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World List abbreviation: Bull. nat. Hist. Mus. Lond. (Zool.)

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

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

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

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

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

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.

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

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

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

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

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.

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

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.

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

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

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

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.

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

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.

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.

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

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.

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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

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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

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

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).

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

| . Fy - —- ‘e ‘eo nas

: FEIN) seb

exe geo), ari

CO yy bts)

sa bes

)

‘o

®,

)

| 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.

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.

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

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. 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

| 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

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

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 \

> AAYL

ee]

)

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

WW Me

a \ SFE

<i!

| Fig.5 P._fimbriatus. Dorsal view of Copepodid stages I-V. A, Female copepodid I; B, Female copepodid II; C, Female copepodid III; D, Male cope-

podid IV; E, Female copepodid IV; F, Female copepodid V; G, Male copepodid V. Scale bar in pm.

100

S. KARAYTUG AND G.A. BOXSHALL

a LLL

shee

AS WS

Oe ae

|Z WO= by

Fig.6 P fimbriatus. Urosome of female copepodids, dorsal. A, Copepodid I; B, Copepodid II; C, Copepodid III; D, Copepodid IV; E, Copepodid

V. Scale bar in um.

First segment of antennule (Fig. 2E) with spinular row on

outer margin. Third segment acquiring 3 small naked setae.

Endopod of antenna (Fig. 3E) with additional small naked seta

distally. Basis of mandible (Fig. 4E) with 4 setae. Maxillule (Fig.

1E) with 1 new naked seta on inner lobe.

Nauplius VI

Body length : mean + standard deviation = 241 + 8.49 um (range

255 to 224 um, n=16), mean body width 152 + 7.28 (range 143 to

162 pm). Body (Fig. 1F) similar to nauplius V, differing as

follows: innermost pair of caudal setae longest, middle pair of

setae plumose, shortest setae naked and laterally located.

Third segment of antennule (Fig. 2F) with total of 13 setae,

including 5 new setae, plus ornamentation of several long

spinules. Endopod of antenna with 9 setae (Fig. 3F), with | extra

seta in proximal group. Mandible (Fig. 4F) with minor changes

in ornamentation of basal setae.

Maxillule with 1 new seta on inner lobe, and with 3 spinules on

outer margin. Anlagen of post-maxillulary limb pairs apparent

on ventral surface of body (Fig. 1F).

Description of Copepodid Stages

Copepodid I

Body length : mean + standard deviation = 382 + 11.5 um (range

373 to 404 um, n=10), mean body width 146 + 5.7 um (range 134

to 152 um, n=10). Body 5-segmented (Fig. 5A), comprising

cephalothorax and 4 postcephalothoracic trunk somites. Second

trunk somite with 1 seta at posterolateral angles representing

third swimming legs. Antennule, antenna, mandible, maxillule,

maxilla, maxilliped and first and second swimming legs present.

Caudal rami (Fig. 6A) as broad as long. Each ramus armed with

6 setae, ornamented with spinular row on ventral surface,

outermost seta plumose with spinular row at base, 1 naked seta

located on dorsal surface posteriorly, 1 short naked seta close to

50 S. KARAYTUG AND G.A. BOXSHALL

Fig.8 P fimbriatus. Development of copepodid antennae. A, Copepodid I; B, Copepodid II; C, Copepodid II; D, Copepodid IV; E, Copepodid V;

F, Copepodid VI. Scale bar in um. [Setal numbering scheme of Boxshall & Evstigneeva (1994) is used.]

LIFE CYCLE OF PARACYCLOPS

terior. Scale bar in pm.

| outermost seta, 2 inner plumose setae distally and 1 naked

| seta on dorsal outer margin. Anal operculum ornamented

| spinular row.

Antennule 5-segmented (Fig. 7A); aesthetasc at antero-distal

_angle of apical segment sharing common base with adjacent

seta, as in all subsequent stages, including adult. Setal formula 3,

| 3,3 + 1 aesthetasc, 3, 7 + 1 aesthetasc.

Antenna 5-segmented (Fig. 8A); coxa unarmed; basis with

| vestigial exopod bearing 2 naked setae, one apically and one

halfway along inner margin. Basis armed with 2 inner angle

setae. Endopod 3-segmented; setal formula 1, 4, 6. Labrum (Fig.

| 9C) with strong denticles along mid posterior margin.

