, 1966) seem to be rather slight. Kilburn (1986) mentioned that Parabathytoma differs from Micantapex in having radular teeth without an elongate base and in the absence of brephic arcuate riblets at the place of the protoconch transition into teleoconch whorls. However, in all the species described below, the presence of long, curved teeth without an elongate base is associated with the presence of arcuate riblets at the place of the protoconch termination, i.e. these species possess characters of both Parabathytoma and Micantapex sensu Kilburn. Thus, the only feature distinguishing these subgenera is the shape of radular teeth. The question is further complicated by the fact that the type species of Parabathytoma is a fossil Pleurotoma striatotuberculata Yokoyama, 1928, while the radula of type species of Micantapex, Bathytoma agnata Hedley & Petterd, 1906, is unknown. Nevertheless, the species listed below are provisionally included into Parabathytoma on the basis of their radular morphology. Figs 90-101 A.V. SYSOEV Bathytoma ( Parabathytoma) prodicia Kilburn, 1986 Figs 90 & 91 Bathytoma (Parabathytoma) regnans (non Melvill, 1918) — Kilburn, 1971, p. 31, figs 2c, 2f, 4b. Bathytoma ( Parabathytoma) prodicia Kilburn, 1986, p. 643, figs 22-23. TYPE LOCALITY. East of Bazaruto Island Mozambique), 300-310 fms. (Southern MATERIAL. stn 119, 1 shell. The shell from the JME material differs from the holotype figured by Kilburn in having less prominent peripheral nodules, shallower anal sinus, and flattened shell base. However, all other essential conchological characters including the shell proportions (D/H = 0.43, Ha/H = 0.52) are similar to B. prodicia. An additional smaller (H = 23.5 mm) specimen collected off Zanzibar by R/V ‘Vityaz’ (stn 4680, 740 m) is in some respects intermediate between the typical B. prodicia and the JME shell. DISTRIBUTION. Southern Mozambique to Zanzibar, 420-1463 m. Bathytoma (Parabathytoma) oldhami (E.A.Smith, 1899) Figs 7 & 92-93 Pleurotoma ( Bathytoma) oldhami E.A.Smith, 1899, p. 238. Pleurotoma oldhami E.A.Smith— Alcock & McArdle, 1901, pl. 9, figs 2, 2a. TYPE LOCALITY. ‘Investigator’, stn 229, off Travancore coast, 360 fms. MATERIAL. stn 145, 1 specimen. The JME specimen is of approximately the same size as the holotype and very similar to the figure of the latter, differing only in slightly broader shell (H = 41.0 mm, D = 15.8 mm vs. 43 and 15 mm in the holotype). The protoconch consists of about 1.5 smooth globose whorls followed by several arcuate axial riblets which gradually become stronger and pass into the teleoconch sculpture. The radular teeth (Fig. 7) are long and strongly curved, of typical shape for the subgenus. The mean tooth lengtiris 0.77 mm. DISTRIBUTION. Southern India and Maldive Islands, 494-658 m. Bathytoma ( Parabathytoma) regnans Melvill, 1918 Figs 8, 16 & 94-97 Bathytoma regnans Melvill, 1918, p. 68, textfig. Bathytoma regnans Melvill — Kilburn, 1986, p. 718, fig. 168 (holotype). TYPE LOCALITY. Indian Ocean, (probably the Bay of Bengal). Investigator Expedition MATERIAL. stn 34, | specimen and 5 shells; stn 188, 2 shells; stn 193, 6 shells. Clathurellinae. 90, 91 — Bathytoma ( Parabathytoma) prodicia Kilburn, 1986, stn 119, H = 32.3 mm; 92, 93— B. (P.) oldhami(E.A. Smith, 1899), stn 145, H = 41.0 mm; 94-97 - B. (P._) regnans Melvill, 1918, stn 188 (94, 97), 34 (95) and 193 (96), H = 27.5 (94), 26.6 (95), 24.4 (96) and 25.5 (97) mm; 98-100 — B. (P.) fissa (von Martens, 1901), stn 176, H = 35.3 (98), 34.7 (99) and 38.4 (100) mm; 101 — Typhlosyrinx praecipua (E.A. Smith, 1899), stn 184, H = 30.2 mm. DEEP-SEA CONOIDEAN GASTROPODS np) The species is rather variable in the shell proportions: the shell may be elongated (Fig. 94) and very similar to the holotype figured by Kilburn (1986), or more stout and broad (Figs 95 & 96). The H/D ratio varies from 2.03 to 2.55 (2.44 in the holotype). The prominence of the columellar pleat can also vary up to almost complete absence. However, there are rather constant features which are characteristic of the species and distinguish it from B. fissa (see below). These are a strongly projecting peripheral keel, distant primary spiral cords with a thin secondary rib in most interspaces, and low position of the peripheral keel, which is bordered or almost bordered by a deeply channelled suture even on the last spire whorls. The protoconch is similar to that of B. oldhami. Radular teeth are typical for the subgenus, awl-shaped, curved, and rather short. The mean tooth length is 0.46 mm at H = 26.6 mm. The operculum is small, oval, with subterminal nucleus. DISTRIBUTION. Gulf of Aden and, probably, the Bay of Bengal, 528-1080 m. Bathytoma ( Parabathytoma) fissa (von Martens, 1901) Figs 9, 17 & 98-100 Pleurotoma (Dolichotoma) fissa von Martens, 1901, p. 18. Genota (Dolichotoma) fissa—von Martens, 1903 [1904], p. 87, pl. 1, fig. 14. TYPE LOCALITY. ‘Valdivia’, stn 264 (Somalia), 1079 m. MATERIAL. stn 176, 2 specimens and 12 shells. The species differs from B. regnans in having closely set spiral cords, with almost equally strong granular secondary cords in the interspaces, and less prominent peripheral keel which is situated rather high on last spire whorls. The sutures are less distinctly channelled than in B. regnans. The shell proportions and the prominence of peripheral keel may vary (Figs 98-100). There are one to three more or less developed columellar pleats; usually two pleats are present and the upper one may be subdivided into two by a groove. The radula and operculum are similar to those of B. regnans. The mean tooth length is 0.51 mm at H = 35.3 mm. Shells from an additional sample (R/V ‘Akademik Mstislav Keldysh’, stn 1089, Tajoura Rift, Gulf of Aden, 857-900 m) show the same characteristic features and the same range of variability but are of larger size (up to 42.5 mm in height) and possess | (usually) or 2 columellar pleats. DISTRIBUTION. Somalia and Gulf of Aden, 665-1079 m. Genus TYPHLOMANGELIA G.O Sars, 1878 Type species: Pleurotoma nivale Loven, 1846 (monotypy) Typhlomangelia adenica Sysoev, new species Figs 4, 14 & 89 MATERIAL. stn 26, | shell (paratype No. 1993106); stn 185, 2 specimens (holotype No. 1993104 and paratype No. 1993105). DESCRIPTION OF HOLOTYPE. The shell is small, broad, turreted, solid, covered by thin light-brown periostracum, and consists of 4 remaining whorls. Protoconch and probably some upper whorls are missing. The whorls are strongly angled at the periphery, the subsutural slope is concave. There is a weak A.V. SYSOEV subsutural fold. Sutures are very shallow and indistinct. The body whorl is large, and the shell base is weakly convex and not differentiated from the canal. The axial sculpture consists of numerous narrow oblique folds reaching the lower suture and abruptly disappearing on the subsutural slope. The folds form pointed tubercles just below the subsutural slope. On the body whorl, they rapidly weaken downwards and do not reach the shell base. There are 19 folds on the body whorl and 15 on the penultimate. The subsutural slope is smooth except for growth lines. A single spiral cord is situated on the subsutural fold, 24 cords are present below the subsutural slope on the spire whorls, and there are about 15 low, widely and evenly spaced cords on the body whorl, which become weaker towards the anterior end and finally disappear. The aperture is oval, not differentiated from the straight and short canal. The inner lip is covered with a white callus which is thickened and becomes distinctly bordered on the canal. The anal sinus is deep, broad and rounded, slightly asymmetrical, its deepest point is situated just below the middle of subsutural slope. H = 10.2, Hb = 6.6, Ha = 4.7, D = 5.6 mm. The paratypes are very similar to the holotype but smaller (H = 7.7, D = 4.5 in paratype from stn 185 and H = 6.9, D = 4.1 in paratype from stn 26). The operculum is rather large in comparison to the aperture size, broadly leaf-shaped, with a terminal nucleus. Radular teeth are typical of the genus, awl-shaped, with short and straight shaft and relatively broad base, 0.2 mm in length. The species differs from all other species of the genus in its very small, broad shell, with a very short and straight canal. DISTRIBUTION. Gulf of Aden, 2000-2312 m. Typhlomangelia maldivica Sysoev, new species Figs 6, 15 & 86-88 MATERIAL. stn 143, 7 specimens (holotype No. 1993107 and 6 paratypes No. 1993108). DESCRIPTION OF HOLOTYPE. The shell is elongate fusiform, slender, with a high spire, rather solid, covered with thin grayish-brown periostracum, and consists of protoconch and 9 teleoconch whorls. The protoconch consists of about two globose smooth whorls, the surface is partly eroded. Definitive whorls are angled at the periphery, concave on the subsutural slope, with distinct subsutural fold. The body whorl occupies 0.58 of the shell height, the shell base is weakly convex and passes smoothly into the slightly twisted canal. The sutures are distinctly channelled, especially in last whorls. The axial sculpture consists of rounded folds below the subsutural slope; they are most prominent and tuberculate in their uppermost parts and weaken towards the lower suture. The folds tend to become smoother on last spire whorls, and on the body whorl they are very low and restricted to the whorl periphery. There are 13 folds on the body whorl and 12 on the penultimate. Spiral sculpture is represented by rather strong and widely spaced cords. One (or two on last whorls) strong cord is situated on the subsutural slope, 1-3 cords are developed below the subsutural slope of spire whorls (1 on initial whorls and 3 on the last one), and about 20 cords cover the body whorl and the canal. On the body whorl periphery, a weaker cord is situated in each interspace between primary cords. The subsutural slope of the initial whorls is smooth, but on subsequent whorls 1—S thin riblets are developed, with a stronger one in the centre of the slope. The aperture is rather narrow, oval. The canal is moderately long. The anal sinus is deep, rounded, slightly DEEP-SEA CONOIDEAN GASTROPODS asymmetrical, with the apex situated just below the middle of the subsutural slope. H = 30.2, Hb = 17.4, Ha = 13.9, D= 8.8 mm. The paratypes vary in details of spiral sculpture, especially on the subsutural slope, where several rather strong and evenly spaced riblets may be developed. In some paratypes the spiral cords are more numerous, closely spaced on the canal and more or less irregularly distributed over the body whorl surface. There may be from 13 to 15 axial folds on the body whorl. The largest paratype has H = 32.0 and D=9.1 mm. Operculum is small, oval, with terminal nucleus. Radular teeth are long and narrow, more or less curved, without a solid base, rather large (0.74 mm on the average at H = 27.8 mm). The new species resembles the type species of Typhlomangelia, Pleurotoma nivale Loven, 1846, differing well in the elongate shell with a high spire and prominent spiral ribs. DISTRIBUTION. Maldive Islands, 797 m. Genus TYPHLOSYRINX Thiele, 1925 Type species: Pleurotoma (Leucosyrinx) vepallida von Martens, 1902 (original designation) Typhlosyrinx praecipua (E.A.Smith, 1899) Fig. 101 Pleurotoma (Surcula) praecipua E.A.Smith, 1899, p. 239; Annandale & Stewart, 1910, pl. 21, figs 4, 4a. Typhlosyrinx praecipua (E.A.Smith) — Powell, 1969, p. 360-361, pl. 272, figs 2, 3. TYPE LOCALITY. ‘Investigator’, stn 229, off Travancore coast (India), 360 fms. MATERIAL. stn 184, | shell. The shell from stn 184 (H = 30.2 mm) corresponds quite well with the description and illustration of the type specimen. Axial folds (11 on the penultimate whorl) abruptly disappear on the border between the penultimate and body whorls. The spiral sculpture is developed only on the whorl periphery and the shell base, and consists of low, broad, flattened, wavy riblets unequal in width and separated by narrow grooves. DISTRIBUTION. India and Gulf of Aden, 658—1270 m. Genus GLYPHOSTOMA Gabb, 1872 Type species: Glyphostoma dentiferum Gabb, 1872 (monotypy) Glyphostoma maldivica Sysoev, new species Figs 102 & 103 MATERIAL. stn 145, | shell (holotype No. 1993109). DESCRIPTION. The shell is rather small, thick, solid, yellowish-white, and consists of the protoconch and 5 3/4 teleoconch whorls. The protoconch consists of 3 whorls; its initial part is represented by 1.5 rapidly increasing semi-transparent, smooth whorls followed by more solid angled whorls sculptured below the periphery by a narrow keel. The teleoconch whorls are concave below the suture and angled at the periphery. The sutures are clear, shallow, and wavy. The shell base is weakly convex and passes smoothly into the canal. The 23 shell surface is distinctly and minutely granular, the granulation is better seen in the interspaces between the axial folds. The growth lines are mostly indistinct. The axial sculpture is represented by rounded folds extending from the subsutural slope to lower suture and, on the body whorl, to the canal. There are 24 such folds on the body whorl and 19 on the penultimate. On the subsutural slope, the axial sculpture consists of numerous curved and rather weak folds with sharpened crests. They generally represent the continuation of main axial, folds but there may also be interstitial folds; as a result, the subsutural slope of the body whorl is covered with 32 folds. The spiral sculpture is represented by