March 31, 1964

INDO-PACIFIC MOLLUSCA, vol. 1, no. 5

Turrinae 227

OS

:T$K

*1 ^ SMto *ii*n

THE FAMILY TURRIDAE IN THE INDO-PACIFIC

Part 1. The subfamily Turrinae

by A. W. B. Powell

Auckland Institute and Museum Auckland, New Zealand

The turrid gastropods comprise one of a group of five related families which form the superfamily Toxoglossa within the order Neogastropoda— 1 ) Speightiidae, 2) Turridae, 3) Thatcheriidae, 4) Conidae, and 5) Terebridae. This monograph is the first of a series intended to cover the family Turridae. The turrids are the largest family group among the marine gastropods, and they occur from the intertidal zone to the abyssal depths and in all seas, including the Arctic and the Antarctic.

The term Toxoglossa refers to the extraordinary toxic property associated with a special type of radula which is found in many turrids, and reaches its climax of development in the Conidae. The rad- ula of most toxoglossate gastropods is reduced to a single pair of slender marginals in each transverse row. In Conus, the teeth are barbed and lie loose in the radula sac from which they are ejected as individual harpoons.

Morphological Characters of the Turridae

The Shell of the Turridae — There is no charac- teristic turrid shape by which all members can be readily recognized, for many turrids simulate in shape such groups as the Buccinidae, Muricidae, Fasciolariidae, Columbellidae, Mitridae, Conidae, and even Terebridae. However, the one shell fea- ture common to the members of the Turridae is a slit or shallow to deep sinus on the outer lip vari- ously located between the suture and the periphery of the last whorl. In size, the adult turrid shell may vary from about 1 to 160 mm.

In the subfamily Turrinae, a well-developed sinus is situated either at the periphery or on a minor ridge immediately above it. Most Turrinae are fusi- form with a tall spire and a long tapered anterior canal, although in some genera, such as Xenuro- turris, the base of the shell is truncated. Typical Turrinae closely resemble Fusinus (family Fascio-

lariidae), the only apparent shell difference being in the presence of a turrid sinus on the outer lip. Characteristic labial profiles of the Turrinae are shown in plate 173, and of other subfamilies in plate 174.

The Protoconch of the Turridae — In most gas- tropod families the protoconch is fairly constant and characteristic, and is generally considered to be of importance in determining phylogenetic re- lationships. A radical difference in protoconch type usually suggests strong taxonomic dissimilarity. Some degree of difference in the size and number of the nuclear whorls exists within a species, de- pending upon the salinity variations in the environ- ment; and there may be produced, again within the same species, either a short pelagic or a non-pelagic stage (in Brachystomia, Thorson, 1946, p. 206, and Rasmussen, 1951, pp. 210-221). Thorson (1950, p. 33) has shown that northern Arctic Naticidae have a nonpelagic development and hence a protoconch of large whorls, while Naticidae of temperate, warm seas have a short pelagic stage and a proto- conch of small, more tightly-coiled whorls. How- ever, Thorson placed all these species in the genus Natica, although, in fact, they belong to quite dif- ferent genera having quite different opercula.

In turrids, there are species and genera which have almost identical adult shell characters, but which have quite dissimilar protoconchs. The sig- nificance of this phenomenon is not, as yet, com- pletely understood, and in the present absence of

1 2 3 4 5

Plate 172. Five families of the supcrfamily Toxoglossa or Conacea. Fig. 1, Speightiidae (Speiglitia) . Fig. 2, Turridae (Turns). Fig. 3, Thatcheriidae (Thatcheria) . Fig. 4, Coni- dae (Conus). Fig. 5, Terebridae ( Terebra ).

228 Turrinae

A. W. B. Powell

Turridae

TITRRIS LOPHIOTOMA POL VST IRA GEMMULA

Plate 173. Labial profiles of genera in the subfamily Tur- rinae showing the characteristic shape and location of the sinus or labial slit.

life history and embryological facts, it is advocated that subgenera, or even genera, be recognized on the basis of the protoconch characters, even though the adult shells are almost identical. Examples of these “pairs” of higher taxa are the Lophiotoma and Lophioturris complex of species, and the Tomo- pleura and Maoritomellci group.

In all likelihood, the size of the protoconch and its number of whorls are correlated with the length of the free-swimming stage of the veliger. A proto- conch of few, rapidly-expanding whorls suggests a short free-swimming life, while one of several, slowly-expanding whorls or one with a sinusigerous- like outer lip suggests a long-living, and hence far- travelling, larval life.

The Dentition of the Turridae — The radula in the Turridae is of several very distinctive types ranging from the prototypic form, with central, lateral and marginal teeth, through a series char- acterized by the absence of laterals but with a com- pensating enlargement of the central tooth, to fi- nally the toxoglossate state, in which only the marginals, one on each side, remain.

The prototypic radula is found in the subfamily Clavinae in genera such as Clavus, Drillia and S pirotropis. The second group, minus laterals, oc- curs in the Cochlespirinae (Aforia and Leucosyrinx) as well as in the Turrinae, and in Ptychosyrinx and

EOTURRIS XENUROT 'JRRIS EPIDIRONA

Turridrupa. The third group, in which all but the marginals have disappeared, culminates in a radula form very close to that of the Conidae. The Man- geliinae, Daphnellinae and also some of the Clavinae have this toxoglossate style of dentition.

The radula form in the Turrinae and in some members of the Turriculinae is a modified toxo- glossate type, having only marginals present, but these are “wishbone” in shape. They in turn show some relationship with the radula of the Clava- tulinae in which the marginals are massive, often with one of the two extremities of the marginals existing as a separate plate. In addition there is an incipient or vestigial central tooth.

It was found that the prototypic style of radula was of common occurrence in many of the Clavinae, but on the other hand, quite a number of genera, Phenatoma for instance, with all the external shell characteristics of that subfamily, were found to possess only slender toxoglossate-like marginals.

Recently, Robinson (1960, Proc. Zool. Soc., 135, p. 319) has demonstrated that the English species of Mangelia are comparable with the Conidae in that the radula is truly toxoglossate, complete with the neurotoxic apparatus as well as a long proboscis capable of sufficient extension to harpoon prey.

These animals have a greatly coiled poison gland which opens ventrally into the oesophagus imme- diately posterior to the opening of the buccal sac. The poison gland is a coiled tube, and the swollen end of this gland is a propulsive organ, termed the

TURRICULINAE CLAVATULINAE COCHLESPIRINAE CLAVINAE

DAPHNELLINAE MANGELIINAE

Plate 174. Labial profiles characteristic of other subfamilies ( Cymatosyrinx ); Daphnellinae ( Daphnella ); Mangeliinae

in the Turridae. Turriculinae ( Turricula ); Clavatulinae ( Elrema ).

( Clavatula ) ; Cochlespirinae ( Ancistrosyrinx) ; Clavinae

[22 - 662]

March 31, 1964

INDO-PACIFIC MOLLUSCA, vol. 1, no. 5

Turrinae 229

Figs.

Plate 175

1, 6 Gemmula (V nedogemmula) unedo (Kiener).

Japan (see text p. 22-761 ).

2, 3 Lophiotoma ( Lophioturris ) indica (Roding).

Cuyo Id., Philippines ( p. 22-931).

4, 5 Lophiotoma ( Lophioturris) leucotropis ( Adams and Reeve) (p. 22-932).

7, 8 Gemmula (U nedogemmula) deshayesii (Dou- met). Hongkong (7), Japan (8) (p. 22-762).

9 Lophiotoma (Lophioturris) indica (Roding). Philippines (p. 22-931).

10,11 Lophiotoma (Lophioturris) polytropa ( Hel- bling). Luzon Id., Philippines (p. 22-933).

12 Xenuroturris cingulifera (Lamarck). Zanzibar (p. 22-962).

13 Xenuroturris millepunctata (Sowerby). New Caledonia (p. 22-963).

14,15 Xenuroturris cerithiformis (Tinker). Hawaiian Islands (p. 22-964).

16 Lophiotoma (Lophioturris) indica (Roding). Moluccas.

17,18 Xenuroturris millepunctata (Sowerby). New Caledonia (p. 22-963).

19,20 Xenuroturris cingulifera (Lamarck). Japan (p. 22-962).

21,22 Xenuroturris castanella (Tinker). Hawaiian Is- lands (p. 22-964).

(all 2/3 natural size)

[22 - 663]

230 Turrinae

A. W. B. Powell

Turridae

muscular bulb, which functions in the harpooning of prey by means of poison-charged, detached radu- lar teeth.

The fully developed toxoglossate dentition does not seem to be correlated with definite shell charac- teristics; in fact, it appears to cut across any sub- family arrangement based upon shell features alone.

The subfamily groups in use in the Turrinae seem to form more or less natural groups that appear to have some geographical and chronological signifi- cance, yet the toxoglossate dentition is likely to develop in any one of them. It is probable, there- fore, that a change from the prototypic to the toxo- glossate state can take place independently in any one of the “subfamily” groups, and if so, no doubt as a direct response to a change to predaceous feeding.

If this is not the case, the inference is that the system of subfamilies at present in use is not morphologically sound, and merely indicates con- venient groups of species based largely upon exter- nal resemblances. The following illustrations show some of the characteristic radula types in the Tur- ridae:

The Operculum of the Turrids — The operculum is here considered to be of secondary importance as a taxonomic character, for after all, its shape is consequent to the shape of the aperture; opercular growth naturally takes the most convenient form to fill the apertural space. Opercular growth is achieved by means of a logarithmic spiral such as in Turbo or, as in the turrids, by the addition of concentric rings or the addition of excentric rings. For species with a long narrow aperture, the oper- culum is lanceolate or leaf-shaped. Excentric growth with an apical nucleus is the obvious mechanical method of growth in such types. The clavatulids, on the other hand, have a very differ- ent apertural shape, due to the lower median plac- ing of the parietal angulation of the inner lip, and the adpressed suture which is clasped high over the preceding whorl. Thus by fan-wise excentric growth from a medio-lateral position the apertural space is best filled. A variation of the leaf-shaped operculum takes place where the aperture does not contract rapidly toward the anterior canal. The re- sultant squarish aperture is accommodated by an operculum in which the position of the nucleus may move over toward the outer lip in order to facilitate the mechanism of opercular growth.

Again in some abyssal genera such as Steiraxis, the operculum may become degenerate and remain

Plate 176 Example of paucispiral and polygyrate proto- eonchs in the Turridae. Fig. a, Micantapex angustatus Powell; fig. b, Bathytoma bartrumi Laws.

Plate 177. Radulae types in the Turridae. Fig. a, prototype with the central, lateral and marginal teeth present (for- mula: 1 + 1 + 1 + 1 + 1), Drillia umbilicata Gray from West Africa. Fig. b, Ptychosyrinx bisinuatus ( Martens ) from East Africa in which the central tooth is enlarged to compensate for the loss of the laterals ( formula: 1 + 0 + 1+ 0+1). Fig. c, Clionella sinuata (Born) from South Africa with a small central and without laterals ( formula : 1+0+1+0+1).