Mandible (Fig. 9A) with well developed gnathobase bearing

row of sharp blades medially, and 1 spinulose seta at inner distal

| angle. Vestigial mandibular palp with 2 long plumose setae and 1

5)II

—

Fig.9 P. fimbriatus. Copepodid I. A, Mandible, posteroventral; B, Maxillule, ventral; C, Labrum, ventral; D, Maxilla, anterior; E, Maxilliped, pos-

naked seta distally. Inner posterior margin of mandible with two

spinular rows.

Maxillule with strong praecoxa and reduced 2-segmented

palp (Fig. 9B). Praecoxal endite armed with 4 setae articulating

at base and 4 spines fused to segment. Proximal segment of palp

derived from coxa and basis, bearing 2 naked and | spinulose

inner margin setae, plus outer seta representing exopod. Distal

segment of palp representing endopod, armed with 3 naked

setae.

Maxilla (Fig. 9D) consisting of praecoxa, coxa, basis and

2-segmented endopod. Praecoxa with single indistinct endite

armed with 2 plumose setae. Coxa with inner seta at midlength

and distal endite bearing strong plumose seta and naked seta

apically. Basis with 3 setae distally. Endopod with 2 setae on

proximal segment and 3 setae terminally.

52 S. KARAYTUG AND G.A. BOXSHALL

WVU \

\\(\\ N

Fig. 10 P fimbriatus. Leg 1, anterior. A, Copepodid I; B, Copepodid II; C, Copepodid III; D, Copepodid IV; E, Copepodid V; F, Copepodid VI

(adult). Scale bars in pm.

LIFE CYCLE OF PARACYCLOPS 53

, OWA «~SN

BWR]. AX

Ait ~ SSS gail! “ry

gn /| WM Y, ert" \ WN

BS \\ Wiese \ & TS

Poni 1

| \ \

| Fig.11 P fimbriatus. Leg 2, anterior. A, Copepodid I; B, Copepodid I]; C, Copepodid II; D, Copepodid IV; E, Copepodid V; F, Copepodid VI

(adult). Scale bar in pm.

]

Goan lPPep,

Feat? we? ie

perenne Tree,

ELIE

SSS

S. KARAYTUG AND G.A. BOXSHALL

Fig. 12 P. fimbriatus. Leg 3, anterior. A, Copepodid II; B, Copepodid III; C, Copepodid IV; D, Copepodid V; E, Copepodid VI (adult). Scale bar in

um.

Maxilliped (Fig. 9E) distinctly 4-segmented, comprising

syncoxa, basis and 2-segmented endopod. Syncoxa armed with

2 spinulose setae. Basis with 2 setae. Endopod segment 1| with 1

plumose seta; segment 2 with 3 setae.

Swimming legs 1 (Fig. 10A) and 2 (Fig. 11A) each with

2-segmented protopod. Intercoxal sclerites unornamented.

Coxa of leg 1 with spinular row posteriorly. Basis of both legs

with outer angle seta ornamented by spinular row at base, inner

basal margin with fine spinules. Both legs with 1-segmented

rami. Inner margin of exopod and outer margin of endopod of

both legs with row of pinnules. Spine and seta formula as

follows:

LIFE CYCLE OF PARACYCLOPS

Coxa Basis Exopod Endopod

Leg] 0-0 1-0 IV,4 1,1+1,3

Leg2 00 1-0 IV,3

Copepodid IT

Body length : mean + standard deviation = 455 + 17.2 ym (range

439 to 488 um, n=10), mean body width 168 + 9.1 um (range 152

to 178 um, n=10). Differing from Copepodid I as follows: body

(Fig. 5B) 6-segmented, comprising cephalothorax and 5

postcephalothoracic trunk somites. Third trunk somite with 1

seta on outer margin representing fourth swimming legs. Last

somite with posterior margin spinular rows ventrally and

laterally (Fig. 6B).

Antennule with 3 spinules on first segment. Setal formula; 5, 4,

: 4+ | aesthetasc, 3, 7 + 1 aesthetasc (Fig. 7B).

Antenna 4-segmented (Fig. 8B); basis with spinular row near

inner margin. Inner setae set close together at distal angle.

' Vestigial exopod represented by spinulose seta. First endopodal

_ segment with | seta, second with 5 setae along distal margin and

_ bearing small spinular row on outer margin. Third endopodal

_ segment with 6 setae distally, ornamented with spinular row on

outer distal margin.

| Coxa of leg 1 with spinulose seta at inner distal angle (Fig.

| 10B). Basis with spinulose seta on inner distal margin

ornamented with spinular rows at base of inner seta and

| posteriorly, between exopod and endopod. Intercoxal sclerite

_ ornamented with setules distally. First segment of exopod with

| spinular row along outer margin. First and second segments of

endopod with pinnules along lateral margins.