strong ribs almost equal in prominence to the axial folds. At the intersection with axial sculpture, the ribs form rounded tubercles. The interspaces between spiral ribs are covered by closely set threads. The ribs on the subsutural slope are much smaller corresponding to much smaller axial folds. The aperture is elongate-oval. The inner lip bears two rather prominent pointed tubercles in its middle part and several smaller ones in the lower part. The outer lip is sharpely and wavely edged and bordered by a heavy curved varix. The inner surface of the aperture bears one strong tubercle in the upper part and a group of 5 tubercles below. The anal sinus is deep, U-shaped, bordered with callus and constricted at its entrance by a heavy tuberculated callus pad. The canal is straight along most its length and slightly curved backwards near the end. H = 17.6, Hb = 11.8, Ha = 9.8, D= 8.6 mm. In general outline the new species is most similar to Clathurella perlissa E.A.Smith, 1904 from the Andaman Islands but differs in the character of sculpture and apertural armament. The species corresponds well to the genus Glyphostoma in all important conchological characters such as characteristic protoconch, granular surface, prominent subsutural slope with different sculpture (in contrast to Etrema), and strong intersecting spiral and axial ribs. DISTRIBUTION. Maldive Islands, 494 m. Glyphostoma supraplicata Sysoev, new species Figs 104 & 105 MATERIAL. stn 176, 6 shells (holotype No. 1993110 and 5 paratypes No. 1993111). DESCRIPTION OF HOLOTYPE. The shell is rather small, slender, fusiform, relatively solid, white, consists of protoconch and almost 7 teleoconch whorls. The protoconch consists of 3.5 whorls, the tip is small and papillate. Initial embryonal whorls are smooth, the two last whorls angled, with a cord-like peripheral keel, the second weaker keel encircles the lower suture. The spire is tall, occupies about 0.4 of the shell height. Definitive whorls are angled at the shoulder and in the lower part, the subsutural slope is weakly concave or almost flat. The sutures are clear, shallow, slightly wavy. The entire shell surface is densely and minutely granulated, but the granulation more or less disappears on prominent parts of the sculpture. Growth lines are thin and irregularly elevated. Axial sculpture consists of strong, oblique, rounded folds separated by interspaces approximately equal in width to the folds. The folds reach the canal, gradually weakening on the shell base. On the subsutural slope, they are much weaker, narrower, almost obsolete near the upper suture, and curved in correspondence to the anal sinus scars. There are 15 folds on the body whorl and 13 on the penultimate. Spiral sculpture is represented by broad cords DEEP-SEA CONOIDEAN GASTROPODS below the shoulder (2 on the spire whorls and 18 on the body whorl plus canal). The cords are separated by 2—3 times wider intervals. They are often subobsolete in interstices between axial folds but very strong when overrided the latter. On the body whorl, the cords gradually diminish in width and prominence towards the canal end. There are 1-4 (usually 2) thin threads in the interspaces between cords, except between those on the canal, and 3-4 low rounded threads on the subsutural slope. Strong rounded tubercles are formed at the intersection between axial folds and spiral cords resulting in a beaded appearance of the spiral sculpture. The aperture is rather narrow, gradually narrowing towards the canal. The inner lip bears about 13 weak transverse plicae in its lower part, the plicae become closer to each other towards the canal end. There is a moderately developed parietal callus pad which is weakly tuberculate in its lower part. The outer lip has a sharpened edge, with a strong varix behind. Inside the aperture, there are 5 transverse plicae most prominent in the region corresponding to the varix, and a strong tubercle just below the anal sinus. The sinus is semi-tubular, U-shaped, broad and rounded, directed outside in relation to subsutural slope. The canal is long, slightly twisted, and obliquely truncated at the end. H = 22.6, Hb = 13.4, Ha = 11.3, D=7.9 mm. Paratypes vary in minor details of sculpture, e.g. in the prominence of axials on the subsutural slope, and in more or less strong nodules at the intersections between spiral and axial elements. The largest paratype is 22.6 mm high (protoconch missing). The new species is quite similar to G. sultana (Thiele, 1925) from East Africa, differing in the more slender shell (H/D ratio is 2.70-2.91 vs. 2.55 in the holotype of G su/tana) and in the presence of weakened but distinct axial folds on the subsutural slope. A peculiar feature of the new species is the presence of a second keel encircling the suture on last protoconch whorls. DISTRIBUTION. Gulf of Aden, 655—732 m. Subfamily DAPHNELLINAE Deshayes, 1863 Genus CRYPTODAPHNE Powell, 1942 Type species: Cryptodaphne pseudodrillia Powell, 1942 (original designation) Cryptodaphne gradata (Schepman, 1913) Fig. 108 Pleurotomella gradata Schepman, 1913, p. 445, pl. 30, fig. 2. Cryptodaphne gradata (Schepman) — Shuto, 1971, p. 11, pl. 2, figs 4-6. TYPE LOCALITY. ‘Siboga’, stn 159, Halmahera Sea, 411 m. MATERIAL. stn 176, | shell. The shell from the JME material basically conforms to the description and illustrations of the holotype given by Schepman and Shuto (see synonymy). However it differs from the latter in some remarkable characters. The shell is much larger than that in P45) the holotype (18.5 mm vs. 10.0 mm), with a proportionally higher spire, at approximately the same number of teleoconch volutions (6+ vs. 6). The anal sinus is deeper, with its apex situated lower on the subsutural slope. The initial three whorls of the teleoconch bear oblique axial folds below the peripheral angulation. These folds form nodes on the angulation, making it crenulated, and rapidly weaken towards the lower suture. On subsequent whorls, they become less developed and disappear on the body whorl. The spiral sculpture lacks a regular alternation of strong and weak spirals mentioned by Shuto. The sculpture has a cancellated appearance due to the intersection with growth lines. Also there are widely spaced, thin, and weak but distinct spiral threads on the subsutural slope which were not described by either Schepman or Shuto. Nevertheless, these differences are not essential and seemingly do not extend beyond the range of intraspecific and geographical variability of C. gradata. DISTRIBUTION. Halmahera Sea and Gulf of Aden, 411—732 m. Genus FAMELICA Bouchet & Warén, 1980 Type species: Pleurotomella catharinae Verrill & Smith, 1884 (original designation) Famelica tajourensis Sysoev & Kantor, 1987 Fig. 112 Famelica tajourensis Sysoev & Kantor, 1987, p. 1257, fig. g, d, e, zh. TYPE LOCALITY. ‘Akademik Kurchatov’, stn 1095, Tajoura Rift, Gulf of Aden, 1330-1406 m. MATERIAL. stn 188, | shell. The JME shell agrees well with the type material and differs mainly in having thin and transparent shell walls compared with rather thick and solid in the type specimens. The species is rather different from the type species of the genus Famelica, but very similar to F: monotropis (Dautzenberg & Fischer, 1896), which was included into Famelica by the authors of the genus (Bouchet & Warén, 1980). On the other hand, the genus seems to be rather heterogeneous in respect to species originally included into it. Nevertheless, I do not know any other genus which can accomodate F: tadjourensis. A similar genus is Pagodidaphne Shuto, 1983, but it differs in having a less distinct keel on the whorl shoulder and much less elongate body whorl with a short canal. DISTRIBUTION. Gulf of Aden, 528—1406 m. Genus GYMNOBELA Verrill, 1884 Type species: Gymnobela engonia Verrill, 1884 (subsequent designation Cossmann, 1896) Figs 102-112 Clathurellinae and Daphnellinae. 102, 103 — Glyphostoma maldivica Sysoev, new species, holotype; 104, 105 — G supraplicata Sysoev, new species, holotype; 106, 107 — Gymnobela adenica Sysoev, new species, holotype (106) and paratype, stn 185, H = 7.1 mm (107); 108 — Crypto- daphne gradata (Schepman, 1913), stn 178, H = 18.5 mm; 109-111 — Gymnobela africana Sysoev, new species, holotype (109, 110) and paratype, stn 118, H = 68.0 mm (111); 112 — Famelica tajourensis Sysoev & Kantor, 1987, stn 188, H = 10.6 mm. 26 Gymnobela adenica Sysoev, new species Figs 106 & 107 MATERIAL. stn 185, 2 specimens (holotype No. 1993112 and paratype No. 1993113). DESCRIPTION OF HOLOTYPE. The shell is small, broadly biconic, thin, yellowish-white, and consisting of 5 remaining whorls. The protoconch is missing, and the upper teleoconch whorls are eroded. The whorls are angled below the periphery. The subsutural slope is almost flat on the upper spire whorls and concave on the body whorl. The uppermost part of subsutural slope is slightly raised forming an indistinct subsutural fold. The sutures are shallow. Growth lines are mostly indistinct, some of them form clear, narrow, oblique folds regularly set on the upper third of the subsutural fold and approximately twice as numerous as main axial folds. The latter are strongly oblique, narrow, with sharpened crests, and tuberculate at the place of whorl angulation. The folds are separated by narrow intervals, abruptly disappear on the subsutural slope and extend to the lower suture on the spire whorls and to the upper part of the shell base. There are 21 folds on the body whorl and 20 on the penultimate. Spiral ribs (about 30 on the body whorl plus canal) are strong, flattened and uniform except for narrower ones on the canal. Intervals between the ribs are approximately equal to ribs in width. The subsutural slope is densely covered with thin, low, rounded and closely set riblets (about 14 on each of two last whorls). The shell base with a distinct bend passes into a short and straight canal. The aperture is rather small, subrectangular, with the inner lip distinctly bent. The canal is narrow, attenuated at its end. The anal sinus is moderately deep, broadly rounded, its deepest point is situated in the middle of subsutural slope. H = 9.5, Hb= 6.9, Ha = 5.1, D= 6.0 mm. The paratype is very similar to the holotype except for smaller size (H = 7.1, D = 5.3 mm, 3 teleoconch whorls), smaller H/D ratio, and longer folds formed by growth lines which often occupy the whole subsutural slope. There are 24 main axial folds on the body whorl. The last protoconch whorl preserved is covered with typical diagonally cancellated sculpture. The species differs from other species of the genus in its small broadly biconic shell with numerous strongly oblique axials and short attenuated canal. DISTRIBUTION. Gulf of Aden, 2000 m. Subgenus BATH YBELA Kobelt, 1905 Type species: Thesbia nudator Locard, 1897 (subsequent designation Dall, 1918) Gymnobela (Bathybela) africana Sysoev, new species Figs 109-111 MATERIAL. stn 118, 1 specimen (holotype, No. 1993114) and 1 shell (No. 1993115). DESCRIPTION OF HOLOTYPE. The shell is large, broadly fusiform, thin but solid, reddish-brown to light-brown, and with a slightly glossy surface. It consists of 6 whorls, the protoconch is missing. The whorls are obtusely angled above the periphery, moderately convex below the angulation and slightly concave above it. The sutures are rather shallowly impressed, clear, straight or wavy in some places. The body whorl is large, A.V. SYSOEV occupies about 0.7 of the shell height; the shell base is weakly convex, with a distinct bend passes into the straight canal. The axial sculpture consists of narrow oblique folds, often with sharpened crests. The folds begin in the lower part of the subsutural slope and extend to the lower suture on the spire whorls and only to the periphery on the body whorl. They are most prominent in the place of the whorl angulation. There are 17 folds on the body whorl and 15 on the penultimate one. The growth lines are indistinct on the whorl surface except the subsutural slope, some of them are thickened and raised. The thickened growth lines, however, do not form the regularly arranged plicae on the subsutural slope which are characteristic of many deep-sea Daphnellinae. The spiral sculpture is represented by rather wide, flattened, wavy ribs unequal in size and separated by narrow grooves. The subsutural slope is smooth except for several very feeble spiral lines. The aperture is broad, the inner lip forms a distinct bend, and the columella is twisted. The anal sinus, judging from the growth lines, is shallow and broad, its deepest point is situated immediately above the middle of subsutural slope. The dried soft body has no operculum. H = 54.7, Hb = 38.5, Ha = 30.1, D= 22.7 mm. The shell of the paratype is larger than in the holotype (H = 68.0, Hb = 48.2, Ha = 38.5, D = 30.0 mm). It was dead-collected and the surface is rather worn. The paratype differs from the holotype mainly in its shorter axial folds developed on the body whorl only at the place of angulation and obsolete in the last quarter of body whorl, and in the canal curved backwards. The new species is rather similar to Spergo sibogae Schepman, 1913 from Indonesia differing in much broader shell (H/D = 3.0 in S. sibogae and 2.3—2.4 in G africana). It also shows some similarity to Pontiothauma pacei E.A.Smith, 1906 from India and Ceylon differing in somewhat more slender shell with short axial folds and faint spiral sculpture. DISTRIBUTION. East Africa eastward of Mombasa, 1789 m. Subgenus THETA Clarke, 1959 Type species: Pleurotomella (Theta) lyronuclea Clarke, 1959 (original designation) Gymnobela ( Theta) daphnelloides (Dall, 1895) Figs 113 & 114 Mangelia (Spergo) daphnelloides Dall, 1895, p. 683-684, pl. 31, fig. 11. TYPE LOCALITY. ‘Albatross’, stn 3476 (Hawaiian Islands), 298 fms. MATERIAL. stn 118, 1 shell. The species was originally described within the subgenus Spergo which is considered as a full genus by all modern authors. However the species differs from the type species of Spergo, Mangilia (Spergo) glandiniformis (Dall, 1895), in all important conchological characters included by Dall in the original diagnosis of the subgenus (the latter was based to a considerable extent on the soft body characteristics which are presently treated as insignificant for taxonomy at the generic level), i.e. in having well developed sculpture and rather deep anal sinus. On the other hand, the species is very similar to the group of species assigned by Bouchet & Warén (1980) to the genus Theta. The | | Type species: Awateria personata Powell, | designation) DEEP-SEA CONOIDEAN GASTROPODS 119 27 Figs 113-122 Daphnellinae and Mangeliinae. 