Fig. d, Turns babylonia (Linnaeus) from the southwest Pacific with only “wish-bone”-shaped marginals (formula: 1+0 + 0 + 0+1). Fig. e, Hormospira maculosa (Sowerby) from West Mexico whose formula may be either 0+1 + 1 + 1+0 ( marginals absent? ) or 1+0h 1 -0+1 ( laterals absent). Fig. f, Phenatoma novaezelandiae (Reeve) from New Zealand with only slender, barbed marginals (formula: 1+0 + 0 + 0+1). Fig. g, Microdrillia optima Thiele from off East Africa with only awl-shaped marginals (formula: 1+0 + 0 + 0+1). Fig. h, Propebela turricula (Montagu) from northern Europe (formula: 1+0 + 0 + 0+1). Fig. i, Daphnella cancellata Hutton from New Zealand with mar- ginals only ( formula: 1 + 0 + 0 + 0 + 1) (all greatly enlarged, but not to scale).

[22 - 664]

March 31, 1964

INDO-PACIFIC MOLLUSCA, vol. 1, no. 5

Turrinae 231

permanently ovate, of small size and without sub- sequent growth accretions.

The operculum is found to be absent in the Mangeliinae, or at least in the few species that have been examined. On the other hand, the complex of Boreal species, such as Oenopota, “ Bela auct.” and “Lora,” although closely resembling mangelids, possess an operculum.

The Daphnellinae are also supposed to lack an operculum, but one species, Typhlodaphne puris- sima (Strebel, 1908), from deep-water off South Georgia, has a vestigial operculum which is ovate, with an apical nucleus, and is very small in relation to the size of the aperture (Powell, 1951, Discovery Rep. 26, p. 196, fig. N.130).

Obviously, the operculum in the light of these considerations cannot be given a high taxonomic value.

Nomenclature of the Turrids — The Linnaean school did not distinguish turrids as a separate group but assigned them individually, according to appearance, to the genera Mur ex, Buccinum or S trombus. The earliest use of a group name for these shells seems to be the subfamily Pleurotomi- nae of Swainson, 1840, in his “Treatise on Mala- cology” as one of four subfamilies of the Strom- bidae. However, H. and A. Adams, 1853, in “The Genera of Recent Mollusca” were the first to ele- vate the group to family rank, i.e., Turritidae; they admitted three subfamilies, the Turritinae, the Clavatulinae and the Defranciinae. Then for more than half a century the family name Pleurotomidae of Chenu, 1859, was in general use. With the rec- ognition of Roding’s Museum Boltenianum as nomenclaturally valid, Pleurotoma Lamarck, 1799, became a synonym of Turns Roding, 1798.

The acceptance of a family name based upon Turns Roding, 1798, to replace Pleurotomidae, based upon a synonym, is allowed under Article 40(a) of the 1961 edition of “The International Code of Zoological Nomenclature,” and further, Article 40(b) rules that a name adopted by virtue of the provisions of section (a) takes the date of the rejected name.

The revised family name, however, has been ren- dered in two ways, i.e., Turritidae and Turridae. The first use of the former seems to have been by H. and A. Adams, 1853, in “The Genera of Recent Mollusca,” but most modern authors since Hedley, 1922, have preferred the shorter rendering. The family name here adopted is TURRIDAE ( = Pleu- rotominae = Turritidae = Pleurotomidae ) Swainson, 1840.

Plate 178. Opercula of Turridae. Fig. 1, Hormospira mac- ulosa (Sowerby), west Mexico. Fig. 2, Crassispira pluto Pilsbry and Lowe, west Mexico. Fig. 3, Aoteadrillia otogoen- sis Powell, 50 fms. off Otago Heads, New Zealand. Fig. 4, Xenuroturris cingulifera (Lamarck), Queensland. Fig. 5, Tomopleura pouloensis ( Jousseaume), Aden. Fig. 6, Pusion- clla nifat ( Bruguiere ) , northwest Africa. Fig. 7, Aforia mag- nifica (Strebel), Palmer Archipelago, Antarctic.

Plate 179. Gross anatomy of a turrid ( Mangelia powisiana Dautzenberg ) . bm, buccal mass; eg, right cerebral ganglion; ct, ctenidium; dg, digestive gland; dpg, duct of poison gland; f, foot; gd, genital duct; k, kidney; me, mantle edge; os, osphradium; ot, oral tube; pb, proboscis; pg, muscular sac, the so-called poison gland; pig, right pleural ganglion; poe, posterior oesophagus; r, rectum; rm, retractor muscle of proboscis; rs, radular sac; s, siphon; sg, salivary gland; ssh, suspensory sheath; t, tentacle (from Fretter and Gra- ham, 1962, British Prosohranch Molluscs, Ray Society, Lon- don, p. 167, fig. 104).

[22 - 665]

232 T urrinae

A. W. B. Powell

Turridae

All Lophiotoma (Lophiotoma) acuta (Perry) and its forms, except figs. 11, 12, which are Lophiotoma alhina (Lamarck). Figs. 1, 2 Bega Id., Viti Levu Id., Fiji.

3 Rani, west end of New Guinea.

4 Heron Id., Queensland, Australia.

5 Mindoro Id., Philippines.

6 marmorata form, west end New Guinea.

7, 8 Mozambique City, Portuguese East Africa.

9 Zanzibar Id., East Africa.

Reef, Queensland, Australia.

LI, 12 Lophiotoma albina (Lamarck). Moluccas.

13 L. acuta form microsticta. Japan.

14 L. acuta form jickelii. Philippines.

L5, 16 Zamboanga, Philippines and Palau Islands.

17 form microsticta. Ryukyu Islands.

18 Ryukyu Islands.

19 form iickelii. Red Sea.

( all natural size )

Plate 180 (see text p. 22-913)

10 King’s

[22 - 666]

March 31, 1964

INDO-PACIFIC MOLLUSCA, vol. 1, no. 5

Turrinae 233

Plate 181 ( see text p. 22-977 )

Figs. 1, 2 3, 4

5

6

7, 8 9, 10 II, 12

13

Turns babylonia (Linnaeus). Southwest Pacific. Turns babylonia (Linnaeus). Philippines and west New Guinea.

Tunis babylonia (Linnaeus). Narrow form; Philippines.

Tunis babylonia (Linnaeus). Pale form; west New Guinea.

Tunis garnonsii (Reeve). Zanzibar.

Tunis crispa (Lamarck). Hongkong, 50 fms. Tunis crispa (Lamarck). Luzon Id., Philippines (11); Noumea, New Caledonia (12).

Tunis crispa yeddoensis ( Jousseaume). Japan.

14,15 Tunis cryptorrhaphe (Sowerby). Bohol Id., Philippines.

16, 17 Tunis spectabilis (Reeve), Philippines (16) and Moluccas ( 17 ) .

18 Tunis garnonsii (Reeve). Slender form; Lubang

Id., Philippines.

19 Tunis annulata (Reeve). Tosa, Japan.

20 Tunis undosa (Lamarck). Mindoro Id., Philip-

pines.

(all 2/3 natural size)

234 Turrinae

A. W. B. Powell

T urridae

For subdivisions of the Turridae the reader is re- ferred to the writer’s 1942 paper (see bibliogra- phy), in which the following subfamilies were adopted and diagnosed: Turrinae Swainson, 1840 (emended); Turriculinae Powell, 1942; Cochle- spirinae Powell, 1942; Clavatulinae H. & A. Adams, 1848; Conorbinae Powell, 1942; Clavinae Powell, 1942; Borsoniinae Bellardi, 1875; Mangeliinae Fis- cher, 1887 and Daphnellinae Hedley, 1922.

In the same paper two new families related to the Turridae were proposed: Speightiidae and Thatcheriidae. Charig (1963, Bull. Brit. Mus. (Nat. Hist.) Geol. 7(9), pp. 255-297) relegated the latter to subfamily rank within the Turridae but this question will be discussed at the appropriate place in a later number of “Indo-Pacific Mollusca.”

Plate 182. Development of a turrid ( Philbertia linearis Montagu from Denmark). Fig. a, egg capsule with young embryos. Fig. b, embryos nearly ready to hatch. Fig. c, hatched larva. Fig. d, older larva. Fig. e, older larva swim- ming showing the ciliated velar lobes ( from C. Thorson, 1946. Reproduction and Larval Development of Danish Marine Bottom Invertebrates, p. 237).

Doubtless when more is known concerning the soft parts of turrids there may be considerable modification of the above subdivisions, but mean- while they serve as a workable scheme and form the basis for future discussion.

Phylogeny of the Turridae — The earliest known undoubted turrid is apparently Clinura anassa Murphy & Rodda, 1960, from the Bald Hills for- mation, California, considered to be Cretaceous, late Albian to late Cenomanian. This species, how- ever, is not a Clinura, which is a near ally of That- cheria. Its affinity is more likely with the forerun- ners of the Turriculinae.

The previous earliest record for the family, Tur- riculina unica Gregorio, from the Liassic (Jurassic) of Sicily, is now considered non turrid, and has been placed by Wenz ( 1938, p. 391 ) , with a query, in the family Coelostylinidae of the Loxonematacea.

The family Turridae is undoubtedly an older one than the Conidae and is considered the main stem from which the several toxoglossate families have at different times arisen.

The prototypic radula found in a number of tur- rid genera suggests that the derivation of the fam- ily was from the taenioglossids rather than from the rachiglossids.

Turrids were already well represented in the Upper Cretaceous, with six of the nine now rec- ognised Recent subfamilies in evidence, and this indicates a much earlier inception for the family, but unfortunately there are no authentic records in support of such an assumption. Since the pres- ence of some form of sinus in the outer lip is the only reliable indicator of a fossil turrid, it is unlikely that the early stages of differentiation for the fam- ily will ever be recognised.

Regarding the origin of the subfamily Turrinae, it seems likely that there is much closer affinity among the Turrinae, Turriculinae, Cochlespirinae and Clavatulinae than with the remaining sub- families.

Detailed phylogenetic discussion will appear at appropriate places throughout the subsequent parts of this work.

Geographical Distribution of the Turrinae — The modern geographical distribution of the Turrinae differs markedly from patterns of the early Tertiary. For instance, Gemmula and a number of short-lived offshoots from it, characterized the Eocene hori- zons of Europe, England and the southern United States, but now Southeast Asia and vicinity has be- come the focal point for that genus and its allies. In fact, there is in that area an explosive develop-

[22 - 668]

March 31, 1964

INDO-PACIFIC MOLLUSCA, vol. 1, no. 5

Turrinae 235

ment with no less than thirteen living genera of the Turrinae alone represented. The Tertiary faunas of India, Burma, Java and Japan show that the Tur- rinae were already strongly entrenched in these areas from the Miocene onward.