Coxa of leg 2 (Fig. 11B) with naked seta on inner distal

| margin; ornamented with spinular rows proximally on outer

| margin of both anterior and posterior surfaces and along outer

| margin. First segment of exopod with spinular rows laterally

| and proximally on posterior surface. Exopodal and endopodal

| segments with pinnules along inner and outer margins

| respectively.

Third leg (Fig. 12A) with 2-segmented protopod; coxa with

| spinular row proximally near outer margin. Basis with naked

outer seta; ornamented with tiny spinules at base and hairs along

inner margin. Intercoxal sclerite unornamented. Exopod and

endopod 1-segmented, bearing pinnules along inner and outer

| margins respectively. Spine and seta formula as follows:

| Coxa Basis Exopod Endopod

iLeg1 0-1 1-1 1-0; III,5 0-1; 1,1+1,4

iLeg2 0-41 1-0 1-0; III,4 0-1; 1,1+1,3

jLeg3 0-0 1-0 IV,3 WEP Es)

| Copepodid IIT

Body length : mean + standard deviation = 511 + 22.8 um (range

|478 to 534 um, n=10), mean body width 179 + 5.3 um (range 173

}to 186 um, n=10). Differing from copepodid II as follows: body

(Fig. 5C) 7-segmented, comprising 4-segmented prosome

bearing swimming legs | to 4, and 3-segmented urosome (Fig.

6C).

Antennule 6-segmented (Fig. 7C): fourth segment partly

divided. Setal formula: 6, 2, 5, 4 + 1 aesthetasc, 3, 7 + |

aesthetasc. Antenna (Fig. 8C) with spinular row on inner margin

of second segment proximally. Second endopodal segment with

6 setae. Labrum with long spinular rows along lateral margins

distally.

Exopod of legs 1 (Fig. 10C) and 2 (Fig. 11C) with new seta on

inner margin of first segment. Intercoxal sclerite of leg 2

ornamented with rows of setules on anterior surface. Second

endopodal and exopodal segments each with new seta. Coxa of

third leg (Fig. 12B) with new seta at inner angle and bearing

spinular row on outer margin. Intercoxal sclerite ornamented.

Exopod and endopod 2-segmented. Leg 4 (Fig. 13A) with

2-segmented protopod; coxa with spinular row on outer margin.

Basis with outer angle seta and bearing spinular row on laterally.

Exopod and endopod l-segmented. Intercoxal sclerite

unornamented. Leg 5 (Fig. 6C) represented by 2 setae. Spine and

seta formula as follows:

Coxa Basis Exopod Endopod

Leg! 0-1 1-1 I-1;111,5 0-1;1,1+1,4

Leg2 0-1 1-0 I-1;IV,5 0-1;1,1+1,4

Leg3 0-1 1-0 I-0;111,4 0-1,1,1+1,3

Leg4 0-0 1-0 IV,3 ers

Female copepodid IV

Body length : mean + standard deviation = 573 + 19.5 um (range

547 to 608 um, n=10), mean body width 200 + 14 um (range 178

to 221 um, n=10). Differing from copepodid III as follows: body

(Fig. SE) 8-segmented with 4-segmented urosome (Fig. 6D);

spinular rows present mid-dorsally along posterior margins of

urosomites 2 and 3.

Antennule 6-segmented (Fig. 7D) with entire fourth segment.

Setal formula; 11, 4, 5, 4 + 1 aesthetasc, 3, 7 + 1 aesthetasc.

Antenna with 2 short spinular rows dorsally on inner side of first

segment. Second endopodal segment with 7 setae (Fig. 8D).

Mandible with spinular row on ventral margin of coxa.

Maxilliped with spinular rows midway along outer margin and

on anterior surface near inner setae.

Legs 1-4 with 2-segmented rami (Figs 10D, 11D, 12C, 13B).

Coxa of leg 4 with naked seta at inner angle proximally and

spinular row on outer margin. Spine and seta formula as follows:

Coxa Basis Exopod Endopod

Leg! 0-1 1-1 I-1;III,5 0-1;1,1+1,4

Leg2 0-1 1-0 I-1;IV,5 0-1;1,1+1,5

Leg3 0-1 1-0 I-1;IV,5 0-1;1,1+1,4

Leg4 0-1 1-0 I-0;1V,5 0-1;1,11,3

Leg 5 with 3 setae; leg 6 represented by 2 simple setae at

posterolateral angle of second urosomite (Fig. 6D)

Male copepodid IV

Body length, (range 565 to 643 um, n=2), body width, (range 195

to 204 um, n=2). Differing from female Copepodid IV as

follows: anterior part of body (Fig. 5D) not much broader than

posterior compared with female; constriction between somites

bearing 4th and Sth legs (Fig. 14A) not as distinct.