113, 114— Gymnobela (Theta) daphnelloides (Dall, 1895), stn 118, H = 27.0; 115 — Mioawateria exten- saeformis (Schepman, 1913), stn 26, H = 9.9 mm; 116, 117 — Xanthodaphne maldivica Sysoev, new species, holotype; 118-121 — Benthomangelia brachytona (Watson, 1881), stn 119, H = 17.1 mm (118, 119) and lectotype, BM(NH) 1887.2.9.1034, H = 15.1 mm (120, 121); 122 — B. trophonoidea Thiele, 1925, stn 185, H= 10.0 mm. latter, in its turn, is too poorly distinguished from Gymnobela to warrant a generic status and should be regarded as a subgenus. DISTRIBUTION. Hawaiian Islands and East Africa, 545-1789 m. The present record is a great range extension for the species. Genus MIOAWATERIA Vella, 1954 1954 (original Mioawateria extensaeformis (Schepman, 1913) Fig. 115 Pleurotomella extensaeformis Schepman, 1913, p. 446, pl. 30, fig. 5. Gymnobela extensaeformis (Schepman) — Thiele, 1925, p. 190(224), pl. 41(29), fig. 9. Magnella extensaeformis (Schepman) — Shuto, 1971, p. 15-16, pl. 2, figs 10-12. TYPE LOCALITY. ‘Siboga’, stn 212, Banda Sea, 462 m. 28 MATERIAL. stn 26, 2 specimens. Of the two specimens available (H = 9.9 and 6.3 mm, both without protoconch), the smaller is comparable in size to those illustrated by Schepman (8.5 mm, or 8.0 mm according to Shuto, 1971) and Thiele (6.25 mm) and quite similar to them. The larger specimen (Fig. 115) differs in having a broader canal, less excavated subsutural slope, and much weaker peripheral spiral keel on the body whorl. The spiral sculpture of both specimens is subobsolete on the body whorl periphery and upper part of the shell base becoming stronger towards the canal. This is in contrast to both Schepman’s and Thiele’s figures, but agrees well with Shuto’s (1971) illustration of the holotype. The same species was probably figured by Thiele (1925) as the North-Atlantic Gymnobela extensa (Dall, 1881). Thiele’s figure, apparently based on a specimen from Sumatra, differs from his illustration of M. extensaeformis in having obsolete spiral ribs on the body whorl except for those on the lower shell base and canal, but, as mentioned above, this is the feature characteristic of M. extensaeformis. Recently this species was assigned by Shuto (1971) to Magnella Dittmar, 1960. Later, Maxwell (1988) synonymized the latter genus with the New Zealand genus Mioawateria Vella, 1954. This synonymization seems to be reasonable, though the status of Mioawateria itself remains somewhat uncertain due to great similarity to Gymnobela Verrill, 1884. The main difference between two latter genera is the shape of anal sinus which is very shallow in Mioawateria. At the same time, the shape of sinus in Gymnobela is rather variable, and there are species, traditionally included into Gymnobela, quite comparable to Mioawateria in this character (e.g. G blakeana (Dall, 1881)). Thus, after comparative examination of broad range of Gymnobela species in respect to the anal sinus shape, Mioawateria may be either a large and very widely distributed genus or a synonym of Gymnobela. DISTRIBUTION. East Africa (Gulf of Aden to Kenya), Sumatra, Banda Sea, 439-2312 m. The present record is the deepest one. Genus XANTHODAPHNE Powell, 1942 Type species: Pleurotoma (Thesbia) membranacea Watson, 1886 (original designation) Xanthodaphne maldivica Sysoev, new species Figs 116 & 117 MATERIAL. stn 143, 1 specimen (holotype No. 1993116). DESCRIPTION. The shell is narrowly fusiform, slender, thin but solid, light-brown, with glossy surface, consists of the protoconch and 9 teleoconch whorls. The protoconch is partly broken off, but the remaining 1.5 whorls are covered with the typical diagonally cancellated sculpture. The teleoconch whorls are slightly concave in the upper part and weakly convex below. Early teleoconch whorls are angled in the lower part just near the suture, this angulation rapidly shifts upwards and becomes less prominent and practically disappears on the 6th whorl. There is a weak subsutural fold on initial teleoconch whorls. The sutures are very shallow, distinct, and more or less straight. The growth lines are clear, thin, numerous, and strongly curved. The sculpture is represented only by low, wide ribs on the canal; these rapidly become obsolete on the shell base. The shell base is weakly convex and smoothly passes into the canal. The canal is A.V. SYSOEV straight, not differentiated from narrow aperture. The inner lip is covered with a very weak callus which becomes thicker towards the canal extremity. The anal sinus is wide and moderately deep, subsutural, ‘reversed L’-shaped, its deepest part is situated just below the suture. The outer lip strongly projects forward below the sinus. H = 29.0, Hb = 17.1, Ha = 14.3, D= 9.0 mm. The new species differs from all known species of the genus in its slender and narrow shell almost completely devoid of spiral sculpture. DISTRIBUTION. Maldive Islands, 797 m. Subfamily MANGELIINAE Fischer, 1883 Genus BENTHOMANGELIA Thiele, 1925 Type species: Surcula trophonoidea Schepman, 1913 (original designation) Benthomangelia brachytona (Watson, 1881) Figs 118-121 Pleurotoma ( Drillia) brachytona Watson, 1881, p. 415. Pleurotoma ( Spirotropis) brachytona Watson — Watson, 1886, p. 324-325, pl. 18, fig. 3. TYPE LOCALITY. ‘Challenger’, stn 191, off the Arrou Island, south-west of Papua, 800 fms. MATERIAL. stn 119, 1 shell. The specimen from the stn 119 (H = 17.1 mm) is quite similar to Watson’s illustration and to the photograph of one of two syntypes (which should be designated as lectotype) (Figs 120 & 121, H = 15.3 mm according to Watson) (the second syntype is represented by broken and heavily worn shell) stored in the NHM. It differs from the lectotype only in more slender shell (H/D = 2.20 vs. 2.03 (Watson’s measurements) or 1.96 (measured by the photograph) in Watson’s specimen) with fewer axial folds. DISTRIBUTION. East Africa and Indonesia, 1463 and 1207-1463 m. Benthomangelia trophonoidea (Schepman, 1913) Fig. 122 Surcula trophonoidea Schepman, 1913, p. 62(426)-63(427), pl. 28, fig. 3. Mangelia (Benthomangelia) trophonoidea (Schepman) — Thiele, 1925, p. 190(224)-191(225), pl. 27(39), fig. 25, text-fig. 25. Benthomangelia trophonoidea (Schepman) — Okutani, 1966, p. 23, text-fig. 11. ? Marshallena gracilispira Powell, 1969, p. 370-371, pl. 281, fig. 2. TYPE LOCALITY. ‘Siboga’, stn 45, Flores Sea, 794 m. MATERIAL. stn 185, 1 shell. The shell from the JME material is small (H = 10.0 mm, which is smaller than all the previously recorded specimens, i.e. 16 mm (Schepman), 15.9 mm (Thiele, measured by the figure), and 15.5 mm (Okutani, measured by the figure)) and apparently rather young. It differs from the original description and figure in having prominent tubercles on the subsutural fold of all the shell whorls and in less curved and more developed axial ribs on the subsutural slope. The presence of short plicae on early whorls, DEEP-SEA CONOIDEAN GASTROPODS which disappear towards the body whorl, was mentioned by Shepman for the much larger paratype. The prominent axial ribs on the subsutural slope are seen in Thiele’s figure of the species. Judging from the published figures of B. trophonoidea, the species is rather variable, though it can be mentioned that the proportions of the shell (H/D and Hs/H) are rather constant in different specimens. Thiele’s figure illustrates a somewhat more slender shell with better differentiated and more straight siphonal canal and much longer axial ribs, almost reaching the canal (they do not reach the whorl periphery in the holotype). Okutani published an illustration of a shell with a high and narrow spire. However, the variability of B. trophonoidea does not exceed that of the Atlantic representatives of the genus (Bouchet & Warén, 1980). Marshallena gracilispira Powell, 1969, described from Borneo and Philippines, 558-717 m, is probably a synonym of B. trophonoidea; from both original description and figure it is impossible to find any essential characters distinguishing the former species from the latter. However, this question cannot be resolved without an examination of type specimens and more material. DISTRIBUTION. Gulf of Aden, Indonesia, and southern Japan, 660-2000 m. The present record is the most western and most deep-sea locality. The following species were also mentioned from the John Murray Expedition bathyal samples by A.W.B.Powell (1964, 1969) but not found in the material studied: Lucerapex denticulata (Thiele, 1925) — Powell, 1964, p. 286 (stn 176, Gulf of Aden, 732 m; stn 184, Gulf of Aden, 1270 m). Nihonia circumstricta (von Martens, 1901) — Powell, 1969, p. 334 (stn 110, off Pemba Id., 333 m). Leucosyrinx julia Thiele, 1925 — Powell, 1969, p. 338 (stn 34, Gulf of Aden, 1040 m). Typhlosyrinx vepallida (von Martens, 1902) — Powell, 1969, p. 360 (stn 184, Gulf of Aden, 1270 m). Marshallena philippinarum (Watson, 1882) — Powell, 1969, p. 369-370 (three stations without numbers indicated, Gulf of Aden and off Pemba Id., 1061, 1022, and 802 m). DISCUSSION A total of 50 species of deep-sea conoidean gastropods were found in the JME collection. They belong to 3 families, 6 subfamilies, 22 genera and 3 subgenera. Lower conoideans (families Drilliidae and Turridae) prevail in the material: 30 species vs. 20 in Conidae. Among subfamilies, the most species-rich appeared to be Cochlespirinae (9 species) and Clathurellinae (11 species). The material studied seemingly does not represent well the fauna of the North-Western Indian Ocean; this is evidenced by low percentage of small species and juvenile individuals of larger species as compared, for example to East African bathyal fauna described by Thiele (1925). This is probably due to methods of collection. Additional evidence for this may be the very high share of lower conoideans, often respresented by large species, (60%) which far exceeds the respective values for East African fauna (about 41%, calculated from Thiele’s (1925) list) and for other world-wide faunas (Sysoev, 1991). Nevertheless, some remarks on the JME collection can be made. The fauna has a typical bathyal appearance, with only few 29 representatives of characteristic shallow-water (e.g. Drillia) and abyssal (e.g. Gymnobela ( Theta) genera. The high percentage of new species in the material studied (about 1/3) indicates that the North-Western Indian Ocean is still insufficiently explored. This is additionally confirmed by the very low overlap with faunal lists obtained by other expeditions: for example, only 5 out of 59 bathyal species reported by Thiele (1925) for East Africa were found in the JME collection (plus 4 species recorded by Powell — see above); the same is true for the bathyal fauna of Southern India collected by the ‘Investigator’ (7 species found out of 37 reported by Winckworth, 1940). Most species were found at only one station, and thus the main areas covered by the JME investigations (i.e. Gulf of Aden, Zanzibar area, and Maldive Islands) have very few common species: 2 species (Comitas subsuturalis and Gemmula amabilis) were found in all three areas, 2 species (Gemmula bisinuata) and Borsonia symbiophora) — in the two first regions, and one species (Comitas erica) — in the two last. All species also found outside the region studied are apparently widely distributed in the Indo-Pacific. Of particular interest in this connection are the findings which greatly extend the geographic range of respective species: Horaiclavus splendidus previously known from Japan, Leucosyrinx claviforma from North-Western Australia, and Gymnobela daphnelloides from Hawaii. ACKNOWLEDGEMENTS. I am greatly indebted to Dr J.D.Taylor and Ms K. Way of The Natural History Museum, London for the loan of the material and providing relevant information. Dr J.D.Taylor and Dr P.Mordan of The Natural History Museum kindly edited the manuscript. Of great value were the taxonomic comments of an anonymous referee. I thank Dr Yu.I.Kantor of the A.N.Severtzov Institute of Problems of Evolution of Russian Academy of Sciences, Moscow, for his assistance. REFERENCES Abbott, R.T. & Dance, S.P. 1990. Compendium of seashells. 4th edition. 