However, very few members of the Turrinae have managed to penetrate farther eastward into the Pacific than the Western Pacific Arc, that geo- graphical feature running from the Ryukyu Islands through the Philippines, New Guinea and New Caledonia to Fiji. The exceptions are Lophiotoma acuta, which has reached the Marshall Islands and Samoa; Xenuroturris cingulifera and millepunctata , both of which are represented by derivative species in the Hawaiian group; Turris spectabilis, recorded from the Marshall and the Phoenix groups; Turris cryptorrhaphe, also from the Marshalls; and a new subspecies of crispa from the Hawaiian Islands.

The Turrinae appear to be absent from the So- ciety group and from most of the oceanic islands and atolls of the central Pacific. It is of interest, also, that the Turrinae are very rare at some of the atoll groups of the Indian Ocean, for Miss V. Orr, who recently collected intensively at Cocos-Keeling Islands, located only one member of the Turrinae, although small mangelids and daphnellids were there.

In tropical western America, the Caribbean and Florida, however, there is a development of large Turrinae, particularly in the genus Polystira, a shell that closely parallels the Indo-Pacific Lophiotoma, but has its own ancestry in that area by undoubted development from Pleuroliria of the Eocene and Oligocene of the southeastern United States. The genus Gemmula, wide-ranging both geographically and in time, also has representatives in tropical western America and off Florida.

In European-Mediterranean-West African wa- ters, however, Gemmula is not represented, al- though it reached Pliocene times in England. The related Fusiturris, however, which has an unbroken ancestry back to the Paleocene, reaches Recent times in the Mediterranean and West Africa, and is the only large genus of the Turrinae still living in those areas.

The larger Turrinae of tropical America, the Mediterranean and the Indo-Pacific have obviously developed independently after the breaking up of that ancient equatorial waterway, the Tethys Sea. The cosmopolitan Gemmula, however, has in the main, retained its generic identity despite geo- graphic segregating influences. Certainly a number of mutations have arisen from it at various times

during its long life from the Paleocene to the pres- ent, but nevertheless the tropical west American type of the genus, hindsiana Berry (formerly gem- mata Reeve ) and the deep-water Caribbean perisce- lida Dali, have all the essential characteristics of the Indo-Pacific members.

List of Recognized Taxa in the Turrinae

Below are listed the recognised generic and sub- generic taxa of the T urrinae, with species and sub- species. Taxa not represented in the Indo-Pacific are in square [brackets], and with these only the type species is cited. Fossils are prefixed by a dag- ger. Where a species occurs both Recent and fossil the dagger occurs after the name.

Included in the list are southern Australian Re- cent and Tertiary Turrinae, as well as New Zealand Tertiary members. Such species are not excluded since they have had an Indo-Pacific origin during past periods of the Tertiary when warm waters extended much further south than they do at present.

Subfamily Turrinae

[Fusiturris Thiele, 1929]

[undatiruga Bivona, 1832], Type Gemmula Weinkauff, 1875

[hindsiana Berry, 1958] (=g emmata Reeve, 1843). Type

aethiopica (Thiele, 1925) amabilis (Weinkauff, 1875) f bimarginata (Suter, 1917) f birmanica (Vredenburg, 1921) f clifdenensis (Powell, 1942) congener (E. A. Smith, 1894) subsp. cosmoi (Sykes, 1930) subsp. diomedea Powell, new subsp. f subsp. mekranica (Vredenburg, 1925) jmiocoronifera Powell, new name dampierana Powell, new species f disjuncta Laws, 1936 ducalis (Thiele, 1925) f duplex (Suter, 1917) f gellibrandensis Chappie, 1934 gemmidina (Martens, 1902) gilchristi (Sowerby, 1902) graeffei (Weinkauff, 1875) hawleyi (Iredale, 1931) hombroni (Hedley, 1922) t imitatrix ( Martin, 1916 ) f iris (Vredenburg, 1921) f kaiparaensis (Marshall, 1918) f karangensis (Martin, 1895)

[22 - 669]

236 Turrinae

A. W. B. Powell

Turridae

kieneri (Doumet, 1840) fsubsp. ryuktjuensis MacNeil, 1960 fsubsp. woodwardi (Martin, 1884) f kishimaensis Shuto and Ueda, 1963 f kotorai ( Nomura & Zinbo, 1935 ) f lawsi Powell, 1942 \longwoodensis Powell, 1942 \margaritata (Marshall, 1918) martini (Tesch, 1915 ) t monilifera (Pease, 1860) murrayi Powell, new species f orba Marwick, 1931 f ornata (Marshall, 1918) f pakistanica (Eames, 1952) f peraspera Marwick, 1931 f polita (Marshall, 1919) praesignis (E. A. Smith, 1895) t pulchella Shuto, 1961 f reticulata (Marshall, 1919) rotatilis (Martens, 1902) f samueli (Tenison-Woods, 1879) sihogae (Schepman, 1913) sibukoensis Powell, new species f sindiensis (Vredenburg, 1925) f soriensis (Eames, 1952) speciosa (Reeve, 1843) f thyrsus (Vredenburg, 1921) vagata (E. A. Smith, 1895)

\waihaoensis Finlay, 1924

Unedogemmula MacNeil, 1960 (subgen. of Gemmula)

\bemmeleni (Oostingh, 1941) binda (Garrard, 1961) deshayesii Doumet ( 1839 ) hastula (Reeve, 1843) f hay deni (Vredenburg, 1925) f ickei (Martin, 1906)

fsubsp. virginoides (Vredenburg, 1925) indica (Deshayes, 1832)

\ina MacNeil, 1960 f koolhoveni (Oostingh, 1938) f sondeiana (Martin, 1895) unedo (Kiener, 1839-40). Type

Pinguigemmula MacNeil, 1960 luzonica Powell, new species f okinavensis MacNeil, 1960. Type philippinensis Powell, new species thielei (Finlay, 1930)

[Cryptogemma Dali, 1918]

[benthina Dali, 1908], Type

[Carinoturris Bartsch, 1944]

[adrastia Dali, 1919]. Type

Ptychosyrinx Thiele, 1925

bisinuata (Martens, 1901). Type lobata (Sowerby, 1903) f timorensis (Tesch, 1915)

subsp. teschi Powell, new subspecies truncata (Schepman, 1913) fKuroshioturris Shuto, 1961 (subgen. of Ptychos- yrinx)

\asukana (Yokoyama, 1926)

\hyugaensis (Shuto, 1961). Type f nipponica (Shuto, 1961)

\totomiensis ( Makiyama, 1931 ) f[Coronia Gregorio, 1890] f [childreni Lea, 1833]. Type f [Trypanotoma Cossmann, 1893] f [terebrijormis Meyer, 1886]. Type f [Sinistrella Meyer, 1885]

f [americanus Aldrich, 1885]. Type [[Infracoronia Harris & Palmer, 1947] (subgen. of Sinistrella )

f [ludoviciana Vaughan, 1896]. Type f[Hesperiturris Gardner, 1945] f [nodocarinata Gabb, I860]. Type f [Campylacrum Finlay & Marwick, 1937 f debile Finlay & Marwick, 1937 f sanum Finlay & Marwick, 1937. Type f [Eopleurotoma Cossmann, 1889] f multicostata Deshayes, 1834. Type f[Oxyacrum Cossmann, 1889] (subgenus) f [obliterata Deshayes, 1834]. Type f[Eoturris Finlay & Marwick, 1937 ]complicata (Suter, 1917). Type f multicincta (Marshall, 1917) f neglecta (Suter, 1917)

}uttleyi Suter, 1917 f[Epalxis Cossmann, 1889] f [crenulata Lamarck, 1803]. Type Lucerapex Iredale, 1936

casearia (Hedley & Petterd, 1906). Type subsp. regilla Iredale, 1936 carola (Thiele, 1925)

“denticulata” (Thiele, 1925) indagatoris (Finlay, 1927) molengraaffi (Tesch, 1915 ) f f murrayana (Pritchard, 1904) adenica Powell, new species fOptoturris Powell, 1944 f edita (Powell, 1944)

\optata (Harris, 1897). Type f paracantha (Tenison-Woods, 1877) f kyushuensis Shuto, 1961

[22 - 670]

March 31, 1964

INDO-PACIFIC MOLLUSCA, vol. 1, no. 5

Turrinae 237

Epidirella Iredale, 1931 xanthophaes (Watson, 1886). Type f sayceana (Chapman, 1912)

Epidirona Iredale, 1931 f adelaidensis (Ludbrook, 1941) beachportensis ( Cotton & Godfrey, 1938 ) Candida Laseron, 1954 carinata Laseron, 1954 costifera Laseron, 1954 flindersi ( Cotton & Godfrey, 1938 gabensis (Hedley, 1922) hedleyi Iredale, 1931. Type jaffaensis (Verco, 1909) molleri Laseron, 1954 multiseriata (E. A. Smith, 1877) nodulosa Laseron, 1954 perksi (Verco, 1896) philipineri (Tenison- Woods, 1877) f powelli Ludbrook, 1957 quoyi (Desmoulins, 1842) schoutanica ( May, 1911 ) f suppressa (Finlay, 1927) torquata (Hedley, 1922) tuberculata Laseron, 1954 f vardoni (Tate, 1899)

Lophiotoma Casey, 1904 abbreviata (Reeve, 1843) subsp. lifuensis ( Sowerby, 1907 ) subsp. ustulata (Reeve, 1846) acuta (Perry, 1811). Type albino (Lamarck, 1822) f albinoides ( Martin, 1883 ) brevicaudata (Reeve, 1843) ruthveniana (Melvill, 1923)

Lophioturris Powell, new subgenus indica (Roding, 1798). Type leucotropis (Adams & Reeve, 1850) \odengensis (Martin, 1895) polytropa (Helbling, 1779) f pseudofascialis (Martin, 1883) f [Pleuroliria Gregorio, 1890] f [supramirifica Gregorio, 1890]. Type [Polystira Woodring, 1928]

[albida Perry, 1811]. Type fEchinoturris Powell, 1942 f finlayi Powell, 1935. Type f[Veruturris Powell, 1944 f bisculpta (Powell, 1944)

[cochleata (Powell, 1944) f quadricarinata (Powell, 1944). Type f subconcava (Harris, 1897) f tomopleuroides (Powell, 1944)

f [Cinguliturris Powell, new subgenus of Veru - turns

\tatei (Cossmann, 1896). Type Viridoturris Powell, new genus corona Laseron, 1954. Type Xenuroturris Iredale, 1929 castanella (Tinker, 1952) cingulifera (Lamarck, 1822). Type subsp. erythraea (Weinkauff, 1875) cerithiformis (Tinker, 1952) incredula ( Iredale, 1931 ) kingae Powell, new species millepunctata (Sowerby, 1908)

Turris Roding, 1798 ?ambages Barnard, 1958 amicta (E. A. Smith, 1877) annulate (Reeve, 1843)

Uabtjlonia (Linnaeus, 1758). Type crispa (Lamarck, 1816)

subsp. intricata Powell, new subsp. subsp. variegata (Kiener, 1839-40) subsp. yeddoensis (Jousseaume, 1883) cryptorrhaphe (Sowerby, 1825) garnonsii (Reeve, 1843) f selwyni (Pritchard, 1904) f septemlirata (Harris, 1897) spectabilis (Reeve, 1843) undosa (Lamarck, 1816)

\?ugaliensis Makiyama, 1927 [Antiplanes Dali, 1902]

[perversa (Gabb, 1865 )].f Type [Rectiplanes Bartsch, 1944] (subgen. of Anti- planes )

[santarosana (Dali, 1902)]. Type [Rectisulcus Habe, 1958] (subgen. of Anti- planes )

[motojimai (Habe, 1958)]. Type

Turrinae to Be Dealt with in the Next Part Turridrupa Hedley, 1922, Austroturris Laseron, 1954, Micropleurotoma Thiele, 1929 and Taranis (= Fenestrosyrinx Finlay, 1926 = Alio (Jousseaume) Lamy, 1934). Bathytoma, Micantapex and allied genera, despite the peripheral site of the sinus seem to have more in common with the Borsoniinae.