Appendages as in female fourth copepodid except as follows:

antennule 6-segmented (Fig. 15SA) but segment 3 broader.

Segments 3 and 4 with distinctive short spiniform setation

56 S. KARAYTUG AND G.A. BOXSHALL

9 PPPPEPEPPP PPPP

KS Y

S23

ees

Fig.13 P fimbriatus. Leg 4, anterior. A, Copepodid III; B, Copepodid IV; C, Copepodid V; D, Copepodid VI (adult). Scale bar in um.

elements (arrowed in Fig. 16D). Setal formula: 11, 4, 5 + spine, 4 (range 217 to 260 um, n=10). Differing from female Copepodid

+ spine + | aesthetasc, 3, 7 + 1 aesthetasc. IV as follows: body 9-segmented (Fig. 5F) with 5-segmented

urosome (Fig. 6E); anal somite with dense spinular row along

Female copepodid V posterior margin extending from ventral surface to dorsal

margin, either side of anal slit. Caudal rami about 3 times as

Body length : mean + standard deviation = 726 + 38.47 um long as broad.

(range 647 to 769 um, n=10), mean body width 246 + 12.4 um Antennule 7-segmented (Fig. 7E) : setal formula; 18, 6, 5,2 +1

LIFE CYCLE OF PARACYCLOPS

j |

WV VA AAA

ape re

MV VY

cee

L¢e\—

ee SSS ™ RSs

———

See

Fig.14 P fimbriatus. Male. A, Copepodid IV urosome, dorsal; B, Copepodid V, dorsal; C, adult urosome, ventral; D, adult urosome, dorsal; E,

Adult leg 5, ventral; F, Adult leg 6, ventral. Scale bars in ym.

S. KARAYTUG AND G.A. BOXSHALL

/ j roa)

ve / i ;

\ i \

\\ ‘

\ \

\\ \

Fig. 15 P. fimbriatus. Male antennules, lateral. A, Copepodid IV; B, Copepodid V. Scale bar in um.

LIFE CYCLE OF PARACYCLOPS

Fig. 16 Scanning electron micrographs of P. fimbriatus. A, Nauplius VI antenna, ventral; arrow indicates minute second exopodal segment; B,

Adult female antennule, aesthetasc on segment 5; C, Male copepodid V antennule, dorsal; D, Male copepodid IV antennule, ventral; arrow indi-

cates the spine on segment 4; E, Adult female antennule, double fusion on terminal segment, lateral. Scale bars A = 20 um, B = Sum, C= 40 um, D

= 25 um, E=5 pum.

aesthetasc, 2, 3, 7 + 1 aesthetasc. Second endopodal segment of

antenna (Fig. 8E) with 8 setae; distal seta VIII stronger than

others. Praecoxal arthrite of maxillule with 5 setae. Proximal

spinulose seta ornamented with long spinules. Praecoxa of

maxilla with short spinular row on outer margin. Basis of

maxilliped with 2 spinular rows laterally.

Leg | (Fig. 10E) with 3-segmented exopod and endopod,

bearing spinular row posteriorly on second segment of exopod.

Intercoxal sclerite with anterior spinular rows distally. Leg 2

(Fig. 11E) with praecoxa bearing spinular row on outer margin.

Exopod and endopod 3-segmented: first segment of exopod

with proximal spinular row anteriorly, second exopodal and

60 S. KARAYTUG AND G.A. BOXSHALL

Fig.17 P. fimbriatus. A, Adult female, dorsal; B, Adult female antennule, dorsal; C, Adult male, dorsal. Scale bars in pm.

endopodal segments with spinular rows posteriorly. Leg 3 (Fig. spinular row near outer proximal angle, spinular row on

12D) with 3-segmented exopod and endopod bearing posterior midanterior surface and very fine spinules along posterior

spinular rows on first and second segments of exopod and margin. Exopod and endopod 3-segmented: first and second

second segment of endopod. Intercoxal sclerite with transverse segments of exopod with anterior spinular rows distally.

spinular row on anterior surface. Leg 4 (Fig. 13C) with praecoxa Intercoxal sclerite with transverse spinular row on anterior

bearing spinular row on outer margin. Coxa with posterior surface. Spine and seta formula as follows:

LIFE CYCLE OF PARACYCLOPS 61

Fig. 18 P fimbriatus. Adult female. A, urosome, ventral; B, urosome, dorsal. C, leg 5, ventral. Scale bars in pm.