411 p. American Malacologists, Inc., Melbourne, Florida. Adams, A. 1867. Descriptions of new species of shells from Japan. Proceedings of the Zoological Society of London: 309-315. Alcock, A. & Anderson, A.R.S. 1898. Illustrations of the Zoology of the Royal Indian Marine Survey Ship Investigator, under the command of Commander T.H. Heming, R.N. Mollusca. Part 2. pl. 7-8. Calcutta. Alcock, A., Annandale N., & MacGilchrist, A.C. 1907. I/lustrations of the Zoology of the Royal Indian Marine Survey Ship Investigator, under the command of Commander T.H. Heming, R.N. ( Retired). Mollusca. Part 4. pl. 14-18. Calcutta. 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Einige neue Meer-Conchylien von der deutsche Tiefsee-Expedition. Sitzungs-berichte der Gesellischaft naturforschender Freunde zu Berlin: 14-26. 1902. Sitzungs-berichte der Gesellischaft naturforschender Freunde zu Berlin: 19. — 1903 [1904]. Die beschalten Gastropoden der deutschen Tiefsee-Expedition 1898-1899. A. Systematisch-geographischer Teil. Wissenschaftliche Ergebnisse der deutschen Tiefsee-Expedition auf dem Dampfer ‘Valdivia’ 1898-1899. Jena, G. Fischer, 7: 1-146. McLean, J.H. 1971. Family Turridae. pp. 686-766 Jn: Keen, A.M. Seashells of tropical West America; marine molluscs from Baja California to Peru. 2nd ed. Stanford, Stanford University Press. Melvill, J.C. 1910. Descriptions of twenty-nine species of marine Mollusca from the Persian Gulf, Gulf of Oman, and North Arabian Sea, mostly collected by Mr. F.W.Townsend of the Indo-European Telegraph Service. Annals and Magazine of Natural History, ser. 8, 6: 1-17. Maxwell, P.A. 1988. 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Prosobranchia of the Siboga Expedition. Part 5. Toxoglossa. Siboga Expeditie Monograph, 49: 365-452. Shuto, T. 1965. Turrid gastropods from the Upper Pleistocene Moeshima Shell Bed (Molluscan palaeontology from Pleistocene formations in Kyushu -1). Memoirs of the Faculty of Science, Kyushu University, series D, Geology, 26(2): 143-207. 1970. Taxonomical notes on the turrids of the Siboga-Collection originally described by M.M.Schepman, 1913 (Part I). Venus, 28(4): 161-178. — 1971. Taxonomical notes on the turrids of the Siboga-Collection originally described by M.M.Schepman, 1913 (Part III). Venus, 30(1): 5—22. — 1975. Notes on type species of some turrid genera based on the type specimens in the British Museum (N.H.). Venus, 33(4): 161-175. 1983. New turrid taxa from the Australian waters. Memoirs of the Faculty of Science, Kyushu University, series D, Geology, 25(1): 1-26. Smith, E.A. 1894. Report upon some Mollusca dredged in the Bay of Bengal and the Arabian sea. 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Preliminary analysis of the relationship between turrids (Gastropoda, Toxoglossa, Turridae) with different types of radular apparatus in various Recent and fossil faunas. Ruthenica, Russian Malacological Journal, 1(1—2): 53-66. Sysoev, A.V. & Kantor, Yu.I. 1987. Three new species of the deep-sea molluscan genus Famelica (Gastropoda, Toxoglossa, Turridae). Zoologicheskij Zhurnal, 56(8): 1255-1258 (In Russian). Taki, I. & Oyama, K. 1954. Matajiro Yokoyama’s The Pliocene and later faunas from the Kwanto region in Japan. Paleontological Society of Japan Special Papers, 2: \-68. Taylor, J.D., Kantor, Yu.I. & Sysoev, A.V. 1993. Foregut anatomy, feeding mechanisms, relationships and classification of the Conoidea (= Toxoglossa) (Gastropoda). Bulletin of the Natural History Museum, Zoology, 59(2): 125-170. Tesch, P. 1915. Jungtertiare und Quartare Mollusken von Timor. Teil I. Paldontologie von Timor, 5: \—70. Thiele, J. 1925. Gastropoda der deutschen Tiefsee-Expedition. II Teil. Wissenschaftliche Ergebnisse der deutschen Tiefsee-Expedition auf dem Dampfer ‘Valdivia’ 1898-1899. Jena, G.Fischer, 17(2): 1-348 (35-382) Watson, R.B. 1881. Mollusca of H.M.S. ‘Challenger’ expedition — part 9. Journal of the Linnean Society, 15: 413-455. 1886. Report on the Scaphopoda and Gasteropoda collected by H.M.S. Challenger during the years 1873-76. Reports of the Scientific Results of the voyage of H. M.S. Challenger, Zoology, 42: 1-756. Weinkauff, H.C. 1875. Die familie Pleurotomidae. Systematisches Conchylien- Cabinet, 4(3): 1-248, 43 pls. Nurenberg: Bauer & Raspe. Winckworth, R. 1940. A systematic list of the Investigator Mollusca. Proceedings of the Malacological Society of London, 24(1): 19-29. Bull. nat. Hist. Mus. Lond. (Zool.) 62(1): 31-36 Issued 27 June 1996 Reassessment of ‘Calcinus’ astathes Stebbing, 1924 (Crustacea: Anomura: Paguridea: Diogenidae) PATSY A. McLAUGHLIN Shannon Point Marine Center, 1900 Shannon Point Road, Anacortes, WA 98221-4042, U.S.A. CONTENTS : : : {HATO (CW CO) EL. dosdeetcratect deochcBpanaenosecRaRaee peace, cea teti gad CER eae R REP ee MERE Es mee ee OCD Sa eee 31 IMUTRCUTCIITIGINIVA CUM OCS Ime ee on, Cetera ae aE ne Pe cence sone rene rete ene es Aol ce cs aiscde aa Sovressse se gosee dues des sueesnceam omens ae sete Senor e 31 | PEMSUCKMALIG UNCOOL tae rkcats Mexstttcescenaceteesacastccvterescvsecuasee loans ceussnapecssdusseradee Mernncnadnuctohsiacs tasuadacdtacseeess veld Smear eee nne eT, FV CCLESCHIPLOMUR satay eets caawavens cctevie Sie tnanstecvosaceanasnadcecsetuacaccudedSaeaatadssnddsahenct teeubs dk ba sigha sh teak WacndaeeeoeMNSaice eco POSES ERAN OL aeons cet rote se arcs ence een ncaa at Fe oe rae ve Se eT Cao sv a bso seidee LE LSTTA TET eee OER AEG Bs OR ene PE, SS 34 | PSII ERY ny Cee EEOC GEER TEE = Soc Oe REECE UOC R RE Ae ono EERE EEE PRE PE EB toe ering Here EEE BRS EEGREERL A ccanacinn Gecaorc aos Saco 34 REITER sce thne Aasdchiccone Bose sBcebe iciss sac bro O te Peet BA GREED Ee eeeR EMRE EMEA RE REN CCR tS Sy cae ERA eet ney coca Coder o ees 35 | FN CKIT OM EG DELICE Leena tree UMN ees SPL AIS, C5 ORT IM cocscsscavunsnudacicuersdsacvasuidensesvaccounevscuceeevennecedeteceeeceaeneceecenseoeeree OD | Stone MCCSpe tee eee rn Seen Seam riate ie: FERN MAB 2.6 Sinan ccocaaisvessveveveteadv scouts suagealduciasdesvase Meee Soe OOD Synopsis. A reexamination of the syntypes of ‘Calcinus’ astathes Stebbing, 1924 in the collection of The Natural History Museum, London, U.K., has shown that although subsequently assigned correctly to the genus Clibanarius, this taxon is not conspecific with C. virescens Krauss, 1843, as was previously proposed at that time. Clibanarius astathes is redescribed and illustrated; a lectotype is designated. INTRODUCTION Stebbing (1924) described a new species of hermit crab from Delagoa Bay, South Africa, which he mistakenly assigned to Calcinus Dana, 1851. The fact that this species clearly should _ have been placed in Clibanarius Dana, 1852 was recognized by Barnard (1947); however, at that time he made only the notation | that four specimens returned to the South African Museum by Stebbing were indistinguishable from Clibanarius virescens | (Krauss, 1843). Barnard (1950) went on to define those four specimens further by commenting, “I would not have ventured to | dispute the identity of Stebbing’s Calcinus astathes if there had not been four specimens returned to the Museum bearing _ Stebbing’s autographic label (the largest and type specimen | probably retained by him, or perhaps now transferred to the | British Museum). These four specimens are obviously C. | virescens. The dactyl of the 3rd leg is not longer than 6th joint, and has the characteristic shape’. Stebbing’s (1924) original description of Calcinus astathes specified only a ‘group’ of specimens collected from Delagoa | Bay by K.H. Barnard on October 12, 1912. Five specimens, | listed as syntypes of this taxon are part of the Stebbing Collection donated to The Natural History Museum (NHM) [formerly British Museum (Natural History)] by Barnard, although only three initially were listed in the Museum registry _ (NHM 1928.12.1.264-266). Four specimens, dry, and in poor condition, remain in the collection of the South African Museum (SAM) catalogued under the original number (A2121) | ©The Natural History Museum, 1996 listed by Stebbing (1924). Barnard’s hand written label accompanying the dry specimens reads ‘A2121. Delagoa Bay 4 spec. returned by Stebbing labelled (sic) as ‘Calcinus astathes’ also 1 with parasites see Stebbing. 1920‘. Barnard’s label reference to the parasitized specimen must refer to a specimen without chelipeds included by Stebbing (1920), with others (NHM 1928.12.1.267-268, SAM A3270 A2120), identified as Calcinus laevimanus (Randall, 1840), the former specimen similarly placed in synonymy with C. virescens by Barnard (1950: 435). No specimens of Calcinus or Clibanarius astathes were listed among the type collection in the South African museum by Kensley (1974), and none have been found on a recent search (Ms. L. Hoenson, pers. comm.). Despite some confusion in the initial NHM registry entry for “Calcinus’ astathes, at least two points are clear. These five specimens labeled as syntypes of Stebbing’s (1924) species all belong to the same taxon; none are conspecific with Clibanarius virescens. The four specimens remaining in the SAM collection truly do represent C. virescens. The largest of the NHM specimens [and the one presumably unaccounted for by Barnard (1950)] is herein designated as the lectotype of ‘Calcinus’ astathes. MATERIAL AND METHODS In addition to the syntypes, and Barnard’s four specimens of Clibanarius virescens, comparative material of a 32 morphologically similar Clibanarius species, C. longitarsus (De Haan, 1849) from Durban Bay, S.A., has been examined. One measurement, shield length (SL), measured from the tip of the rostrum to the midpoint of the cervical groove provides an indication of specimen size. The type material of C. astathes has been returned to the Natural History Museum, and Barnard’s C. virescens to the South African Museum. Two specimens of C. longitarsus used in the comparison also have been deposited in The Natural History Museum (NHM 1995.163—164), the remaining have been retained in the author’s personal collection. SYSTEMATIC ACCOUNT Clibanarius astathes (Stebbing, 1924) Figs 1,2 Calcinus astathes Stebbing, 1924: 239, pl. 2 (CXVII of continuing series). Clibanarius virescens: Barnard, 1947: 376 (in part); 1950: 435 (in part). Lectotype (herein selected). Female (SL — 7.3 mm), NHM reg. 1928.12.1.264 Paralectotypes. Two females (SL = 6.3, 6.3 mm), 2 males (SL = 5.8, 6.9), NHM reg. 1928.12.1.265—266. TYPE LOCALITY. Delagoa Bay (25°50°S, ‘Mosambique’, Ethiopian region, Indian Ocean, 1912. 82-5088); DIAGNOSIS. Rostrum with simple or bifid termination. Ocular peduncles 0.66—0.75 length of shield; little if at all overreached by antennular peduncles; antennal peduncles not reaching to bases of corneae. Ocular acicles with acute, simple or bifid termination. Basal segment of antennular peduncle with 1 or 2 small spines. Chelipeds subequal, right somewhat larger. Right cheliped with dactyl equal to or slightly longer than palm; dorsomesial margin and dorsal surface with rows of corneous-tipped spines. Dorsomesial margin of palm with row of low spines and tufts of long setae, second adjacent row of spines and one large tubercle at proximal margin, dorsal surface sloping, with few spines distally and on proximal portion of fixed finger. Dorsomesial distal angle of carpus with acute spine, low protuberances and tufts of setae on dorsomesial margin. Merus with few low, somewhat spinulose protuberances on ventromesial margin; ventrolateral distal angle with three acute spines. Left cheliped with few more spinules on dorsal surface of palm. Carpus with two strong corneous-tipped spines on dorsomesial margin and blunt protuberance in line with tubercle on proximal margin of palm. Ventromesial margin of merus with row of few small spines or two small spines distally; lateral face with two spines at ventrolateral distal angle. Ambulatory legs similar from left to right. Dactyls approximately 1.5 times longer than propodi; dorsal surfaces flattened; ventral margins each with row of minute corneous spinules (10-13 in distal half and one or two proximally). Lateral faces of propodi each with very strongly developed dorsolateral margin; dorsal surfaces somewhat flattened; ventrolateral distal angles each with one or two spines. Carpi each with spine at dorsodistal angle. Meri each with acute spine at ventrolateral distal angle. Sternite of third pereopods with subrectangular anterior lobe, anterior margin slightly rounded. Fourth pereopods each with acute spine at dorsodistal margin of carpus. Telson with slightly asymmetrical posterior lobes, separated P.A. MCLAUGHLIN by small median cleft; terminal margins each with three to five small spines, larger on left. REDESCRIPTION Shield longer than broad; anterior margin between rostrum and lateral projections straight or very faintly concave; anterolateral margins sloping; posterior margin roundly truncate. Dorsal surface of shield with scattered setae and distinct “Y’-shaped suture medianly in posterior portion. Rostrum triangular, terminating acutely or minutely bifid, overreaching lateral projections. Lateral projections broadly rounded or obtusely triangular, with terminal spinule or blunt small projection. Ocular peduncles slender, 0.66—-0.75 length of shield, with scattered setae dorsally and mesially; corneae not dilated. Ocular acicles narrowly triangular, dorsally rounded (convex), with simple or bifid acute termination; separated by less than basal width of one acicle and tending to become approximate distally. Antennular peduncles