[22 - 671]

238 Turrinae

A. W. B. Powell

Turridae

Doubtful Taxa in the Turrinae

The following species, mostly from the Tertiary of southeast Asia, cannot be satisfactorily evalu- ated at present, except that all appear to belong to the Turrinae. They are listed below under the genera to which they were ascribed by their re- spective authors.

Clavatula striata Gray, 1826

This is the type of the genus Epideira Hedley, but the type specimen of striata has not been found, and no shell from Western Australia, the presumed type locality, has been found which fits the rather undiagnostic description: “Shell ovate, turreted, whitish brown, with eleven or twelve longitudinal axial interrupted ribs forming long tubercles on the centre of the whorls; the whorls with distant impressed spiral lines near the suture with a rather flattened slightly nodulose band; the mouth rather more than one-third the length of the shell; outer lip thin inside, grooved; tail short, with a linear depression on its columella side; axis ten- twelfths, diameter four-twelfths of an inch.” (in King’s Narrative of a Survey . . . Coasts of Australia, London, 2, appendix, p. 485, [1827] 1826).

Genus Epideira Hedley, 1918 (Proc. Royal Soc. New South Wales 51, p. M79). An unrecog- nised genus with the type, by original designa- tion, Clavatula striata Gray, 1826.

Pleurotoma (Eopleurotoma) adela Cossmann and Pissarro, 1909. Palaeont. Indica, N.S. 3, Mem. No. 1 (India (Ranikot Series, Tertiary)).

Pleurotoma (Eopleurotoma) jhirakensis Cossmann and Pissarro, 1909. Palaeont. Indica, N.S. 3, Mem. No. 1 (India (Ranikot Series, Tertiary)).

Pleurotoma (Gemmula?) simplicissima Thiele, 1925. Deutsch. Tiefsee-Exped., 17(2), p. 209, pi. 23, figs. 3, 3a; 5 x 1.8 mm. (off West Coast of Su- matra, 470 metres ) .

This is certainly not a Gemmula for the simple median carina is plain, and the rest of the shell smooth except for axials on the first two post-nu- clear whorls. It may not even belong to the Tur- rinae for the nature of the sinus is neither figured nor described.

Pleurotoma (Hemipleurotoma) bonneti Cossmann, 1900. Faune Pliocenique de Karikal, p. 30, pi. 2, figs. 11, 13 (India, Karikal (Pliocene)).

Pleurotoma (Hemipleurotoma) bonneti bhagotho- rensis Vredenburg, 1925. Mem. Geol. Surv.

India 50(1), p. 51, pi. 12, fig. 5 (N.W. India, Sind, Nari of Rhagothoro Hill (post Eocene) ). Pleurotoma (Hemipleurotoma) humilis iravadica Vredenburg, 1921. Rec. Geol. Surv. India 53, p. 98, pi. 12, fig. 13 ( Burma, Payagyigon (Kama Series, Neogene)).

Pleurotoma yenanensis Noetling, 1895. Mem. Geol. Surv. India 27(1), p. 42, pi. 10, fig. 3 (Burma (Tertiary)).

Pleurotoma (Hemipleurotoma) yenanensis narica Vredenburg, 1925. Mem. Geol. Surv. India 50(1), p. 48, pi. 1, fig. 7 (N.W. India, Sind, Nari of Bhagothoro Hill (post-Eocene)).

Turns vandervlerki Beets, 1941. Geol. Mijnbouwk Genoot. voor Nederl. en Kalonien, Geol. Ser. 13, p. 7, pi. 7, figs. 273, 274 (East Borneo, Mangkalihat ( Upper Miocene ) ) .

Turns (Gemmula) husamaru Nomura, 1940. Rec. Oceanogr. Works Japan 12, 1, p. 113, pi. 1, figs. 4a, b (Japan, off Tiba Prefecture). See pi. 207

Turns tigrinaeformis Nomura, 1935. Sci. Rep. To- hoku Univ., 18(2), p. 113, pi. 7, figs. 32a, b Formosa [Taiwan Island] (Pliocene)).

Bibliography

This is a list of some of the major works on the Turridae which includes monographs, and papers dealing with the turrids of a specific geographical area or those in which ideas on classification were advanced.

Bellardi, L. 1878. I Molluschi dei terreni terziarii del Pie- monte e della Liguria 2, Torino, pp. 1-364.

Bouge, L. J. & Dautzenberg, Ph. 1914. Les Pleurotomes de la Nouvelle-Caledonie et de ses Dependances. Journ. de Conchyl. 61, pp. 123-214.

Casey, T. L. 1904. Notes on the Pleurotomidae with De- scription of Some New Genera and Species. Trans. Acad. Sci. St. Louis 14(5), pp. 123-170.

Cossmann, M. 1896. Essais de Paleoconchologie comparee, Paris 2, pp. 58-179.

Dali, W. H. 1889. Reports on the Results of Dredging un- der the Supervision of Alexander Agassiz, in the Gulf of Mexico (1877-78) and in the Caribbean Sea (1879-80), by the U. S. Coast Survey Steamer “Blake.” Bull. Mus. Comp. Zool. 18(29), pt. 2, pp. 1-492.

Dali, W. H. 1908. Reports on the Dredging Operations off the West Coast of Central America to the Galapagos, to the West Coast of Mexico and in the Gulf of California in charge of Alexander Agassiz, carried on by the U. S. Fish Commission Steamer “Albatross” during 1891. Bull. Mus. Comp. Zool. 43(6), pp. 205-487.

Dali, W. H. 1918. Notes on the Nomenclature of the Mol- lusks of the Family Turritidae. Proc. U. S. Nat. Mus. 54, pp. 313-333.

[22 - 672]

March 31, 1964

INDO-PACIFIC MOLLUSCA, vol. 1, no. 5

Tuninae 239

Glibert, M. 1960. Les Conacea Fossiles du Cenozoique Etranger. Mem. Inst. Roy. Sci. Nat. Belg. 2(64), pp. 1-132.

Grant, U. S. & Gale, H. R. 1931. Catalogue of the Marine Pliocene and Pleistocene Mollusca of California and Ad- jacent Regions. Mem. 1, San Diego Soc. Nat. Hist., pp. 477-612.

Harris, G. D. 1937. Turrid Illustrations, Mainly Claibornian, Pal. Americana, 2(7), pp. 1-122.

Harris, G. F. 1897. Catalogue of Tertiary Mollusca in the Department of Geology, British Museum (Natural His- tory), pp. 1-407.

Hedley, C. 1922. A Revision of the Australian Turridae. Rec. Aust. Mus. 13(6), pp. 213-259.

Keen, A. Myra, 1958. Sea Shells of Tropical West America. Stanford Univ. Press, pp. 1-624.

Kiener, L. C. 1839-40. Iconographie des coquilles vivantes. Pleurotome 5, pp. 1-84.

Kira, T. 1955. Coloured Illustrations of the Shells of Japan. Hoikusha, Osaka, pis. 1-67.

Laseron, C. F. 1954. The New South Wales Turridae. Roy. Zool. Soc. N.S.W., Zool. Handb., pp. 1-56.

Melvill, J. C. 1917. A Revision of the Turridae (Pleuro- tomidae) Occurring in the Persian Gulf, Gulf of Oman and the North Arabian Sea as Evidenced Mostly Through the Results of Dredgings Carried out by Mr. F. W. Townsend, 1893-1914. Proc. Malac. Soc. London 12, pp. 140-201.

Powell, A. W. B. 1942. The New Zealand Recent and Fossil Mollusca of the Family Turridae. With general notes on Turrid nomenclature and systematics. Bull. Auck. Inst. Mus. 2, pp. 1-192.

Powell, A. W. B. 1944. The Australian Tertiary Mollusca of the Family Turridae. Rec. Auck. Inst. Mus. 3(1), pp. 1-68.

Rasmussen, E. 1951. Faunistic and Biological Notes on Marine Invertebrates. Pt. 2. Vidensk, Medd. Naturh. Foren. Kbli. 113, pp. 201-249, 29 figs.

Reeve, L. A. 1843-46. Monograph of the Genus Pleurotoma. Conchologia Iconica, 1, pis. 1-369.

Reeve, L. A. 1846. Monograph of the Genus Mangelia. Conchologia Iconica 3, pis. 1-8.

Schepman, M. M. 1913. Siboga Expedition Monograph 49(l)e, Part 5; Toxoglossa, pp. 365-452.

Thiele, J. 1925. Gastropoda der Deutschen Tiefsee-Expedi- tion, 17(2), pp. 1-382.

Thiele, J. 1929. Handbuch der Systematischen Weichtier- kunde. Jena, pp. 1-376.

Thorson, G. 1946. Reproduction and Larval Development of Danish Marine Bottom Invertebrates, Medd. Komm. Havunders., Kbh., ser. Plankton 4, pp. 1-523.

Thorson, G. 1950. Reproductive and Larval Ecology of Marine Bottom Invertebrates. Biol. Revue 25, pp. 1-45.

Watson, R. B. 1886. Report on the Scaphopoda and Gas- teropoda Collected by H. M. S. Challenger During the Years 1873-76. Challenger Rep., Zool., 15, pp. 1-756.

Weinkauff, H. C. 1875. Systematisches Conchylien-Cabinet von Martini und Chemnitz. Die Familie Pleurotomidae, pp. 1-248.

Woodring, W. P. 1928. Miocene Mollusks from Bowden, Jamaica, pt. 2. Pub. no. 385, Carnegie Inst, of Washing- ton. pp. 144-201.

van der Vlerk, I. M. 1931. Caenozoic Amphineura, Gas- tropoda, Lamellibranchiata, Scaphopoda. Leidsche Geol- ogische Mededeelingen, Leiden, 5, pp. 206-296.

Acknowledgements

This is the first of what is hoped may be a long series of monographs on Indo-Pacific Turridae. This research is being made possible by the gener- osity of a large number of people who have helped with the project in various ways.

Firstly, the writer is greatly indebted to Mr. W. B. Dixon Stroud who, besides taking a keen interest in the work, generously made available through the Academy of Natural Sciences of Philadelphia a very considerable monetary grant. This grant has enabled the writer to visit museums and other sci- entific institutions in the United States, Great Brit- ain and Australia to examine collections and to re- fer to type material and literature not available in New Zealand.