Coxa Basis Exopod Endopod

Leg! 0-1 1-I I-1;1-1;11,1,4 0-1;0-1;1,1,4

Leg2 0O- 1-0 I-1;I-1;1IL1,5 0-1;0-2;1,1,4

Leg3 0-1 1-0 I-1;1-1;111,1,5 0-1;0—2;1,1,4

Leg4 0-1 1-0 J-1;I-1;1,1,4 0-1;0—2;1, 11,2

Leg 6 represented by 1 plumose and 1 naked seta at

posterolateral angle of second urosomite (Fig. 6E)

Male Copepodid V

Mean body length: mean + standard deviation = 695 + 44.83 um

(range 634 to 743 um, n=7), mean body width 216 + 8.1 um

(range 200 to 221.73 um). Differing from male Copepodid IV as

follows: body 9-segmented: first to fourth urosomites

ornamented along posterior margins (Fig. 14B). Anal somite

densely furnished with spinules along distal margin and

extending either side of anal operculum.

Appendages as in fourth copepodid except as follows:

antennule (Fig. 15B) 7-segmented; proximal 3 segments much

enlarged, swollen (Figs 16C) first segment with one modified

spiniform seta (stippled on Fig. 15B); segment 3 with 6 similarly

modified elements and segment 4 with 1 such element. Setal

formula; 19 + 1 spine, 5, 5 + 6 spines, 2 + 1 aesthetasc + | spine, 2,

3, 7 + | aesthetasc.

Leg 6 (Fig. 14B) represented by 3 setae, not visible in dorsal

view.

Adult female

Body length: mean + standard deviation = 935 + 61.07 um (range

833 to 1013 um, n=10), mean body width 307 + 14.9 um (range

291 to 330 um, n=10). Body (Fig. 17A) comprising 4-segmented

prosome and 5-segmented urosome. Prosome _ with

cephalothorax and 3 free pedigerous somites decreasing in width

from anterior to posterior. Cephalothorax narrowing anteriorly;

widest aproximately in middle. Third and fourth pedigerous

somites with lateral groups of bristles at posterolateral angles.

Urosome (Figs 18A,B) consisting of 5th pedigerous somite,

genital double-somite and 3 free abdominal somites. Genital

double-somite about as long as broad. Genital double-somite

and first 2 free abdominal somites with surface ornamentation

dorsally and ventrally fifth pedigerous somite with

ornamentation along posterior margin. Anal somite with

spinular row ventrally extending round to anal operculum (Fig.

19A). Anal operculum smooth; row of spinules present in anal

cleft either side of midline (Fig. 19D).

Caudal rami about 3.5 times longer than broad with

ornamentation comprising rows of pits or cuticular depressions

on ventral surface (Fig. 19B). Generally held wide apart, slightly

divergent. Caudal rami with 6 setae; seta I missing; outer lateral

seta (III) with spinular row at base extending dorsally and

ventrally; small seta (II) on dorsolateral surface with spinular

row behind it and extending ventrally.

Antennule 8-segmented (Fig. 17B), quite short. First segment

with spinular row ventrally. Segment 2 with partial suture line.

Fourth segment longest. Segment 5 distinctive with short

aesthetasc (Fig. 16B); apical segment with aesthetasc fused to

adjacent seta at base (Fig. 16E). Setal formula: 8, 12, 6,5,2+ 1

aesthetasc, 2, 3, 7 + 1 aesthetasc. One element on seventh

segment possibly a setiform aesthetasc (see discussion below).

Antenna 4-segmented (Fig. 8F), comprising coxobasis and

3-segmented endopod. Coxobasis with complex ornamentation

S. KARAYTUG AND G.A. BOXSHALL

as figured and armed with 2 inner setae, and 1 outer spinulose

seta representing exopod. First endopodal segment with inner

distal seta and midsurface spinular row. Second endopodal

segment with 9 setae, of which 5 on inner margin and 4 arranged

along inner part of distal margin; segment ornamented with

spinules along outer margin. Third endopodal segment armed

with 7 setae around apex; segment ornamented with spinular

row along outer margin.

Labrum with complex ornamentation on ventral surface (Fig.

20A). Anterior part broader than posterior part. Posterior

margin forming strong teeth.

Mandible (Fig. 20B) consisting of well developed coxal

gnathobase and reduced palp. Gnathobasic blades mostly

simple, dorsal seta with spinules along inner rim. Palp

represented by 3 setae, one long and spinulose, the other

plumose. Third seta short and naked. Central surface of coxa

with 3 spinular rows and another spinular row on margin at base

of gnathobase.