overreaching distal margin of corneae little if at all. Ultimate and penultimate segments with scattered setae. Basal segment with one or two very small spinules on ventrolateral distal margin. Antennal peduncles reaching approximately to bases of corneae; with supernumerary segmentation. Basal segment without spine on laterodistal margin, but with acute spine on ventrodistal margin laterally. Second segment with dorsolateral distal angle produced, with small terminal spine, dorsomesial distal angle rounded. Third segment with small spine on ventrodistal margin. Fourth and fifth segments with scattered setae. Antennal acicle reaching almost to distal margin of fourth peduncular segment, triangular, armed on mesial margin with one to three spines and tufts of setae, terminating in acute spine. Antennal flagellum overreaching chelipeds, and approximately as long as ambulatory legs; each article with two or three minute bristles. Chelipeds subequal, right somewhat larger. Right cheliped with dactyl equal to or slightly longer than palm; dorsomesial margin with row of corneous-tipped spines, dorsal surface with two rows of appreciably stronger, corneous-tipped spines; surfaces all with numerous tufts of moderately long setae; cutting edge with two prominent calcareous teeth in proximal half and broad terminal corneous hoof-shaped claw. Palm slightly longer than carpus; dorsomesial margin with row of low spines and tufts of long setae, second adjacent row of spines and one large tubercle at proximal margin, dorsal surface sloping, with no delimitation of dorsolateral margin, surface with few spines distally and on proximal portion of fixed finger, also with tufts of long setae; fixed finger with two rows of small spines on dorsal surface, all surfaces with tufts of long setae; cutting edge with three calcareous teeth in proximal half, distal-most strongest; terminating in corneous hoof-shaped claw. Carpus slightly more than half length of merus; dorsomesial distal angle with acute spine, low protuberances and tufts of setae on dorsomesial margin; dorsal surface with indications of points of original tufts of setae (no longer present), dorsolateral margin not delimited; mesial face with few scattered tufts of setae. Merus subtriangular; dorsal margin with tufts of setae; ventromesial margin with few low, somewhat spinulose protuberances and tufts of setae; ventrolateral distal angle with three acute spines, ventrolateral margin proximally and lateral REASSESSMENT OF *‘CALCINUS*‘ ASTATHES 33 | a 5 a - wy) Fig. 1 Clibanarius astathes (Stebbing, 1924), A-E, female lectotype (7.3 mm); F, male paralectotype (6.9 mm). A, shield and cephalic appendages; B, right second antennal peduncular segment enlarged; C, carpus and chela of left cheliped (lateral view); D, third left pereopod (lateral view); E, dac- tyl of third left pereopod (mesial view); F, telson. Scales equal 5.0 mm (A, C-E) and 3.0 mm (B, F). 34 face ventrally with low protuberances and tufts of setae. Ischium with few low spinules on ventromesial margin. Left cheliped with few more spinules on dorsal surface of palm; carpus with two strong corneous-tipped spines on dorsomesial margin and blunt protuberance in line with tubercle on proximal margin of palm; ventromesial margin of merus with row of few small spines or two small spines distally and two, plus low protuberance, medially, with additional protuberance proximally; lateral face with two spines at ventrolateral distal angle and low protuberances on ventrolateral margin. Ambulatory legs similar from left to mght. Dactyls approximately 1.5 times longer than propodi; in dorsal view straight, in lateral view slightly curved; dorsal surfaces flattened, and with rows of tufts of stiff setae, lateral faces each with longitudinal row of tufts of stiff setae; ventral margins with row of tufts of stiff setae and row of minute corneous spinules, 14 (spaced distally to proximally 8,5,1) on left third (lectotype), right third with 10, left second with 13, all in distal half; paralectotypes usually with 10—13 spinules in distal half and one or two proximally. Propodi slightly less than twice length of carpi; lateral faces each with distinctly developed dorsolateral margin and tufts of setae; dorsal surfaces somewhat flattened and with numerous tufts of setae; ventrolateral distal angles each with one or two spines, ventral margins with tufts of setae. Carpi 0.66 to 0.90 length of meri; each with spine at dorsodistal margin and tufts of setae on dorsal and lateral faces. Meri with tufts of setae dorsally and ventrally; ventrolateral distal angles each with one acute spine and ventral margins of second also with low protuberances. Ischia with tufts of setae on ventral margins. Fourth pereopods each with acute spine at dorsodistal margin of carpi. Sternite of third pereopods with subrectangular anterior lobe, anterior margin slightly rounded. Uropods asymmetrical. Telson with transverse suture; posterior lobes slightly asymmetrical, separated by small median cleft; terminal margins each with three to five small spines, larger on left. CoLourR. Unknown. DISTRIBUTION. At present recognized only from the type locality, Delagoa Bay, South Africa. P.A. McLAUGHLIN AFFINITIES. Clibanarius astathes shares with C. padavensis De Man, 1888, dactyls of the ambulatory legs that are longer than the propodi. Barnard (1926) listed the latter species from Delagoa Bay, and subsequently (Barnard, 1950) gave a brief diagnosis of that species. The ratio of cornea diameter to ocular peduncle length cited by Barnard is less than that given by De Man (1888) in his original description, or later by Alcock (1905), and the illustrated propodus of the left third pereopod (Barnard, 1950: fig. 80d) is appreciably shorter and stouter than that described for C. padavensis. However, if the colour patterns described by Barnard actually were taken from his specimens, one must assume that his identification was correct, and that C. astathes and C. padavensis exist sympatrically in the Delagoa area. I have not had the opportunity to examine Barnard’s (1950) specimen(s) of C. padavensis, nor other specimens of this species; however, De Man’s (1888) very detailed description points to several characters that would distinguish C. padavensis from C. astathes in the absence of colour. These include longer and more slender ocular peduncles; multispinose ocular acicles; longer antennal acicles, which reach beyond the proximal margins of the penultimate peduncular segments; longer and more slender chelae; and subcylindrical propodi of the ambulatory legs. Clibanarius astathes also bears a very strong resemblance to C. longitarsus. Clibanarius astathes differs from the Durban Bay population of C. /ongitarsus (12 males, 3 females, SL = 1.9-10.4 mm) that I have examined, in having: 1) fewer spines on the ventrolateral distal margin of the antennular peduncle (one or two, as opposed to three to six in C. longitarsus); 2) two strong corneous-tipped spines on the dorsomesial margin of the carpus of the left cheliped (only one was observed in numerous specimens of C. Jongitarsus of varying sizes); 3) strongly delineated dorsomesial propodal margin on the third pereopod (rounded or very faintly ridged in C. Jongitarsus); 4) roundly rectangular anterior lobe on the sternite of the third pereopods (this lobe is subquadrate, and often with a central blister-like protuberance in C. /ongitarsus). Other characters, such as the rows of spines on the fixed finger of the left cheliped, the armature of the ventral margins of the meri of the chelipeds, the Fig. 2 Clibanarius astathes (Stebbing, 1924), male paralectotype (6.9 mm). A, ieft cheliped; B, right cheliped. Scale equals 12 mm. REASSESSMENT OF ‘CALCINUS* ASTATHES presence of a spine at the ventrolateral distal angle of the merus of the third right and left pereopods, the number and spacing of the corneous spinules on the ventral margins of the ambulatory dactyls, and the spination of the terminal margins of the telson appear quite variable in C. Jongitarsus. Too few specimens of C. astathes are known for any evaluation of morphological variation. REMARKS. Stebbing’s (1924) comments on the genus Calcinus emphasized as did his earlier remarks (Stebbing, 1914), the development of the maxillipeds, which seems to suggest that he was not really familiar with the overall morphology of Calcinus species. Finding similar maxillipedal development in his new taxon, Stebbing (1924) assigned astathes to this genus, with apparent disregard for the numerous characters which set Clibanarius apart from Calcinus. It also would appear that the telson of the specimen he described and illustrated (Stebbing, 1924: 240, pl. 2T) was not closely examined, as it was characterized and depicted as being a simple plate with a smoothly rounded terminal margin. In actuality, the telson has a slight transverse suture dividing it into anterior and posterior lobes; the posterior lobes are separated by a small median cleft, and the terminal margins each have a few distinct spines. Barnard (1950) was correct to transfer Stebbing’s (1924) taxon to Clibanarius, although this transfer was obscured by his placement of the species in synonymy with C. virescens. Calcinus _ astathes Stebbing, 1924 was still listed by Gordan (1956) in her comprehensive tabulation of hermit crab species, and more | recently was included by Morgan (1991) in his world-wide listing of known Calcinus species. Barnard’s (1950) decision regarding | the conspecificity of Clibanarius astathes and C. virescens appears to have been based on specimens incorrectly labeled by Stebbing, and not on the actual type material of C. astathes. Not | only the fact that the Natal-Mozambique areas of South Africa | are type localities of both species, but also the inadequacy of Stebbing’s (1924) original description and _ illustrations, / undoubtedly account for Barnard’s (1950) synonymy having been accepted, without question, by subsequent carcinologists. One of the principal characters upon which Barnard (1950) | based his identification of C. virescens was the shortness of the dactyls of the third pereopods in relation to the propodi, / although, as pointed out by Lewinsohn (1982), this character was not mentioned in Krauss’ (1843) original description of the species. Not only Barnard (1950), but Fize and Serene (1955), and Gherardi and McLaughlin (1994) reported that the dactyls of their specimens were shorter than the propodi; however, Buitendiyk (1937), Miyake (1978), and Lewinsohn (1982) described the dactyls and propodi as being equal in length. Rahayu and Forest (1993) found the dactyls longer than the propodi in small specimens, but shorter in large specimens. Stebbing (1924), like Krauss (1843), made no mention of the length ratios of the dactyls and propodi of the ambulatory legs of C. astathes. Despite the variations in this major diagnostic character }observed in C. virescens, C. astathes sensu stricto differs markedly from Krauss’ (1843) taxon. The dactyls of the ambulatory legs of C. astathes are approximately half again the length of the propodi. Additionally, the ventral margins of the ambulatory dactyls are armed with 10 to 14 tiny corneous spinules in C. astathes, in contrast to the five to eight strong corneous spines seen in C. virescens. As previously indicated, C. astathes bears a far greater similarity to that group of 33) Clibanarius species characterized by long pereopodal dactyls and very short antennal acicles. ACKNOWLEDGEMENTS. I am indebted to Paul Clark, The Natural History Museum, London, for the loan of Stebbing’s syntypic material, and to Barbara Cook and Liz Hoenson, South African Museum, Cape Town, for providing the specimens upon which Barnard based his synonymy. The photographs are to work of E.J. McGeorge. This a scientific contribution from the Shannon Point Marine Center, Western Washington University. REFERENCES Alcock, A. 1905. Anomura. Fasc. I. Pagurides. — Catalogue of the Indian decapod Crustacea in the collections of the Indian Museum 2: 1-197. Calcutta, Indian Museum. Barnard, K. H. 1926. Report on a collection of Crustacea from Portuguese East Africa. Transactions of the Royal Society of South Africa 13(2): 119-129. 1947. Descriptions of new species of South African decapod Crustacea, with notes on synonymy and new records. The Annals and Magazine of Natural History (11) 13: 361-392. 1950. Descriptive catalogue of South African decapod Crustacea (crabs and shrimps). Annals of the South African Museum 38: \-837. Buitendijk, A. M. 1937. Biological results of the Snellius expedition. IV. The Paguridea of the Snellius Expedition. Temminckia 2: 251-280. Dana, J.D. 1851. Conspectus crustaceorum quae in orbis_ terrarum circumnavigatione, Carolo Wilkes e classe reipublicae foederatae duce, lexit et descripsit. (Preprint from)Proceedings of the Academy of Natural Sciences, Philadelphia 5: 267-272. — 1852. Conspectus crustaceorum, etc., Conspectus of the Crustacea of the Exploring Expedition under Capt. Wilkes, U.S.N., including the Paguridea, continued, the Megalopidea, and the Macroura. Paguridea, continued, and subtribe Megalopidea. (Preprint from) Proceedings of the Academy of Natural Sciences, Philadelphia 6: 6-28 (1854). Fize, A. & Seréne, R. 1955. Les Pagures du Vietnam. Jnstitut Océanographique Nhatrang Note 45: ix, 1-228. Gherardi, F. & McLaughlin, P. A. 1994. Shallow-water hermit crabs (Crustacea: Decapoda: Anomura: Paguridea) from Mauritius and Rodrigues Islands, with the description of a new species of Calcinus. Raffles Bulletin of Zoology 42(3): 613-656. Gordan, J. 1956. A bibliography of pagurid crabs, exclusive of Alcock, 1905. Bulletin of the American Museum of Natural History 108: 253-352. Haan, W., De. 1833-1850. Crustacea. In: P.F. von Siebold, Fauna Japonica wive Descriptio animalium, quae in itinere per Japoniam, jussu et auspicilis superiorum, qui summum in India Batava Imperium tenent, suscepto, annis 1823-1830 collegit, notis, observationibus et adumbrationibus illustravit: 4: ix—xvi, vii—xvii, i—xxxi, 1-244, pls. 1-55. Lugdunum Batavorum. Krauss, F. 1843. Die Stidafrikanischen Crustaceen. Eine Zusammenstellung aller bekannten Malacostraca. Bemerkungen tiber deren Lebensweise und geographische Verbreitung, nebst Beschreibung und Abbildung mehrerer neuen Arten. 