To the following people the writer gratefully ac- knowledges excellent working facilities provided at their respective institutions and other privileges granted which included the loan of extensive ma- terial: Dr. R. Tucker Abbott, Academy of Natural Sciences of Philadelphia, Dr. Harald A. Rehder, United States National Museum, Dr. L. R. Cox and Mr. Norman Tebble, British Museum (Natural History), Dr. Yoshio Kondo, B. P. Bishop Museum, Honolulu, Dr. D. McMichael, Australian Museum, Sydney, Dr. W. J. Clench, Museum of Comparative Zoology, Harvard, Mr. Allyn G. Smith, California Academy of Sciences, Dr. K. van Winkle Palmer, Palaeontological Research Institution, Ithaca, Dr. A. Myra Keen, Stanford University, Dr. W. K. Emerson and Mr. W. Old, American Museum of Natural History, Dr. A. Solem, Chicago Museum of Natural History and Dr. Dillwyn John, National Museum of Wales (the order of mention is in re- lation to the amount of time spent at each of the above institutions ) .

To Miss Virginia Orr of the Academy of Natural Sciences of Philadelphia it is gratefully acknowl- edged that this work would have been much more difficult of accomplishment had it not been for her meticulous attention to innumerable enquiries re- lating to literature references and above all for the onerous, very considerable help she gave towards the compilation of an illustrated index to the Tur- ridae, a task involving the taking of thousands of photographs.

The writer is deeply indebted also to the follow- ing people who have helped with the project in various ways: Mr. and Mrs. John Q. Burch, Los Angeles, Dr. P. Burgess, Honolulu, Mr. & Mrs.

[22 - 673]

240 Turrinae

A. W. B. Powell

Turridae

Crawford N. Cate, Los Angeles, Dr. R. K. Dell, Dominion Museum, New Zealand, Dr. C. A. Flem- ing, New Zealand Geological Survey, Mr. R. W. Foster, Museum of Comparative Zoology, Harvard, Mr. T. Garrard, Sydney, Dr. Alison Kay, University

of Hawaii, Mrs. J. Kerslake, Sydney, Mrs. M. E. King, Honolulu, Dr. Hope Macpherson, National Museum, Victoria, Mr. P. Nuttall and Mr. John Peake, Rritish Museum (Natural History), Mr. D. Thaanum and Mr. Clifton S. Weaver, Honolulu.

Dr. A. W. Baden Powell is the Assistant Director of the Auckland Institute and Museum in New Zealand, and is the Curator of its Section of Mol- lusca. An internationally known conchologist and author of many scientific and popular articles and books on New Zealand molluscs, Dr. Powell has published extensively on land as well as marine forms. He has devoted many years to the study of the family Turridae, both fossil and Recent, and plans to complete his Indo-Pacific studies in a series of monographs for this journal. Dr. Powell is an accomplished artist and an ardent philatelist, spe- cialising in New Zealand and British Common- wealth stamps. He was born in New Zealand on April 4, 1901.

[22 - 674]

March 31, 1964

INDO-PACIFIC MOLLUSCA, vol. 1, no. 5

Fusiturris 241

Family Turridae

Subfamily Turrinae

Genus Fusiturris Thiele, 1929

Type: Pleurotoma undatiruga Bivona, 1832

Members of this genus still live in the Mediter- ranean and southward along the equatorial coast of West Africa. It has an European ancestry back to the Paleocene and is the European-West African counterpart of the Indo-Pacific Lophiotoma. Fusi- turris and Gemmula have had a parallel develop- ment from the European Paleocene upwards, but apparently Gemmula is no longer living in the European-West African area.

Shell up to 50 mm. (2 inches) in height, elon- gate-fusiform, with a tall spire and long straight anterior canal. Protoconch small, narrowly conic and of three smooth whorls. Spire turreted; sculp- ture of wavy thin axials crescentically thickened at the narrowly rounded periphery. Sinus peripheral, moderately deep and narrowly U-shaped. Opercu- lum leaf -shaped and with an apical nucleus.

Synonymy —

1929 Fusiturris Thiele, Handbuch der Systematischen Weichtierkunde, Jena, 1, p. 361 [type by mono- typy of section of Turris: T. (F.) undatiruga (Bivona ) ] .

1929 Tyrrhenoturris Coen, Atti. Soc. Ital. Milano, 68: p. 297 ( type by Powell’s 1942, p. 22, subsequent designation: Pleurotoma undatiruga Bivona).

Characteristic species —

RECENT : undatiruga Bivona, 1832 ( = balteata and corrugata Kiener, 1839-40, = similis Dautzen- berg, 1891), torta Dautzenberg, 1912.

PLIOCENE: porrecta Wood, 1848.

MIOCENE: aquensis Grateloup, 1832; inermis Hoernes, 1856; mercati Bellardi, 1877; reevei Bel- lardi, 1847.

OLIGOCENE: conifera Edwards, 1861; difficilis Giebel, 1864; duchastelii Nyst, 1836; explanata Koenen, 1890; flexiplicata Kautsky, 1925; koeneni Glibert, 1860; plana Giebel, 1864; selysi Koninck, 1837.

EOCENE: infraeocaenica Cossmann, 1889; lau- brierei Cossmann, 1889; prestwichi Edwards, 1861; wetherelli Edwards, 1861.

Plate 183. Fusiturris undatiruga (Bivona, 1832). Recent; Mediterranean. 46 mm. Type of Fusiturris.

Plate 184. Marginal radular tooth of Fusiturris undatiruga (Bivona, 1832) (from J. Thiele, 1929, p. 361, fig. 441).

[22 - 685]

242 Fusiturris

A. VV. B. Powell

Turridae

[These occasional blank areas occur between genera and subgenera to permit the insertion of new material and future sections in their proper systematic sequence .]

[22 - 686]

March 31, 1964

INDO-PACIFIC MOLLUSCA, vol. 1, no. 5

Gemmula 243

The Gemmate Series

The genus Gemmula has the most extensive Re- cent geographical range of any of the Turrinae and it also extends back to at least the beginning of the Tertiary. It is the most vigorous member of the Turrinae and undoubtedly represents the main stem of the subfamily. Gemmula is well-represented in the Tertiary of southern United States, Europe, India, Burma, Indonesia, Japan, Australia and New Zealand. The greatest development of Gemmula is now in the Indo-West Pacific. Very few species still exist in Caribbean-Panamic waters, and as al- ready mentioned there are none from the Mediter- ranean and West Africa where the genus is re- placed by Fusiturris.

Both Gemmula and Fusiturris seem to have had a common origin in the European Paleocene or ear- lier and both were well-represented throughout the Tertiary. All of the Eocene-Oligocene derivatives of Gemmula were relatively short-lived, as for ex- ample the European-American Eopleurotoma, the European Epalxis, H emipleurotoma and Oxyacrum , as well as a group of New Zealand Upper Creta- ceous-Eocene genera, Campylacrum, Eoturris and possibly Tholitoma.

Gemmula has the posterior sinus as a deep slit on the peripheral carina and in general the shell resembles Lophiotoma, except for the character- istic gemmate sculpture on the peripheral sinus rib.

The radula in Gemmula consists of a pair of marginals in the shape of a “wishbone,” a type of dentition common to Tunis, Lophiotoma, Epicli- rona and some members of the Turriculinae.

Genus Gemmula Weinkauff, 1875

Type: Gemmula hindsiana Berry, 1958 It may appear that only small differences sepa- rate the bracket of names, Tunis, Gemmula and Lophiotoma, especially the fact that the presence of gemmules on the peripheral carina is the only obvious difference between Gemmula and Lophio- toma, which has a plain peripheral sinus band. However, these differences, which appear slight when Recent material only is reviewed take on more significance when phylogeny is considered.

Synonymy —

1875 Gemmula Weinkauff, Jahrbiich. der Deutschen Mai. Gesell., 2: p. 287. Type by subsequent designation by Cossmann, 1896, p. 62: Pleurotoma gemmata Reeve, 1843 (non Conrad, 1835) = Gemmula hind- siana Berry, 1958.

Plate 185. Gemmula hindsiana Berry, 1958. Gulf of Cal- ifornia. Formerly Pleurotoma gemmata Reeve, 1843, and Hinds, 1843; non Conrad, 1835 ( from Harris, 1937, Paleont. Americana 2 (7), pi. 1. fig. 33). Type of Gemmula.

Key to the Gemmate Genera

A. Protoconch polygyrate and axially costate; peripheral keel gemmate throughout.

1. Shell elongate-fusiform, sinus deep and narrow Gemmula, p. 22-695

2. Shell truncated anteriorly; sinus broadly V-shaped Ptychosyrinx, p. 22-851

3. Shell broadly conic; keel placed low; sinus open 45° .... Pinguigemmula, p. 22-789

B. Protoconch polygyrate and smooth, except for half whorl of axials; keel gemmate on early

whorls only; shell elongate-fusiform Unedo gemmula, p. 22-761

C. Protoconch paucispiral, globose and smooth; sinus broadly V-shaped.

1. Shell truncated anteriorly Kuroshioturris, p. 22-865

2. Shell elongate-fusiform and smooth; except for a peripheral row of tubercles.

Lucerapex, p. 22-837

[22 - 695]

244 Gemmula

A. W. B. Powell

Turridae

1931 Eugemmula Iredale, Records Australian Mus., 18(4): p. 226. Type by original designation: E. hawleyi Iredale, 1931.

In addition to the large number of Indo-Pacific species either mentioned or described in the fol- lowing text, Gemmula is abundantly represented in the Tertiary of both Europe and the United States of America.

Characteristic European Tertiary Species —

PALEOCENE: gryi Ravn, 1939. EOCENE: acu- tangularis Deshayes, 1834; aspera Edwards, 1861; callifera Edwards, 1861; cancellata Deshayes, 1834; gentilis Sowerby, 1850; goosensi Boury, 1899; lon- gaeva Edwards, 1861; monerma Edwards, 1861; nilssoni Deshayes, 1865; pleheia Sowerby, 1850; reticulosa Edwards, 1861; simillima Edwards, 1861; subcarinata Rouault, 1850; submonilifera Boury, 1899; taeniolata Edwards, 1861; tenuistriata Desh- ayes, 1834; uniserialis Deshayes, 1834; varians Ed- wards, 1861.

OLIGOCENE: bosqueti Nyst, 1843; humilis Koe- nen, 1890; laticlavia Beyrich, 1848; lunulifera Koe- nen, 1890; nodigera Koenen, 1890, odontella Ed- wards, 1861; odontophora Koenen, 1890; parkinsoni Deshayes, 1865; subdentata Goldfuss, 1844.