Maxillule (Fig. 20C) consisting of powerful praecoxa and

reduced 2-segmented palp. Praecoxal arthrite armed with 7 setae

articulating at base and 4 spines fused to segment; proximalmost

seta spinulose, spines naked. Proximal segment of palp derived

from coxa and basis, bearing 2 naked and | spinulose inner

margin setae, plus outer seta representing exopod. Distal

segment of palp, representing endopod, armed with 2 setae with

spinules along margin and | naked seta.

Maxilla 5-segmented (Fig. 20D), comprising praecoxa, coxa,

basis and 2-segmented endopod. Praecoxa with spinular row on

outer margin. Praecoxal endite with 2 setae, 1 of which

spinulose. Coxa with proximal endite represented by single seta,

distal endite with well developed process carrying strong

spinulose seta and naked seta apically. Basis drawn out into

powerful spinulate claw and armed with strong accessory claw

with spinular row along convex margin and naked seta. First

endopodal segment carrying 2 spinulose setae, second carrying 3

setae.

Maxilliped 4-segmented (Fig. 20E), comprising syncoxa,

basis and 2-segmented endopod. Syncoxa armed with 3 inner

margin setae representing endites; ornamented with spinular

row near middle of inner margin and 2 spinules on outer margin.

Basis armed with 2 inner setae, | of which with spinules;

ornamented with 2 transverse rows of spinules near outer distal

angle and another spinular row midway along outer margin.

First endopodal segment bearing claw-like seta with 5 spinules at

midlength. Second endopodal segment with 3 setae, 2 of which

bearing spinules midway; outermost naked.

Legs | to 4 each with complex ornamentation on anterior

and posterior surfaces of coxa as figured; also ornamented on

intercoxal sclerite. Legs 1 to 3 with spinular rows on posterior

surface of exopodal segments | and 2, and endopodal segment

Leg 1 (Fig. 10F) with 3-segmented protopod. Praecoxa

represented by triangular sclerite at outer proximal angle. Coxa

with inner plumose seta. Basis with outer angle seta and setiform

spine on inner margin; bearing spinular row on posterior

surface. Both seta and spine with spinular rows at bases.

Legs 2 (Fig. 11F) and 3 (Fig. 12E) with 3-segmented

protopods. Coxa with inner setiform spine. Outer seta on basis

with spinular row at base. First segment of exopod with spinular

row on anterior surface.

Leg 4 (Fig. 13D) with 3-segmented protopod. Praecoxa

represented by thin, hooped sclerite. Coxa with inner setiform

spine. Basis with outer seta bearing spinular row at base;

spinular row on inner margin anteriorly. Segments | and 2 of

LIFE CYCLE OF PARACYCLOPS

Fig. 19 Scanning electron micrographs of P. fimbriatus. A, Adult female, anal operculum, lateral; B, adult female, caudal rami showing ornamenta-

tion of pits in integument; C, Nauplius II, ventral; D, Adult female, anal operculum, dorsal. Scale bars A = 20 pm, B = 40 um, C = 60 um, D = 25

um.

exopod with spinular row on distal margin anteriorly. Spine and

seta formula as follows:

Coxa Basis

0-1

0-I

0-1

0-I

Leg 5 (Fig. 18C) comprising single free segment, armed with 1

well developed outer spinulose seta, 1 strong inner spine

ornamented with spinules, and | plumose seta in middle. Leg 6

(Fig. 18A) represented by | naked seta and | tiny spinule

dorsolaterally.

Adult male

Body length: Mean + standard deviation = 732 + 47.45 pm

(range 673 to 769 um, n=10), mean body width 237 + 7.6 um

Sa aN

“WY

S. KARAYTUG AND G.A. BOXSHALL

Fig.20 P fimbriatus. Adult female. A, Labrum, ventral; B, Mandible, posteroventral; C, Maxillule, ventral; D, Maxilla, posterior; E, Maxilliped,

anterior. Scale bar in pm.

>, a Val A

y XXI-XXIII

vor

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

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

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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

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.

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The genera of pelmatochromine fishes (Teleostei, Cichlidae).

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Certain Actiniaria (Cnidaria, Anthozoa) from the Red Sea

and tropical Indo-Pacific Ocean. K. W. England. 1987. Pp.