68 pp. E. Schweizerbart’sche Verlagsvuchhandlung, Stuttgart. Kensley, B. 1974. Type specimens of Decapoda (Crustacea) in the collections of the South African Museum. Annals of the South African Museum 66(4): 55-77. Lewinsohn, Ch. 1982. Researches on the coast of Somalia. The shore and the dune of Sar Uanle. 33. Diogenidae, Paguridae and Coenobitidae (Crustacea Decapoda Paguridea). Monitore Zoologico Italiano, n.s. supplement 16: 33-68. Man, J. G., De. 1888. Report on the Podophthalmous Crustacea of the Mergui Archipelago, collected for the Trustees of the Indian Museum, Calcutta, by Dr. John Anderson, F.R.S., Superintendent of the Museum, parts IV and V. Journal of the Linnean Society, London 22: 177-249. Miyake, S. 1978. The crustacean Anomura of Sagami Bay: 1-200 (English), 1-161 (Japanese). Hoikusha Publishing Co., Tokyo. Morgan, G.J. 1991. A review of the hermit crab genus Calcinus Dana (Crustacea: Decapoda: Diogenidae) from Australia, with descriptions of two new species. Invertebrate Taxonomy 5: 869-913. Rahayu, D.L. & Forest, J. 1993. Le genre Clibanarius (Crustacea, Decapoda, Diogenidae) en Indonésie, avec la description de six espéces nouvelles. Bulletin du Muséum National d'Histoire Naturelle, Paris [1992] (4) 14(A)(2): 745-779. Randall, J.W. 1840. Catalogue of the Crustacea brought by Thomas Nuttall and J.K. Townsend, from the west coast of North America and the Sandwich Islands, with descriptions of such species as are apparently new . . . Journal of the Academy of Natural Sciences of Philadelphia 8: 106-147. 36 P.A. MCLAUGHLIN Stebbing, T.R.R. 1914. Stalk-eyed Crustacea Malacostraca of the Scottish 1924. South African Crustacea (Part XII of S. A. Crustacea, for the Marine National Antarctic Expedition. Transaction of the Royal Society of Edinburgh, Investigations in South Africa). Annals of the South African Museum 19: 50(2): 253-307 (issued separately June 4, 1914). 235-250. 1920. South African Crustacea (Part X of S. A. Crustacea, for the Marine Investigations in South Africa). Annals of the South African Museum, 17(4): 231-272: Bull. nat. Hist. Mus. Lond. (Zool.) 62(1): 37-39 Issued 27 June 1996 On a new species of Ophidiaster (Echinodermata: Asteroidea) from southern China YULIN LIAO Institute of Oceanology, Academia Sinica, 7 Nanhai Road, Qingdao, P.R. China AILSA M. CLARK Formerly of The Natural History Museum, Cromwell Road, London SW7 5BD, UK Synopsis. A new species, Ophidiaster multispinus, from southern Chinese waters, is described. This was previously recorded by A.M. Clark (1982) and by Liao & Clark (in press) as Ophidiaster armatus Koehler, 1910 but both authors now believe that Chinese specimens are specifically distinct. : SYSTEMATIC DESCRIPTION shape, slightly convex, covered with numerous coarse granules, beeen EEE — density in the central part of the plates 12-18/mm~. Abactinal | plates on dorsal surface of arms arranged in three regular | Family OPHIDIASTERIDAE longitudinal series, together with the two marginal series each Genus Ophidiaster L. Agassiz, 1835 side forming seven regular longitudinal series as well as | i transverse ones. Papular areas distinctly arranged in eight | are Aer, longitudinal series but the lowest on each side with only a few | Ophidiaster multispinus sp. nov. pores in each area. No pedicellariae detected. All the marginals, | Fig. 1, pl. 1 except the basal three to five, armed with a fairly conspicuous ; short blunt spine, these forming a longitudinal series but missing Ophidiaster armatus: A.M. Clark, 1982:487, 490; Liao & Clark | é on occasional plates, leaving small gaps in the sequence. Some _ (in press): .[Non Ophidiaster armatus Koehler, 1910]. abactinal plates near the arm tips also bearing one or two small Ho.otyre. IOAS:E1070, Hainan Strait, southern China Spines but these are not at all conspicuous. | (20°15'N, 110°15'E), 55 m, rocky, collected July 10, 1960; Adambulacral plates with two furrow spines, those of paratypes IOAS-E-1071, 4 specimens from Xiamen (Amoy), consecutive plates not separated by granules on the vertical faces Fujian Province, 1975. of the furrows. A large cone-like subambulacral spine on each plate, set back from the furrow within the general granulation. DESCRIPTION. R (major radius) 55-70 mm; r (minor radius) 10 The most proximal six to eleven adambulacral plates with a |mm, br (arm breadth basally) 11 mm. Disc small, arms five, smaller supplemental series of subambulacral spines interposed unequal, cylindrical, only tapering slightly in the distal third to between the furrow spines and the main subambulacral ones. rounded tips. Abactinal plates large, more or less triangular in rr | . Fy - —- ‘e ‘eo nas : FEIN) seb exe geo), ari CO yy bts) sa bes ) ab ‘o Bo a © as 6 @, ®, ) © | Fig.1 Ophidiaster multispinus sp. nov., Holotype. Proximal portion of actinal surface showing: (a) furrow spines; (b) supplemental subambulacral spines; (c) subambulacral spines; (d) actinal spines. The mouth is towards the right. The scale bar = 2 mm. © The Natural History Museum, 1996 i] Y. LIAO AND A.M. CLARK 068 a sot 8 a ze a Plate 1 Ophidiaster multispinus sp. nov. Holotype: (a) dorsal view; (b) ventral view. | NEW SPECIES OF OPHIDIASTER Four or five actinal plates in each intermediate area, armed with single spines corresponding to the subambulacral spines (Fig. 1). Colour (in the dried state) yellowish with faint darker bands. In addition to the holotype, there are four paratypes from Amoy, Fujian Province, far to the east of Hainan Strait. These are much larger than the holotype (which was selected for better comparison with the type material of Ophidiaster armatus) and have R 95, 105, 108 and 115 mm. Three of them have pedicellariae on some abactinal plates. Four other large specimens (R c. 130 mm), from the east side of Hong Kong Island in c. 15 m, studied by A.M.C., show some variation in the extent of the marginal spines, which are most numerous on the inferomarginals and may extend almost to the interradius. REMARKS. This new species was previously referred to Ophidiaster armatus Koehler, 1910 by A.M. Clark (1982) and by Liao & Clark in ‘The echinoderms of southern China’ (in press), both references commenting on the unusually large size. During a visit to the Senckenberg Museum by the senior author in 1993, a direct comparison was made between the small Chinese specimen and the three syntypes of O. armatus from the Aru Islands, Indonesia, of similar size. As a result of this comparison, we conclude that the Chinese specimens are specifically distinct, the differences being evident as shown in Table 1. The maximum size for Ophidiaster armatus, R 66 mm, was observed by H.L. Clark (1938) in material from Queensland, Australia. It is noteworthy that all three syntypes of O. armatus do have some abactinal pedicellariae which Koehler seems to have overlooked, however, the occurrence of pedicellariae is rarely regarded as a character of specific weight in this family. The presence of spines on plates other than the adambulacrals, separates Ophidiaster armatus and O. multispinus from the remaining species of the genus Ophidiaster. 39 Table 1 Comparison between the two species mentioned. Ophidiaster multispinus sp. nov. O. armatus Koehler Maximum R 130 mm. Maximum R 66 mm. Arms stout, br 11 mmat R60 mm. Arms slender, br 9 mm at R 60 mm. Marginal spines fairly conspicuous Marginal spines not at all and more extensive, absent only conspicuous, only present from the first 3—5 plates towards the arm tips Proximal adambulacrals with two Subambulacral spine series series of subambulacral spines single throughout 4 or 5 spinose actinal plates in intermediate area No spinose actinal plates However, in our opinion, this character is not of sufficient importance to justify a generic distinction, especially as the lesser development in O. armatus provides an intermediate condition. REFERENCES Clark, A.M. 1982. Echinoderms of Hong Kong, pp. 485-500. Jn: B.S. Morton & C.K. Tseng (Eds) The marine flora and fauna of Hong Kong and southern China. (Hong Kong, Hong Kong University Press). Clark, A.M. & Rowe, F.W.E. 1971. Monograph of shallow-water Indo-West Pacific echinoderms. London: British Museum (Natural History), 1x + 238 pp. Clark, H.L. 1938. Echinoderms from Australia. Memoirs of the Museum of Comparative Zoology, Harvard 55: viii + 596 pp. Koehler, R. 1910. Asteries et Ophiures des iles Aru et Kei. Abhandlungen Senckenbergischen Naturforschen dem Gesellschaft, 33(3): 265-295. Liao, Y. & Clark, A.M. (in press). The echinoderms of southern China. Bull. nat. Hist. Mus. Lond. (Zool.) 62(1): 41-70 Issued 27 June 1996 The life cycle of Paracyclops fimbriatus (Fischer, 1853) (Copepoda, Cyclopoida) S. KARAYTUG Department of Zoology, The Natural History Museum, Cromwell Road, London SW7 5BD, UK, and School of Biological Sciences, Queen Mary and Westfield College, Mile End Road, London EI 4NS, UK G. A. BOXSHALL Department of Zoology, The Natural History Museum, Cromwell Road, London SW7 5BD, UK CONTENTS MURA CULO erect este meet crocs sas awe tnevsssossys aur oxeesreo mesa aecten eet anc aeacivcaua coors rosnas ip Ses acgat ce couss Saeete races sate eeceac ee eR ees 41 IWMI OS eetee cece sears os bach eacucctsvascysccw ex susuneueuteasceusetuasdecscdscaeteessuccssssedarsteeaeece ssastcuncpavsOTouadesuds cen MOmeeaE eR aoe ete seco 4] DESCRIP tomo fy NAIIATHS Ca CES) tee ceen case c cease metre ene nee eee ees conc senate cuvsutiocscuncneseecusshoceanestetsexsucossvuraunesednae: seneeetteaverasnees 42 DESGHIPMOMOMCOMSDOGICS ta GES saecse See ses ces esate cscs tat aaaaes cae sqncnadcste Seceaedn Su cneioucie\iesudacssudeentcscadeddaetsudsevunedsaceeessounaNeneived 48 NOIYSULS SION secaeeg cs Ses 2S- Sec ce oes le sage guecedie Tstes Snanacd cds eatets idcine ands spieesmpera'vucup sales Rene. PeUGa. ant capees ccm auaeee, Re ecaeoey lake ae eal 67 | PRGKTIOW LEGO CIION US: nce, toca races tas ee garae cceme scence scene ceW usec lose cats Sawiad oe Mean aedsansaca ciovlencte's Soduceets vat cms ws cases Sazttade Coates See 69 FRE LG LEM COS ira ectec teen cases tren saticteacec fon reeceracnancanecusset eves sonengavee cass deez esas veee duccase pa euwerne ducee iu une sy tocmaaiie ceva inecaaaat ene tanauean cet Rao 70 . Synopsis. The complete life cycle of Paracyclops fimbriatus (Fischer, 1853) (Copepoda, Cyclopoida) is redescribed based on cultured material. All 6 nauplius and 5 copepodid stages plus the adults are described in detail. Scanning electron microscopy is used to elucidate the fine structure of some appendages, especially the male antennule, and to study body ornamentation patterns. Sexual dimorphism is first apparent at the copepodid I'V stage which has a more robust antennule | in the male than in the female. provide important information on the phylogenetic INTRODUCTION i | Paracyclops fimbriatus has been reported from a wide range of _ freshwater habitats and the geographical records summarised by | Dussart & Defaye (1985) indicate a cosmopolitan distribution. | It has also been regularly recorded from subterranean waters | (Ito, 1954; Pesce & Galassi, 1987) and from waters with a salinity | of 31%o(Loffler, 1961). Early reports of this species at great | depths in Lake Baikal (Mazepova, 1962) refer to the newly _ recognized P. baikalensis Mazepova, 1978 (Mazepova, 1978). | The genus Paracyclops comprises 24 species and subspecies | (Dussart & Defaye, 1985; Reid, 1987a, 1987b; Strayer, 1988). | The type species, P fimbriatus, was originally described by Fischer (1853), using material from St-Petersburg in Russia. | This taxon has been partially redescribed on numerous occasions (for example Kiefer, 1929; Gurney, 1933; Lindberg, 1941; Dussart, 1969; Einsle, 1971). Some descriptions of developmental stages have also been published; one of the most accurate being that of Gurney (1933) but even that gives j insufficient data to characterise the copepodid stages. Other descriptions have concentrated primarily on the naupliar instars (Gurney, 1933; Dukina, 1956; Ewers, 1930), but these all lack detail by modern standards. Although the general morphology of cyclopoid naupliar stages has been known for a long time it is | only recently that detailed naupliar descriptions of freshwater |cyclopoid copepods have been published (Dahms & Fernando | 1992, 1993, 1994). Systematic studies on naupliar stages can © The Natural History Museum, 1996 relationships among copepods (Dahms, 1991a,b). In the present paper P. fimbriatus has been redescribed in detail, including