MIOCENE: annae Hoernes and Auinger, 1891;

badensis Hoernes and Auinger, 1891; contigua Brocchi, 1814; coronata Goldfuss, 1844; coronifera Bellardi, 1877; cossmanni Peyrot, 1931; cypris Or- bigny, 1852; denticula Basterot, 1825; disjuncta Peyrot, 1931; rotula Brocchi, 1814; spiralis M. de Serres, 1829; stoffelsi Nyst, 1843.

PLIOCENE: antwerpiensis Vincent, 1890; moni- lis Brocchi, 1814; turrifera Nyst, 1853.

Characteristic American Tertiary Species —

EOCENE: alternata Conrad, 1833; carodenta Harris, 1937; casteri Harris, 1937; conjuncta Casey, 1904; coraliger Harris, 1937; lancea Casey, 1904; lerchi Vaughan, 1896; ludocarola Harris, 1937; plen- topsis Harris, 1947; sublerchi Harris, 1937; watele- tella Harris, 1937; weisbordi Harris, 1937.

OLIGOCENE: arnica Casey, 1903; ancilla Casey, 1903; tenella Conrad, 1847.

It is possible that true Gemmula is represented in the Eocene-Miocene horizons of the west coast of North America, but material examined so far discounts this. The nearest approach to a Gemmula is Surcula monilifera Cooper, 1894, but although this Californian Eocene species has a beaded peri- pheral keel, the smooth conical three-whorled pro- toconch is atypical.

Plate 186

Figs. 1 Gemmula speciosa (Reeve). Holotype of Pleuro- toma carinata Griffith and Pidgeon, 1834, non Link, 1808. Australia. 73 mm.

2, 3 Gemmula kieneri ( Doumet ) . 40 fathoms, off Tosa,

Japan. 61 mm.

4 Gemmula murrayi new species. Holotype. Off

Sharam, west coast of the Gulf of Oman, Saudi Arabia. 35.5 mm.

5 Gemmula dampierana new species. Holotype. 7

mi. N.N.W. of Anchor Island. Onslow, Western Australia. 35.5 mm.

[22 - 696]

March 31, 1964

INDO-PACIFIC MOLLUSCA, vol. 1, no. 5

Gemmula 245

Gemmula speciosa (Reeve, 1843)

(PI. 186, fig. 1)

Range — Philippines, China Sea and Arabian Sea.

Remarks — This very distinctive and uncommon species is at once recognized by its broad, cog- wheel-like peripheral keel which is regularly and closely studded with laterally compressed, raised gemmules. Further, there is a distinctive colour pattern of light-brown, continuous, spiral lines con- fined to the peripheral keel and the primary spiral cords. The closely allied Gemmula kieneri (Dou- met) differs in having a heavier and less projecting peripheral keel, stronger interstitial spirals, as well as a more prominent subsutural fold. The colour markings of kieneri are not continuous lines, but are interrupted dots and dashes occurring between the peripheral gemmules and on the primary spirals.

Considerable confusion exists concerning the cor- rect name to be used for this species, due to the erroneous and repeated statements that carinata Gray is synonymous with kieneri Doumet. Cer- tainly, Reeve’s 1843 carinata (Gray) appears to be identical with kieneri, but Gray’s (in Griffith and Pidgeon, 1834) original carinata is undoubtedly the same as Reeve’s speciosa. The latter name must be employed because Gray’s name is preoccupied by Pleurotoma carinata Link, 1808. Because of this confusion, literature records of this species are not included here.

Melvill (1917, p. 145) described a Tunis ( Gem-

mula) granosa guadurensis, from the Mekran Coast in 70 fathoms, as follows: “Testa ut in typo, sed omnino minor, fere immaculata, M. C. Gwadur, one specimen at 70 fathoms. A few others, all much of the same calibre, off Ras Maidani, between Gwadur and Jask. The sculpture of this variety is identical with the type; the size about half, say 38 mm., as against 60-70 mm., the coloration most simple, nearly immaculate.”

Although the actual type of guadurensis was not located, an apparently authentic topotype from the late Mr. J. R. le B. Tomlin shows this alleged sub- species to be merely an immature speciosa, for it fits into the size range of a series of that species from Station 72, John Murray Expedition, from the Gulf of Oman in 73 metres.

These shells are identical in sculpture with spe- ciosa but are almost lacking in coloration, although a faint pattern does show faintly in some of the larger examples. The Tomlin specimen is 37 mm. in height and has 7 post-embryonic whorls, in con- trast to 8 or 9 in adult speciosa. From the above observations it is clear that guadurensis is not a dwarf form, as Melvill believed it to be, but merely an immature speciosa.

Melvill made guadurensis a subspecies of granosa (= kieneri) but that species has fewer peripheral gemmules. Counts showed that in speciosa the gemmules range between 33 and 46 per whorl (43-46 in adults), and in adult kieneri, 31-35 per whorl.

Plate 187. Geographical distribution of Gemmula kieneri (Doumet), G. speciosa (Reeve) and G. murrayi new species.

[22 - 697]

246 Gemmula

A. W. B. Powell

Turridae

Fossil specimens are difficult to assign by illustra- tions alone, but it would appear that of the figured Upper Tertiary shells assigned to carinata by Tesch, 1915, only fig. 47a resembles speciosa, the remainder on plates 76 and 77 being sculpturally more in accord with kieneri.

Description — Adult shell 53.5 to 73 mm. ( 2 to 3 inches) in height. Fusiform, with tall spire and long tapered and slightly flexed anterior canal. Whorls strongly angulate and carinate at just be- low middle whorl height; base rather suddenly con- tracted. Peripheral carina a square-cut prominent flange densely studded with narrow, laterally com- pressed and peaked nodules which give a regular cog-like effect. Primary spirals plain, thin, but sharply raised, three above the carina, one of which is on a moderate subsutural fold, one or two below the carina, and about six on the base, exclusive of the canal. The secondary sculpture consists of from one to three plain weak threads in the interspaces of the primaries. The surface is crowded with weak, crisp, axial threads. Colour pattern of light- brown spirals on a buff ground.

The whole of the peripheral carina is uniformly coloured and all of the primary spirals are sim- ilarly tinted light-brown. There are no interrupted markings, dots or dashes.

Measurements (mm.) —

height	width
73.0	26.0	type of carinata G. and P.
73.0	21.3	Samar Id., 35 fms., Philippines
53.5	19.0	Mantaquin Id., 27 fms., Philippines
Synonymy	-

1834 Pleurotoma carinata (Gray) in Griffith & Pidgeon, Moll. & Radiata, arranged by Baron Cuvier, Lon- don, p. 599, PI. 23, f. 2. Non Pleurotoma carinata Link, 1808; non carinata (Gray); Reeve, 1843, Conch. Icon. 1, pi. 7, fig. 56.

1843 Pleurotoma speciosa Reeve, Conch. Icon., 1, pi. 2, fig. 9.

1884 Pleurotoma ( Gemmula ) speciosa Reeve, Tryon, Manual of Conch. 6, p. 173, pi. 4, fig. 48.

1915 Pleurotoma carinata Gray, Tesch, Palaont. von Timor, pi. 77, fig. 47a (only).

1917 Tunis ( Gemmula ) guadurensis Melvill, Proc. Malac. Soc., 12, p. 145.

Types — British Museum (Natural History), no locality (carinata).

Records — PHILIPPINES: off Badian Island, west Samar, 35 fms. (Albatross Sta. 5426); off Mantaquin Island, east coast, Palawan, 27 fms. (Albatross Sta. 5207) (USNM); Samar (coll, of A. D’Attilio, New York). ARABIA: Gulf of Oman, 73 metres (John Murray Exped. Brit. Mus. (Nat. Hist.)).

Fossil Records — Recorded from the Upper Tertiary of Timor (Tesch, 1915, p. 25, pi. 77, fig. 47a, only).

Gemmula kieneri (Doumet, 1840)

(PI. 186, figs. 2, 3)

Range — Japan, China and the Philippines.

Remarks — Under speciosa Reeve, I have already pointed out that kieneri is very similar, but the peripheral keel in the latter is less projecting, the body whorl not so abruptly or so deeply contracted, and the colour pattern is in the form of macula- tions between the gemmules of the carina and in- terrupted dots and dashes on the primary spirals. In speciosa, all the colour lines are uninterrupted.

The species name granosa Helbling, 1779, has long been in misuse for this species which is com- mon in Chinese and Japanese waters. Helbling’s figure, however, depicts a misshapen specimen with a short anterior canal. There is a band of rounded gemmules subsuturally, the peripheral carina is studded with closely spaced, strong, round gem- mules and all of the spirals below the peripheral keel are shown to be strongly and closely gemmate. It could be just a case of faulty draftsmanship but on the other hand Helbling’s figures of other spe- cies in the same work are accurate and the species easily recognized.

No locality was given by Helbling and it is doubtful if the type specimen is still in existence. The wisest course is to drop the name as indeter- minable and for this Oriental species employ Dou- met’s name, kieneri. The original reference is Murex (Fusus) granosa Helbling, 1779, Abhandl. Priv. Bohm. Math., Prag., 4, p. 116, pi. 2, fig. 22.

Description — Adult shell 56 to 73 mm. (21* to 3 inches ) in height, robust, fusiform, with tall spire and long rather straight anterior canal. Adult whorls, 10 or 11, rather tightly coiled. Spire whorls sculptured with a strong square-cut keel, situated below the middle, not prominently projecting and sculptured with numerous closely spaced rectangu- lar nodules, which are laterally compressed. There is a strong subsutural fold bearing one primary cord and two threads, three or four sharply raised slightly imbricate threads between the subsutural fold and the peripheral keel and one primary cord and several threads between the periphery and the lower suture. Base, exclusive of the anterior canal, with about six primary cords and from one to three interstitial threads. Surface covered with dense lamellate axial growth threads which imbricate the secondary spiral threads. Colour maculated in light- brown on a white ground, with some obscure and irregular small blotches on the subsutural fold, with regular squarish spots between the gemmules on

[22 - 698]

March 31, 1964

INDO-PACIFIC MOLLUSCA, vol. 1, no. 5

Gemmula 247

the carina and with irregularly disposed spots on the primary cords of the base.

Measurements (mm.) —

height width

73.0 21.0 “India” (ANSP)

59.5 20.0 Tosa, Japan

56.0 17.5 Kii, Japan

Synonymy -

1840 Pleurotoma kieneri Doumet, Magasin de Zool. 2, p. 2, pi. 10.

1843 Pleurotoma carinata Gray, Reeve (non Gray, 1834), Conch. Icon. 1, pi. 7, fig. 56.

1909 Pleurotoma carinata Gray, Annandale & Stewart, Illus.

Zool. Investigator, Moll. 6, pi. 20, figs. 3, 4.

1915 Pleurotoma carinata Gray, Tesch, Palaont. von Timor, pis. 76 and 77 (all figs, of carinata except 47a). 1955 Gemmula granosa “Helbling,” Kira, Coloured Illus. Shells of Japan, Osaka, pi. 35, fig. 18.