205-292. £23.95

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Volume 54

No. | The cranial muscles and ligaments of macrouroid fishes

(Teleostei: Gadiformes) functional, ecological and

phylogenetic inferences. G. J. Howes. 1988. Pp. 1-62. £16.90

No. 2 A review of the Macrochelidae (Acari: Mesostigmata) of the

British Isles. K. H. Hyatt & R. M. Emberson. 1988. Pp

63-126. £17.20

A revision of Haplocaulus Precht, 1935 (Ciliophora:

Peritrichida) and its morphological relatives. A. Warren.

1988. Pp. 127-152. £8.50

Echinoderms of the Rockall Trough and adjacent areas. 3.

Additional records. R. Harvey, J. D. Gage, D. S. M. Billett,

A. M. Clark, G. L. J. Paterson. 1988. Pp. 153-198. £14.00

A morphological atlas of the avian uropygial gland. D. W.

Johnston. 1988. £16.60

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Miscellanea.

A review of the Copepod endoparasites of brittle stars

(Ophiuroida). G. A. Boxshall.

A new genus of tantulocaridan (Crustacea: Tantulocarida)

parasitic on a harpacticoid copepod from Tasmania. G. A.

Boxshall.

Unusual ascothoracid nauplii from the Red Sea. G. A.

Boxshall & R. Bottger-Schnack.

New nicothoid copepods (Copepoda: Siphonostomatoida)

from an amphipod and from deep sea isopods. G. A.

Boxshall & K. Harrison.

A new genus of Lichomolgidae (Copepoda:

Poecilostomatoida) associated with a phoronid in Hong

Kong. G. A. Boxshall & A. G. Humes. 1988. £9.00

| Volume 55

No. 1 Miscellanea.

Structure and taxonomy of the genus Delosina Wiesner,

1931 (Protozoa: Foraminiferida). S. A. Revets.

Morphology and morphogenesis of Parakahliella haideri

nov. spec. (Ciliophora, Hypotrichida). H. Berger & W.

Foissner.

Morphology and biometry of some soil hypotrichs

(Protozoa, Ciliophora) from Europe and Japan. H. Berger

& W. Foissner.

Polyclad turbellarians recorded from African waters. S.

Prudhoe, O.B.E.

Ten new taxa of chiropteran myobiids of the genus

Pteracarus (Acarina: Myobiidae). K. Uchikawa.

Anatomy and phylogeny of the cyprinid fish genus

Onychostoma Ginther, 1896. C. Yiyu. 1989. Pp.

1-121. £30.00

No. 2 Studies on the Deep Sea Protobranchia: The Subfamily

Ledellinae (Nuculanidae). J. A. Allen & F. J. Hannah.

1989. £38.00

Volume 56

No. 1 Osteology of the Soay sheep. J. Clutton-Brock, K.

Dennis-Bryan, P. L. Armitage & P. A. Jewell.

A new marine species of Euplotes (Ciliophora,

Hypotrichida) from Antarctica. A. Valbonesi & P. Luporini.

Revision of the genus Eizalia Gerlach, 1957 (Nematoda:

Xyalidae) including three new species from an oil producing

zone in the Gulf of Mexico, with a discussion of the sibling

species problem. D. Castillo-Fernandez & P. J. D.

Lambshead.

Records of Neba!ia (Crustacea: Lepostraca) from the

Southern Hemisphere a critical review. Erik Dahl.

1990. £31.00

No. 2 Tinogullmia riemanni sp. nov. (Allogromiina:

Foraminiferida), a new species associated with organic

detritus in the deep-sea. A. J. Gooday.

Larval and post-larval development of Anapagurus

chiroacanthus (Lilljeborg, 1855) Anomura: Paguroidea:

Paguridae. R. W. Ingle.

Redescription of Martialia hyadesi Rochebrune and

Mabille, 1889 (Mollusca: Cephalopoda) from the Southern

Ocean. P. G. Rodhouse & J. Yeatman.

The phylogenetic relationships of salmonoid fishes. C. P. J.

Sanford.

A review of the Bathygadidae (Teleostei: Gadiformes). G. J.

Howes & O. A. Crimmen. 1990. £36.00

Volume 57

No. | Morphology and biometry of twelve soil testate amoebae

(Protozoa, Rhizopoda) from Australia, Africa and Austria.

G. Liiftenegger & W. Foissner

A revision of Cothurnia (Ciliophora: Peritrichida) and its

morphological relatives. A. Warren & J. Paynter

Indian Ocean echinoderms collected during the Sinbad

Voyage (1980-81): 2. Asteroidea. L. M. Marsh & A. R.G.

Price

The identity and taxonomic status of Tilapia arnoldi

Bilchrist and Thompson, 1917 (Teleostei, Cichlidae). P. H.