all its developmental stages, in order to provide a base for comparison with other Paracyclops species. METHODS Adult females of P fimbriatus were collected by plankton net from Regent’s Park pond (London) on 29.5.1994. Ovigerous females were kept in plastic dishes filled with fresh water filtered through a 35 uym-mesh net. Water was changed at 2 day intervals until the eggs hatched and the nauplius phase was completed. The adult females were removed after hatching of their egg sacs. Copepodid stages were placed in an aquarium tank in a constant temperature room. The nauplii were examined as whole mounts. Broken glass-fibres were used to prevent the nauplii from being compressed by the coverslip and to facilitate rotation which allowed viewing from all sides. Copepodids were dissected in lactophenol. All drawings were made with the aid of a camera lucida using Nomarski differential interference contrast. All measurements were made with an ocular micrometer. Body lengths were measured from the anterior to the posterior end of the body in nauplii, and from the base of the rostrum to the posterior edge of the caudal rami 42 in copepodids. Body width is given as the widest part of the nauplius body or the copepodid cephalothorax. In the spine and seta formula of the swimming legs Roman numerals and Arabic numerals are used for spines and setae respectively. When necessary scanning electron microscopy (SEM) was used to study fine details. Material for SEM was prepared by cleaning specimens in an aqueous solution (one drop in 100 ml of distilled water) of the surfactant detergent RBS pF for 30 min and subsequently sonicating in an ultrasonic cell disrupter for 7 s. Cleaned specimens were washed in 3 changes of distilled water for 5 minutes, dehydrated through a graded acetone series, critical point dried, mounted on aluminium stubs, sputter coated with gold and viewed under an Hitachi S-800 scanning elctron microscope. RESULTS Description of Naupliar Stages Nauplius I Body length : mean + standard deviation = 118 + 2.68 um (range 113 to 122 um, n=20), mean body width 75 + 1.5 um (range 72 to 77 um, n=20). Body oval (Fig. 1A), with posterior spinular row on ventral surface and on either side of caudal setae. Ventral surface with 2 paired patches of spinules posterior to labrum. Oval area of integument located posteriorly on ventral surface. Caudal rami represented by pair of naked setae. Antennule 3-segmented (Fig. 2A). First segment with | spinulose seta anteriorly and group of spinules along outer margin. Second segment with 2 setae, distal one spinulose and slightly shorter than proximal plumose one; ornamented with 2 spinular rows along ventral side. Third segment with 2 naked setae distally and 2 spinular rows along ventral margin. Antenna biramous (Fig. 3A), with 2-segmented protopod comprising coxa and basis. Coxal gnathobase represented by large spine armed with spinular row distally. Basis with 3 small setae along inner margin, 2 of them closely set together in proximal third. Exopod 4-segmented; segment | large with 1 spinulose seta and spinular row on outer margin; second segment with | long naked seta; third segment with | long naked seta and few tiny spinules on outer margin; fourth segment with naked seta proximally and | long spinulose seta plus | naked seta distally; outer margin with spinular row near proximal seta and single isolated setule apically. Endopod unsegmented; armed with 2 short inner setae and 2 long setae terminally. Mandible (Fig. 4A) biramous, with 2-segmented protopod comprising small coxa and large basis. Coxal gnathobase with 1 naked seta. Inner margin of basis with 1 spinulose and 1 naked seta; outer margin with minute spinule. Exopod 4-segmented; first to third segments each with | plumose seta at inner distal angle; segments 3 and 4 with spinular row along outer margin; apex of fourth segment with long plumose, inner seta and short, naked, outer seta, the latter about as long as segment. Endopod 2-segmented; first segment with two spinulose setae and spinular row On inner distal margin; second segment with 4 naked setae. Nauplius IT Body length : mean + standard deviation = 135 + 7.84 um (range 113 to 144 um, n=21), mean body width 88 + 4.5 um (range 83 to 100 um, n=21). Body similar to first nauplius but larger and elongated caudally (Fig. 1B); differing as follows: labrum (Fig. S. KARAYTUG AND G.A. BOXSHALL 19C) with some spinules along lateral margin. Caudal region lacking spinular row adjacent to caudal setae. Antennule (Fig. 2B) armed with third seta on inner terminal edge of distal segment. Antenna (Fig. 3B) with additional small seta at base of large sword-shaped coxal gnathobasic seta. Basis with distal spinular row. Exopod 6-segmented, with | extra seta located on minute second segment (arrowed in Fig. 16A). Terminal segment of first nauplius subdivided; segment five with 1 naked seta and additional spinular row distally. Endopod of antenna with 2 plumose setae of equal size and 1 shorter plumose seta terminally, plus 2 inner lateral naked setae. Coxa of mandible (Fig. 4B) with unilaterally spinulose seta. Additional naked seta present on inner margin of basis. First segment of endopod with one naked seta and one longer plumose seta but with no spinular row on inner distal margin. Second segment of endopod with 4 naked setae and 1 plumose seta. Base of first exopodal segment with additional naked seta. Maxillule appearing as strong plumose seta (arrowed in Fig. 1B). Nauplius IIT Body length : mean + standard deviation = 158 + 6.08 um (range 155 to 168 um, n=14), mean body width 105 + 8.7 um (range 88 to 117 um, n=14). Body (Fig. 1C) similar to second nauplius but larger, differing as follows: lateral spinular row on labrum consisting of much stronger spinules. Caudal margin with pair of strong plumose setae adjacent to longer naked setae. Posterior end of body trilobate. Third segment of antennule (Fig. 2C) acquiring extra naked seta on inner margin. Sixth exopodal segment of antenna (Fig. 3C) with | naked inner seta, 1 strong spinulose subapical seta and | long spinulose apical seta. Mandible with minor changes in relative lengths of endopodal setae (Fig. 4C). Nauplius IV Body length : mean + standard deviation = 181 + 8.4 um (range 166 to 192 um, n=14), mean body width 125 + 8.7 um (range 113 to 144 um, n=14). Body (Fig. 1D) pear-shaped, differing from Nauplius III as follows: caudal margin with pair of minute spines representing Anlagen of caudal setae. Posterior end of body trilobate and becoming elongated. Antennule (Fig. 2D) with a longer row of dentiform spinules distally on first segment. Third segment with 2 additional naked setae, plus extra row of long spinules midway along lateral margin. Coxa of antenna (Fig. 3D) with spinular row midway along outer margin. Basis with 4 setae along inner margin; outer margin with 2 spinular rows on anterior surface and | spinular row on posterior surface. Apical segment of exopod with spinular row distally. Basis of mandible with few mid-anterior surface spinules. Fourth segment of exopod with additional small seta distally (Fig. 4D). Maxillule (Fig. 1D) bilobed with 2 spinulose and | naked seta on inner lobe. Outer lobe with | spinulose, 1 plumose and | long naked seta. Nauplius V Body length : mean + standard deviation = 210 + 8.49 um (range 194 to 226 um, n=14), mean body width 138 + 12.08 pm (range 122 to 157 um, n=14). Body (Fig 1E) similar to nauplius IV, differing as follows: third pair of caudal setae larger. LIFE CYCLE OF PARACYCLOPS 43 ptt => —S oS sae’ — eG SS a ( / oe a(S ee AS \ a) > AAYL y) ee] ) =I \ eel P. fimbriatus. Naupliar stages, ventral view. A, Nauplius I; B, Nauplius II; C, Nauplius III; D, Nauplius IV; E, Nauplius V; F, Nauplius VI. Fig. 1 Antennules, antennae and mandibles omitted. Scale bar in pm. 44 S. KARAYTUG AND G.A. BOXSHALL \ \ 19 91 Lee en ee Fig.2 P. fimbriatus. N aupliar antennules, ventral view. A, Nauplius I; B, Nauplius II; C, Nauplius III; D, Nauplius IV; E, Nauplius V; F, Nauplius VI. Scale bar in ym : LIFE CYCLE OF PARACYCLOPS 45 | Fig.3 P fimbriatus. Naupliar antennae, ventral view. A, Nauplius I; B, Nauplius II; C, Nauplius III; D, Nauplius IV; E, Nauplius V; F, Nauplius VI. Scale bar in um. 46 S. KARAYTUG AND G.A. BOXSHALL Fig.4 P. fimbriatus. Naupliar mandibles, ventral view. A, Nauplius I; B, Nauplius II; C, Nauplius III; D, Nauplius IV; E, Nauplius V; F, Nauplius VI. Scale bar in pm. LIFE CYCLE OF PARACYCLOPS 47 D WW Me a \ SFE , a Val A y XXI-XXIII vor ae Soran Fig.21 P fimbriatus. Male antennule. A, dorsal view. B, Detail of segments XI to XVI, anterior; C, Detail of segments XIII to XX, lateral view. Scale bar in pm. ail a Fig.22 Scanning electron micrographs of P. fimbriatus. Adult male antennule. A, Dorsal view, with segment 5 arrowed; B, Lateral view, with sheath on segment XV arrowed; C, Modified seta on proximal segment; D, Segment proximal to geniculation, with pore on modified element arrowed. Scale bars A = 30 um, B = 25 um, C = 6 um, D = Spm. (range 226 to 247 um, n=10). Body (Fig. 17C) differing from adult female as follows: urosome 6-segmented, comprising fifth pedigerous, genital and 4 free abdominal somites; ornamentation of spinule rows as in Fig. 14D. Genital somite bearing paired genital apertures ventrally (Fig. 14C). Caudal rami about 2.5 times longer than broad, ventral S. KARAYTUG AND G.A. BOXSHALL surface unornamented. Large medial seta (V) relatively longer than in female. Antennule 15-segmented (Figs 21A-C). Segment 1 armed with 9 setae; 1 seta large and modified (Fig. 22C), by ornamentation of strong spinules in proximal and mid sections, tapering to fine point distally; segment ornamented with curved LIFE CYCLE OF PARACYCLOPS row of long spinules and row of minute spinules. Segment 2 with 4 setae. Segment 3 with 2 setae. Segments 4 and 5 partly fused, with segment 5 (arrowed in Fig. 22A) defined only on dorsal side, not ventrally. Segment 4 with 2 setae; segment 5 with 2 setae. Segment 6 with 2 setae. Segments 7, 8 and 9 separated from each other by extensive arthrodial membrane: segment 7 with 2 setae, segment 8 with 2 setae, and segment 9 with 2 setae plus aesthetasc. Segment 10 (= ancestral segment XV) produced on one side into extensive sheath (arrowed in Fig. 22B) enclosing segment 11 ventrally: armed with 2 setae, | ornamented with long spinules unilaterally. Segment 11 bearing curved seta ornamented with double row of strong denticles, plus 1 naked seta. Segment 12 partly fused to segment 13; armed with short seta ornamented with 2 rows of fine spinules, plus short naked seta. Segment 13 partly subdivided by partial suture: armed with short spinulate seta proximally, 2 short naked setae, plus | modified element attached to segment by short stalk, main part of element lying along surface of segment and ornamented with longitudinal ridges and small central pore (arrowed in Fig. 22D). Geniculation located between segments 13 and 14. Segments 14 and 15 partly fused, forming curved subchela-like section: segment 14 armed with | seta, 1 aesthetasc and 2 modified elements each ornamented with longitudinal ridges and a central pore, as distal element on segment 13. Apical segment tapering distally; armed with 11 setae and 1 aesthetasc, mostly originating on outer (= posterior) surface. All other appendages as in female except for fifth (Fig. 14E) and sixth legs (Fig. 14F). Outer spinulose seta of leg 5 ornamented with some long setules distally. Sixth legs forming opercular plates bearing row of large spinules along ventral surface; armed with | inner spine, | well developed spinulose seta and | inner naked seta. DISCUSSION The number of naupliar instars in the Cyclopoida has been the subject of some controversy but it is clear, as Elgmork & Langeland (1970) strongly indicated, that there are normally 6 naupliar instars. This is supported by recent works on free-living freshwater Cyclopoida by Dahms & Fernando (1992, 1993, 1994) and by our data. The most difficult distinction is between nauplius IV and V and these stages have often been confused. The complete naupliar sequence of P fimbriatus was previously described by Ewers (1930) and Dukina (1956). Nauplius I and II were also described by Gurney (1933) but none of these provides setation counts of sufficient accuracy. Ewers (1930) described 6 stages but our descriptions differ as follows: antennule is 3-segmented not 4-segmented; antennary exopod is 4-segmented at N I and becomes 6-segmented, rather than remaining 4-segmented; caudal rami of N IV are represented by 2 pairs of setae and 1 pair of minute seta, rather than just 2 pairs of setae; ventral body surface is ornamented with spinular rows throughout the nauplius phase. In general, however, Ewers’ drawings are so small that it is not worthwhile making detailed comparisons of appendage setation patterns. Six naupliar stages were also described by Dukina (1956). Our descriptions differ from Dukina’s as follows: antennule is 3-segmented; antennary exopod is 4-segmented at N I (given as 5-segmented by Dukina). Dukina’s descriptions of appendage setation also lack sufficient detail for meaningful comparisons. Gurney (1933) described the first 2 naupliar stages, but our 67 findings indicate that the caudal rami of N II are represented by a pair of setae not by 2 pairs as illustrated by Gurney. Apart from this discrepancy our results differ only in