Records — JAPAN : Tosa, 40 fathoms; Kii (ANSP); Sa- gami Bay, 40 fathoms (USNM). HONGKONG (USNM). PHILIPPINES: West of Pacyan Island, 189 fathoms, green mud (Albatross Sta. 5409); between Cebu and Bohol, 162 fathoms (Albatross Sta. 5412) (USNM).

Fossil Records — Recorded from the Upper Tertiary of Timor (Tesch, 1915, p. 25, as carinata ), pis. 76 and 77, all figures of “carinata,” except 47a.

Gemmula kieneri subspecies ryukyuensis MacNeil, 1960

(PI. 188, fig. 3)

Range — Okinawa, Miocene or Pliocene.

Remarks — “This subspecies has a broader sub- sutural slope than is common for the species and the subsutural collar is weaker. It has fewer and larger nodes on the peripheral carina. All of the specimens at hand have two or three coarse spirals below the periphery, those below on the base of the body whorl and columella becoming much weaker. The latter condition is approached by oc- casional specimens of the more typical form of the species but on ryukyuensis it is consistent” (Mac- Neil, 1960, p. 103).

MacNeil’s figure shows a shell allied to kieneri by the presence of a bicordate subsutural fold, reminiscent of cosmoi in the three heavier spirals of the upper base, but distinct from both in the fewer peripheral gemmules.

Measurements (mm.) —

height width

47 15 holotype, USNM 562975.

Synonymy —

1960 Gemmula granosa ryukyuensis MacNeil, U. S. Geol. Surv. Prof. Paper 339, p. 103, pi. 14, fig. 24.

Records — OKINAWA and Takabanare-shima ( Miocene, Yonabaru clay member); Okinawa, Shinzato tuff member (Miocene or Pliocene) (Type locality).

Gemmula kieneri subspecies woodwardi (Martin, 1884)

(PI. 188, fig. 2)

Range — Indonesia, Pliocene.

Remarks — Judging from the original illustration, Pleurotoma woodwardi Martin is probably a syn- onym of kieneri Doumet, 1840 ( = granosa auct. ) . Both Makiyama, ( 1927, p. 95 ) and Nomura ( 1935, p. 114) concur in this view but since Oostingh (1938, p. 27) makes it a subspecies of “ granosa ” and I have not seen the relevant material, Oostingh’s view is accepted for the present.

Measurements (mm.) — Size not indicated. Synonymy —

1884 Pleurotoma woodwardi Martin, Samml. Geol. Reichs- Mus., Leiden, Ser. I, 3, p. 56, pi. 4, fig. 57.

1938 Turns (Gemmula) granosa woodwardi Martin, Oostingh, De Ingenieur in Ned.-Indie, Gast. 1(7) p. 27.

Records — PLIOCENE: deep boring (130 metres) near Batavia, Java Id., Indonesia (type locality); South Bantam, Indonesia, Pliocene (Oostingh, 1938).

Plate 188. Fig. 1, “Murex (Fiisus) granosa” Helbling (from the original), an indeterminate species. Fig. 2, Gem- mula kieneri woodwardi (Martin), Pliocene of Sonde, Java. 22.5 mm. (photo courtesy of C. P. Nuttall, Brit. Mus. ). Fig. 3, Gemmula kieneri ryukyuensis MacNeil. Holotype. Mio- cene of Okinawa Id., Ryukyu Ids. 47 mm. (from the orig- inal, pi. 14, fig. 24 ) .

[22 - 699]

248 Gemmula

A. W. B. Powell

Turridae

Gemmula murrayi new species (PL 186, fig. 4)

Range — Persian Gulf.

Remarks — This species stands nearest to speciosa (Reeve). It has the same distinctive colour pat- tern with all the primary spiral cords continuously lined in light -brown on a pale buff ground. It is of smaller adult size and is much narrower than spe- ciosa and the peripheral keel is not so prominent.

Description — Adult shell 35 to 40 mm. (about 1/2 inches) in height, elongate-fusiform, with a tall spire and long straight canal; spire angle 28° to 30°. Whorls 11-11/2, plus a tall narrowly conic axially costate protoconch of 3M whorls. Spire whorls sculptured firstly with a prominent sub- sutural fold which is narrowly crested by a smooth spiral cord and has two secondary spiral threads both above and below it, three or four smooth spiral cords on the broad, steep and lightly concave shoulder area to the not very prominent gemmate peripheral sinus ridge, which is composed of two closely spaced cords with the gemmules vertically fused, 39-40 on the penultimate. One primary spiral and a few very weak threads present between the peripheral carina and the lower suture. On the up- per part of the base there are six rather wide- spaced primary spirals and below, over the neck and canal, there is an alternation of closely spaced primary and secondary spirals. Sinus moderately deep, U-shaped, its apex wider than the peripheral keel. Colour pale creamy buff, the sinus rib and all major spirals lined in light-brown. The main spiral of the subsutural fold is darker brown than any of the other spirals.

Measurements (mm.) —

height width

40.0 11.0 holotype

38.0 11.0

33.0 9.0

Type Locality - 25° 38' 18" N., 56° 26' 36" E„ 73 metres (John Murray Expedition, Sta. 72): off Sharam, west coast of the Gulf of Oman.

Types — The holotype is in the British Museum (Natural History).

Gemmula dampierana new species (PI. 186, fig. 5)

Range — North West Australia, the Dampierian Marine Province.

Remarks — This species is closely allied to the preoccupied Pleurotoma concinna Dunker, 1856.

Dunker described two species with the specific name concinna and confusion later arose through the action of Tryon ( 1884, p. 335 ) in combining both species in the synonymy of reeveana De- shayes, 1863. Actually, Dunker’s two species are very different; that of 1871 in Malak. Blatt. 18, p. 160 is a Hemidaphne, and the earlier one, 1856, Proc. Zool. Soc., p. 356, is a Gemmula. Dunker’s 1856 concinna was described from unknown lo- cality but “Andaman Islands” has since been pen- cilled on the type tablet. This type specimen fits Dunker’s description (in Latin) very well, even to the violet staining of the anterior end.

The species name provided above is not a new name for the preoccupied concinna but a new proposition for a very constant little Gemmula which comes from several dredgings in North West Australia at depths between 46 and 65 fathoms.

This species resembles murrayi n. sp. described above but has a wider spire angle, the peripheral gemmules are fewer, the colour pattern is confined to the subsutural fold and to the interstices of the gemmate keel, and the secondary spiral sculpture is stronger.

Nothing exactly matching Dunker’s concinna is known to the writer, so this preoccupied species is left unnamed until it is rediscovered from some definite locality. A specimen from the Andaman Is- lands in the United States National Museum, at- tributed to concinna, lacks the overall brown colour and violet stained anterior end, and is in fact in- separable from the South African gilchristi.

Description — Adult shell 30 to 35 mm. ( about 1/4 inches) in height, elongate-fusiform, with a tall spire and long straight canal. Spire equal to height of aperture plus canal, angle 33° to 35°. Whorls 8, plus a conical protoconch of 3M whorls, first whorl smooth, remainder axially costate. Spire whorls sculptured firstly with a strong narrowly crested subsutural fold, four crisp spirals on the concave shoulder area, then the prominent gem- mate bicarinate peripheral keel, the gemmules ver- tically fused and numbering 31 or 32 on the pe- nultimate. One strong spiral cord with one or two secondary spirals both above and below it, between the peripheral carina and the lower suture. Upper base with four strong primary cords and one or two intermediates; neck and anterior end with an alter- nation of closely spaced cords and threads. Sinus deep and narrow, U-shaped, its apex no wider than the peripheral carina. Colour white with the sub- sutural folds and interspaces of the gemmules on the carina light reddish brown.

[22 - 700]

March 31, 1964

A. W. B. Powell

Gemmula 249

Measurements (mm.) —

height width

35.5 11.0 holotype

30.7 10.7

Holotype — Western Australian Museum, Perth.

Records — NORTH WESTERN AUSTRALIA: 7 mi.

N.N.W. of Anchor Island, Onslow, 46 fms. (type locality); 10 mi. N.N.W. of Anchor Island, Onslow, 65 fms.; 20 mi. N.W. of Anchor Island, Onslow, 65 fms. (Western Aus- tralian—Hawaiian Expedition, 1960).

Gemmula gilchristi (Sowerby, 1902)

(PI. 189, figs. 1, 2)

Range — South Africa, Zanzibar, Andaman Is- lands and Japan.

Remarks — This species has been frequently mis- identified as monilifera Pease, 1860. It was differ- entiated by Sowerby, 1902, as a new species, Pleurotoma gilchristi, a now well-known South African shell.

Description — Adult shell 34 to 37 mm. ( lh to 1/2 inches) in height, elongate-fusiform, with a tall spire and moderately long, slightly flexed anterior canal. Spire greater than height of aperture plus canal; angle 35° to 37°. Adult whorls 9 plus a narrowly conic protoconch of 3/2 whorls, smooth at first but strongly axially costate over the last VA whorls. Spire whorls with a heavy subsutural fold, a median flange-like gemmate keel and a single strong smooth cord below. The subsutural fold is sharply keeled at the middle and the peripheral or sinus keel is double, densely sculptured with ver- tically fused gemmules, about 44 on the penulti- mate. Three interstitial threads between the sub- sutural fold and the peripheral keel and one in each interspace below the keel. Body whorl with four smooth primary cords below the peripheral keel, number two level with the suture and an alternation of spiral cords and threads over the base. About ten weak closely spaced spirals on the weak anterior fasciole. The whole surface between the spirals is crowded with fine lamellate axial threads. Sinus moderately deep, U-shaped, situated at the termination of the gemmate peripheral keel. Colour uniform golden to light reddish brown for shallow water shells; topotypes (55 fathoms) are white, sometimes with pale-brown between the peripheral gemmules.

Measurements (mm.) —

height	width
34.6	10.7	Kii, Japan
34.0	10.7	Durban
32.5	10.5	Durban
32.0	11.0	holotype
28.7	9.0	Andaman Ids. (USNM)

Synonymy —

1897 Pleurotoma monilifera Pease, Sowerby, Append. Ma- rine Shells S. Africa, p. 2 (non Pease, 1860).

1902 Pleurotoma gilchristi Sowerby, Marine Invest, in S.

Africa, Cape Town, p. 99, pi. 2, fig. 9.

1958 Tunis gilchristi Sowerby, Barnard, Ann. S. African Mus. 44, p. 106, fig. 3h, 6d.

Types — The holotype is in the British Museum ( Natural History ) .

Records — SOUTH AFRICA: off mouth of Tugela River, 55 fms. (type locality for gilchristi ); Zululand and Natal Coast, 27 to 90 fms.; off Cape Natal, 185 to 200 fms.; Hood Point, East London, 49 fms. ZANZIBAR: lM mi. W.S.W. of Ras Mungwe, 8 fms., fine sand, grass and shell ( ANSP). ANDAMAN ISLANDS: (USNM). JAPAN: Kii ( USNM ).

Gemmula monilifera (Pease, 1860)

(PI. 189, figs. 3, 4)

Range — Hawaiian Islands and Fiji Islands.