Greenwood

Anatomy, phylogeny and txonomy of the gadoid fish genus

Macruronus Ginther, 1873, with a revised hypothesis of

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No. 2 The pharyngobranchial organ of mugilid fishes; its structure,

variability, ontogeny, possible function and taxonomic

utility. I. J. Harrison & G. J. Howes

Cranial anatomy and phylogeny of the South-East Asian

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A collection of seasnakes from Thailand with new records

of Hydrophis belcheri (Gray). C. J. McCarthy & D. A.

Warrell

The copepod inhabitants of sponges and algae from Hong

Kong. S. Malt

The freshwater cyclopoid of Nigeria, with an illustrated key

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A new species of Ferdina (Echinodermata: Asteroidea) from

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No. 1 The morphology and phylogeny of the Cerastinae

(Pulmonata: Pupilloidea). P. B. Mordan

A redescription of the uniquely polychromatic African

cichlid fish Tilapia guinasana Trewavas, 1936. P. H.

Greenwood

A revision and redescription of the monotypic cichlid genus

Pharyngochromis (Teleostei, Labroidei). P. H. Greenwood

Description of a new species of Microgale (Insectivora:

Tenrecidae) from eastern Madagascar. P. D. Jenkins

Studies on the deep-sea Protobranchia (Bivalvia): the family

Nuculidae. P. M. Rhind and J. A. Allen £40.30

No. 2 Notes on the anatomy and classification of ophidiiform

fishes with particular reference to the abyssal genus

Acanthonus Gunther, 1878. G. J. Howes

Morphology and morphogenesis of the soil ciliate Bakuella

edaphoni nov. spec. and revision of the genus Bakuella

Agamaliev & Alekperov, 1976 (Ciliophora, Hypotrichida).

W. Song, N. Wilbert and H. Berger

A new genus and species of freshwater crab from

Cameroon, West Africa (Crustacea, Brachyura,

Potamoidea, Potamonautidae). N. Cumberlidge and P. F.

Clark

On the discovery of the male of Mormonilla Giesbrecht,

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No. 1 A new snake from St Lucia, West Indies. G. Underwood

Anatomy of the Melanonidae (Teleostei: Gadiformes), with

comments on its phylogenetic relationships. G. J. Howes

A review of the serranochromine cichlid fish genera

Pharyngochromis, Sargochromis, Serranochromis and Chetia

(Teleoste1: Labroidei). P. H. Greenwood

A revision of Danielssenia Boeck and Psammis Sars with the

establishment of two new genera Archisenia and

Bathypsammis (Harpacticoida: Paranannopidae). R. Huys

and J. M. Gee.

A new species of Syrticola Willems & Claeys, 1982

(Copepoda: Harpacticoida) from Japan with notes on the

type species. R. Huys and S. Ohtsuka

Erratum £40.30

No. 2 The status of the Persian Gulf sea snake Hydrophis

lapemoides (Gray, 1849) (Serpentes, Hydrophiidae). A.

Redsted Rasmussen.

Taxonomic revision of some Recent agglutinated

foraminifera from the Malay Archipelago, in the Millett

Collection, The Natural History Museum, London. P.

Bronnimann and J. E. Whittaker.

Foregut anatomy, feeding mechanisms, relationships and

classification of the Conoidea (=Toxoglossa)(Gastropoda).

J. D. Taylor, Y. I. Kantor and A.V. Sysoev. 1993. Pp.

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No. | A new subfamily and genus in Achatinidae (Pulmonata:

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On Recent species of Spiraserpula Regenhardt, 1961, a

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No. 2 Phylogenetic relationships between arietellid genera

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1994. Pp. 105-277. £40.30

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No. | A revised familial classification for certain cirrhitoid genera

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Studies on the deep-sea Protobranchia (Bivalvia); the

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No. 2 Primary studies on a mandibulohyoid ‘ligament’ and other

intrabucal connective tissue linkages in cirrhitid, latrid and

cheilodactylid fishes (Perciformes: Cirrhitoidei). P.H.

Greenwood

A new species of Crocidura (Insectivora: Soricidae)

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A.L. Smith

Redescription of Sudanonautes Floweri (De Man, 1901)

(Brachyura: Potamoidea: Potamonautidae) from Nigeria

and Central Africa. N. Cumberlidge

Association of epaxial musculature with dorsal-fin

pterygiophores in acanthomorph fishes, and its phylogenetic

significance. R.D. Mooi and A.C. Gill. 1995. Pp.

91-138.

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