that the exopod of antenna is described as 3-segmented rather than 4-segmented as in our material. The copepodid stages of P fimbriatus were also partly described by Gurney (1933). Although our results are in substantial agreement, for example, with antennulary segmentation throughout the copepodid phase, Gurney’s drawings are not sufficiently accurate to permit comparisons of segmental setation. Analysis of the antennulary setation patterns of adult male Paracyclops permits the identification of the pattern of segmental homologies. The basic armature of each antennulary segment in copepods is 2 setae plus one aesthetasc (Giesbrecht, 1892), with a few exceptions as identified by Huys & Boxshall (1991). Using this basic pattern, the 9 setae on the first segment of male Pfimbriatus indicate that it can be identified as representing 5 ancestral segments (segments I-V). The second segment, with 4 setae, can similarly be identified as derived from 2 ancestral segments (VI-VII). The third to twelfth segments all represent single ancestral segments (VIII to XVII), as indicated by the presence of a maximum of 2 setae on each. The fourth and fifth are incompletely separated (Fig. 22A) but we have treated them as distinct. The thirteenth segment has only 4 setae but is here identified as representing 3 ancestral segments (X VIII-XX). This decision is based on the presence of the neocopepodan geniculation between the thirteenth and fourteenth segments which unequivocally identifies the segmental boundary involved as XX to XXI, and on comparison with other cyclopids such as Euryte robusta Giesbrecht, 1900. In E. robusta males segment XVIII is separate and carries a long naked seta and a short spinulose seta, segment XIX—XX carries a short spinulose seta and a modified spine proximally and a slender seta distally (Huys & Boxshall, 1991). In P fimbriatus the proximal part of the triple segment is defined by a partial suture marking the original plane between segments XVIII and XIX—XX. This part carries only a single spinulose seta and lacks the long seta; the distal part representing XIX—XX carries the same setation as in E. robusta. This confirms our interpretation of the thirteenth segment as a triple segment (XVIII-XX). The fourteenth and fifteenth segments, lying distal to the geniculation represent ancestral segments XXI-XXIII and XXIV—-XXVIII, exactly as Huys & Boxshall (1991) found for E. robusta. Compound antennulary segments, such as the first and second segments of P fimbriatus, were simply referred to as ‘fused’ by Huys and Boxshall (1991) in their comparative analysis of antennulary segmentation patterns in all copepod orders, although such compound segments could be the result of two different developmental processes: 1) secondary fusion of segments that were separated earlier during ontogeny 2) failure of separation during development. The compound first and second segments of the male antennule of P fimbriatus result from the second process, the failure to separate. In contrast, the compound apical segment results from the secondary fusion of the sixth, seventh and eighth segments of the copepodid V stage. (The eighth segment of the copepodid V was already a compound segment, representing three ancestral segments XX VI-X XVIII which are not separately expressed by any known member of the order Cyclopoida). Vertical tracking of the segmental boundaries as identified by their setation elements allows us to identify the homologies of [ape = '> ‘(1661) 1eusxog 2p skny Aq pasodoid anuusjue [esyssour pajuowsas-g7 jeonoyodAy wory suUOIRALJOp ayeoIpuT s[esdUINU ULUIOY] ‘SodvIS [[e UI (ae) SOSe}OY}SoR pure soutds ‘avjaS JO SIOQUINU SP [JOM SP a[RLL ][NPe UT YIPaYs puK UONL[NOIUSd Jo UOIRIO] SUIMOYS ‘saxas YI0g JO sode}s pipodadoo [jv ut sa1Zojowoy ye}UowSes Arejnuusjue Jo UOsIedwOD ONeWIDYIg snIDIuquuf gq €7 “BIA S. KARAYTUG AND G.A. BOXSHALL IWAXX ~AIXX IXX-IXX XX=IIAX WAX IAX AX AIX IIIX IX IX X=XI IIA sMWA-IA A- | ae my [ee th | [oere] | + |lel elle} El el El eel El Le] Le pi ee ees eed | ee WAXX-IAXX NXX ANIXX —_IIIKX-IXX XX — AX A\X— IX IX -IA IN= | ing [8242] Le] le] Petal emeetcieneee |2 E.G 2 ro [ee+2] [2] fe pere| 7 b No oe+ Z fe Vv i IN oe+ 2 |e | ee+ Ilo oe+ ZL |e | V eS G ] pace eases): Maere fee UG. e | > LIFE CYCLE OF PARACYCLOPS the segments of the female antennule and of the antennules throughout the copepodid phase of development. These results are presented in schematic form in Figure 23 which visually indicates homologous segmental boundaries expressed in successive copepodid stages. The aesthetasc derived from ancestral segment XXI provides a reference point throughout the copepodid phase: from the third segment of the first copepodid, to the fifth segment of the adult female and the fourteenth segment of the adult male. This, in concert with other setation features, confirms the unequivocal identification of the XX to XX] articulation (marked by the large arrow) in all stages. Identification of the homology of the more proximal segments relies on the presence of 11 setation elements on the third segment of the male copepodid V stage. These 11 elements testify to the segment’s derivation from 6 unseparated ancestral segments (X V-XX) and indicates that the boundary between the second and third segments represents the XIV to XV articulation. Similarly, the presence of 5 setae on the second segment indicates that the boundary between first and second segments represents the XI to XII articulation. The remaining segmental homologies in all stages, including the adult female, can now be identified by tracking vertically through the male stages back to copepodids I and II and then forwards again to the adult female. The seventh segment of the female antennule is identified as homologous with a single ancestral segment (XXV). The presence of 3 setae on this segment therefore requires explanation since the basic armature of segment XXV is 1 posterior margin seta plus | seta and | aesthetasc on the anterior margin (Huys & Boxshall, 1991). We suggest that one of the anterior elements may be a setiform aesthetasc and this should be investigated ultrastructurally. This is the first time that the homologies of all antennulary segments have been identified in all copepodid stages of both sexes of any cyclopoid copepod, with reference to the hypothetical 28-segmented antennule of the ancestral copepod (Huys & Boxshall, 1991). Within the family Cyclopidae antennulary segmental numbers are frequently reduced and, indeed, many genera are characterised by their low segment numbers. The possible involvement of heterochrony in the reduction of antennulary segmental numbers has already been noted. Gurney (1933) suggested that the reduced numbers of antennulary segments in some species may be accounted for as the persistence of a larval character in the adult’, pointing out that during development the last copepodid (Co V) typically has an 11-segmented antennule and that this number is commonly found in adults, as for example in Microcyclops. Several authors working on cyclopids were aware of the importance of determining homologies between the different generic segmentation patterns and made detailed comparisons. Gurney (1933) reviewed earlier work by Claus (1893), Manfredi (1923), Lucks (1929) and Gelmini (1928) on the sequence of segmental subdivision during development in Cyclops species. He presented a tabular system showing derivations of a 17-segmented antennule and confirmed this pattern using his own data on C. strenuus. The strict determination of ancestral homologies of antennulary segments provides new insight into the rather confused world of cyclopid systematics at the generic level. The scheme of segmental development (Fig. 23) indicates that at the moult from copepodid II to copepodid III it is the second segment that subdivides. This contrasts with the scheme presented for P fimbriatus by Gurney (1933) who showed the first segment subdividing at this moult. We base our interpretation on the positions and relative lengths of the setae 69 on these segments. In particular a relatively long seta is positioned at the anterodistal angle of the first segment in copepodid II and 2 similarly long setae are present at the same position in copepodid III. Since no other setae of this length are present we interpret this as evidence of the constancy of the boundary marked by this seta and therefore conclude that the second segment has subdivided. The setation of the second endopodal segment of the antenna increases progressively through the copepodid stages of P fimbriatus in a very regular manner. The first copepodid stage possesses 4 setae distributed around the inner-distal angle of the segment (Fig. 8A). The seta which is located on the angle is slightly stouter than the other 3 and is here identified as seta VIII, using the numbering scheme proposed for another member of the family Cyclopidae by Boxshall & Evstigneeva (1994). The more distal seta is then identified as seta [IX and the 2 more proximally located setae on the inner margin, as setae VII and VI. These 4 setae are presumably homologous with the proximal group of 4 setae present on the margin of the unsegmented endopod of the sixth nauplius stage (Fig. 3F). At each successive moult through the copepodid phase one additional seta is added proximally on the inner margin of the segment (Figs 8A-F). Thus at copepodid II a fifth seta (seta V) is added, at copepodid III a sixth seta (seta IV), and so on, until the final moult to adult (= copepodid VI) at which the ninth and final seta (seta I) is added. The progressive development of the setation suggests that reductions in numbers of setae on the second endopodal segment, which are common within the family Cyclopidae, may be interpreted as resulting from heterochronic events. Cryptocyclops bicolor (Sars, 1863), for example, is a cyclopid with only 7 setae on the second endopodal segment. This number is typical of the copepodid IV stage in both Paracyclops and the presumed ancestral cyclopid stock, and may be interpreted as evidence that a neotenic event within the Cryptocyclops lineage has interrupted the progressive addition of setae. The ontogeny of copepodid stages and examination the patterns of leg formation offer some evidence for inferring copepod phylogenetic relationships (Ferrari, 1988). A common pattern of development for legs 14, exhibited by at least 20 genera, was recognized by Ferrari. The pattern of development for the swimming legs of P. fimbriatus is in accordance with this common pattern as follows: Legs 1 2 3 4 N iB: 1°B I 1+1; 1+1; 1°B Il 2+2: 242; 1+1; 1B Ill 2+2: 2+2: 242; 1+1; IV 2+2- Mar) 2+2; Marre V 33: saree 343; 3+3: VI 3+3; 3+3; 3+3; 3+3; (Where N = nauplius; Roman numerals = copepodid stages; 1~B = primary leg bud; 1+1 = reorganized leg with 1-segmented exopod and endopod; 2+2 = leg with 2-segmented exopod and endopod; 3+3 = leg with 3-segmented exopod and endopod). ACKNOWLEDGEMENTS. We are grateful to Dr Rony Huys for his helpful comments on drawing techniques and on the manuscript. S. Karaytug would also like to thank Dr Steve Alston for his assistance in the laboratory. This research has been supported by a postgraduate grant from the University of Balikesir, Turkey to S. Karaytug. 70 REFERENCES Boxshall, G.A. & Evstigneeya, T. 1994. 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Each table should be typed on a separate sheet designed to extend across a single or double column width of a Journal page. It should have a brief specific title, be self-explanatory and be supplementary to the text. Limited space in the Journal means that only modest listing of primary data may be accepted. Lengthy material, such as non-essential locality lists, tables of measurements or details of mathematical derivations should be deposited in the Biological Data Collection of the Department of Library Services, The Natural History Museum, and reference should be made to them in the text. Illustrations. Figures should be designed to go across single (84 mm wide) or double (174 mm wide) column width of the Journal page, type area 235 x 174 mm. Drawings should be in black on white stiff card or tracing film with a line weight and lettering suitable for the same reduction throughout, either 50%, 30% or 25%. After reduction the smallest lettering should be not less than 10 pt (3 mm). All photographs should be prepared to the final size of reproduction, mounted upon stiff card and labelled with press-on lettering. Components of figure-plates should be abutted. All figures should be numbered consecutively as a single series. Legends, brief and precise, must indicate scale and explain symbols and letters. Reprints. 25 reprints will be provided free of charge per paper. Orders for additional reprints can be submitted to the publisher on the form provided with the proofs. Later orders cannot be accepted. CONTENTS 1 Deep-sea conolidean gastropods collected by the John Murray Expedition, 1933-34 A.V. Sysoev 31. Reassessment of ‘Calcinus’ astathes Stebbing 1924 (Crustacea: Anomura: Paguridea: Diogenidae) PA. McLaughlin 37. Onanew species of Ophidiaster (Echinodermata: Asteroidea) from southern China Y. Liao and A.M. Clark 41 The life cycle of Paracyclops fimbriatus (Fischer, 1853) (Copepoda, Cyclopoida) S. Karaytug and G.A. Boxshall Bulletin of The Natural History Museum ZOOLOGY SERIES Vo!. 62, No. 1, June 1996