Plate 189. Figs. 1 and 2, Gemmula gilchristi (Sowerby). 55 fms., off the mouth of Tugela River, South Africa. 32 mm. Figs. 3 and 4, G. monilifera (Pease). Holotype from Brit. Mus. Hawaiian Ids. 29.4 mm. ( photo by Alison Kay ) .

[22 - 701]

250 Gemmula

A. W. B. Powell

T urridae

Remarks — This shell is widely known in collec- tions under Dali’s manuscript name, aelomitra, and in effect this name was legalised in 1952 by the ac- tion of Tinker who figured and described the spe- cies under Dali’s manuscript name in “Pacific Sea Shells,” p. 46.

Dali intended aelomitra as a new name for the Hawaiian. Turris monilifera Pease, 1860, which he presumed was preoccupied by Pleurotoma monili- fera Lea, 1833, an American Eocene species con- sidered by Harris (1937, p. 31) to be synonymous with Eopleurotoma saiji (Lea, 1833). Since Pease and Lea described their respective species under different generic names, and each is now located in a different genus, i.e., Gemmula and Eopleurotoma, there is no name conflict, and monilifera Pease, 1860, may stand.

In 1869, Pease again described a monilifera but under the genus name Pleurotoma ; it also is from the Hawaiian Islands and is presumably the same as his 1860 proposition. This point is of no conse- quence, however, since the 1869 combination is in- validated by Lea’s earlier proposition.

Description — Adult shell 22 to 29 mm. ( about one inch) in height, narrowly fusiform, with a tall spire and moderately long, almost straight, anterior canal. Spire greater than height of aperture plus canal; angle 25° to 27°. Adult whorls 8 or 9, plus a narrowly conic protoconch of three brown whorls, the first two smooth, the third closely axially cos- tate. Spire whorls with a prominent but narrowly crested subsutural fold, followed by a deep shoulder concavity, bearing 3 or 4 crisp spiral threads. Then a heavy peripheral sinus-keel, which is closely studded with two series of gemmules,

vertically fused, resulting in a cog-wheel effect. A single stout smooth spiral cord below the keel and a second one sometimes emergent over the last whorl. Body whorl with four primary smooth cords, followed by weaker cords below, becoming more closely spaced over the neck and anterior end; one or two spiral threads in each interspace. Anal sinus of moderate depth, U-shaped, its apex occupying the full width of the peripheral carina. Colour yel- lowish to light reddish brown, with the gemmate peripheral keel and the lower base picked out in white.

Measurements (mm.) —

height	width
29.4	9.5	holotype
28.0	9.0	all off W
25.5	8.5
23.0	7.5
22.7	7.5
Synonymy	-

1860 Turris monilifera Pease, Proc. Zool. Soc., London, p. 398.

1869 Pleurotoma monilifera Pease, Amer. Jour. Conch. 5, p. 68 (not of Lea, 1833).

1875 Pleurotoma ( Gemmula ) monilifera Pease, Weinkauff, Tahrb. Deutsch. Malak. Ges. 2, p. 289, pi. 9, figs. 1, 3.

1952 Turris aelomitra (Dali, ms.) Tinker, Pacific Sea Shells, p. 46, pi. 1 (lower row).

Types — Pease’s holotype is in the British Mu- seum (Natural History). We are indebted to Dr. Alison Kay of Hawaii for its photograph.

Records — HAWAIIAN ISLANDS : “Sandwich Islands,” Cuming coll., (type locality); Oahu, off Waikiki, 33-50 fms.; entrance to Honolulu Harbour, 6-8 fms.; entrance to Pearl Harbour, 12-25 fms.; Maui, off Kaanapali; off Mt. Lihau, 4-12 fms. (D. Thaanum coll.); Oahu, off Diamond Head, 100 fms. (P. Burgess). FIJI: Momi Bay, S.E. of Nabilo Light, 13-15 fms. (W. Jennings); “Viti Islands” (A. Garrett, ANSP).

[22 - 702]

March 31, 1964

INDO-PACIFIC MOLLUSCA, vol. 1, no. 5

Gemmula 251

The Congener Series

Key to the Recent subspecies of G. congener

A. Base with 4 spirals more prominent than the rest

Shoulder area moderately wide and shallow Subsutural fold unicarinate, slight

congener cosmoi Subsutural fold bicarinate, slight

congener diomedea

B. Base with spirals gradually diminishing

Shoulder area a deep and narrow cleft Subsutural fold bicarinate, massive

congener congener

The distributional areas for this group of three Recent subspecies is in the tropical Indian Ocean and Persian Gulf through the East Indies to the Philippines ( for congener congener ) ; off the Philip- pines in 100 to 310 fathoms (for congener dio- medea) and in Japanese waters, 50 to 100 fathoms (for congener cosmoi ).

Gemmula congener subspecies congener (E. A. Smith, 1894)

(PL 191, figs. 1-4)

Range — East Africa to the Philippines, 100 to 185 fathoms.

Remarks — This subspecies is characterised by the massive development of both the subsutural fold and the peripheral carina. So strongly devel- oped are these two features that the shoulder is reduced to a narrow deep cleft. The spire varies in height between 1.2 and 1.3 times the height of the aperture plus the canal.

Specimens from two John Murray Expedition East African stations in 212 to 310 metres, are beau- tifully maculated in reddish brown on a white

Plate 191. Figs. 1-4, Gemmula congener congener (E. A. Smith). Figs. 1, 2, from 150 fms., off Colombo, Ceylon. 45 mm.; figs. 3, 4, from 128 fms., off Arena Point, Luzon Id.,

ground; they are spotted on all the spirals and those between gemmules on the bicarinate peri- phery are vertically confluent. The specimens are strongly but sparsely lirate within the outer lip and two examples exhibit tubular distortion of basal spirals. This East African maculated form may yet prove to be subspecifically distinct, for it tends also to have a relatively taller spire. The coloration of congener from elsewhere is, as in cosmoi and dio- medea, confined to a light-brown band coincident with the subsutural fold.

Description — Adult shell 44 to 63 mm. ( 1/4 to 2M inches ) in height. Post-embryonic whorls 9, rather tightly coiled. Subsutural fold of two strong, densely gemmate, closely spaced spiral ridges. The massive peripheral keel is composed of two closely spaced, densely gemmate, rounded heavy spirals. One or two lesser spirals are between the peripheral carinae and the lower suture, and there are about ten primary spirals on the base, plus intermediate threads which extend densely below over the an- terior canal. The primary basal spirals diminish gradually with no especially prominent ones. All of the spirals are rendered gemmate to some de- gree by the crossing of dense strong axial growth lines. Colour (typically) dull- white except for a light-brown band covering the subsutural pair of cords.

Measurements (mm.) -

height	width
63.0	18.0	Balayan Bay, Luzon, 159 fms.
56.0	18.0	off Taal, Luzon, 177 fms.
56.0	18.0	off East Africa, 310 metres
55.0	16.2	off East Africa, 310 metres
52.0	17.0	Bay of Bengal, 128 fms. (holotype)
48.5	15.0	Opol, Mindanao, 214 fms.
45.0	16.0	off Colombo, 150 fms.
38.0	13.0	Andaman Islands, 185 fms.

Philippines. 44 mm. Figs. 5, 6, Gemmula congener diomedea new subspecies. Holotype. 256 fms., off Apo Id., Negros Id., Philippines. 73 mm.

[22 - 713]

252 Gemmula

A. W. B. Powell

Turridae

Synonymy —

1894 Pleurotoma congener E. A. Smith, Ann. Mag. Nat.

Hist., Ser. 6, 14, p. 160, pi. 3, figs. 4, 5.

1913 Pleurotoma (Gemmula) congener Smith, Schepman, Siboga Exped., Pt. 5, 49e, p. 403.

1917 Tunis (Gemmula) congener Smith, Melvill, Proc. Malac. Soc. 12, p. 144.

Type — Indian Museum, Calcutta.

Records — PERSIAN GULF (Melvill, 1917). INDIA: Bay of Bengal, 128 fms. (type locality). CEYLON: west of Colombo, 150 fms. (USNM). ANDAMAN ISLANDS: 185 fms. (Investigator Exped., Brit. Mus. ). EAST AFRICA: 5° 36' 12" S., 39° 13' 12" E., 310 metres; 5° 38' 54" S., 39° 15' 42" E., 212 metres ( Tohn Murray Exped., Brit. Mus.). INDONESIA (Schepman, 1913). PHILIPPINES: off Arena Point, Luzon, 128 fms., Sta. 5382; S.W. of Corregidor Lt., Luzon, 118 fms., mud and shell, Sta. 5272; Tayabas Bay, Luzon, 150 fms., dark green mud, Sta. 5372; Batangas Bay, 170 fms., Luzon, Sta. 5268; off Opol, 214 fms., Mindanao, Sta. 5502; Balayan Bay, 159 fms., Luzon, Sta. 5118 (Albatross Exped., USNM).

Gemmula congener subspecies mekranica (Vredenburg, 1925)

Range — Post-Eocene of India and Miocene of Java and Sumatra.

Remarks — Despite Vredenburg’s detailed de- scription, which occupies 4/2 pages, there appears to be no obvious differences in his subspecies that are not covered by the normal limits of variation admissible for the Recent species. In his own sum- mary, following the description, Vredenburg (p. 57) states that “Compared with Pleurotoma con- gener this fossil is distinguished by its somewhat smaller dimensions,” i.e., as follows: height, 27; width, 10; height of spire, 15; height of body whorl, 16 mm.

Vredenburg (p. 57) then states: “Judging from the figures and descriptions, no essential difference can be discovered between this fossil (i.e. mekranica ) and a shell from the Upper Miocene of Java and Sumatra which Martin has described as coronifer(a) ( a name pre-employed by Bellardi in 1877 for a species from the Miocene of Piedmont).”

Vredenburg’s figures of his mekranica are small and too indistinct for accurate comparison with the living congener, so the subspecies is retained ten- tatively until comparisons of material can be made.

Incidentally, also on page 57, Vredenburg vir- tually introduces another new name with the fol- lowing statement “Compared with Mekran speci- mens, the Gaj specimens are apparently of a still smaller size which, if really characteristic of their horizon, might serve to distinguish them as a vari- ety ‘ gajensis ”

Synonymy —

1925 Pleurotoma (Gemmula) congener mekranica Vreden- burg, Mem. Geol. Surv. India 50(1), p. 54.

1925 Pleurotoma (Gemmula) congener gajensis Vredenburg, Mem. Geol. Surv. India 50(l), p. 57 (nomen nudum ) .

Records— N.W. INDIA: Gaj of Kachh (Mekran beds, post-Eocene ) ; also west of Gharh Hill and between Kanderi and Sari Dasht in Kulanch. JAVA and SUMATRA: (Up- per Miocene) (Vredenburg).

Gemmula congener subspecies cosmoi (Sykes, 1930)

(PI. 192)

Range — Japan, 50 to 100 fathoms.

Remarks — This shell resembles kieneri but is always separable by its loose coiling, or more rap- idly increasing whorls, the single subsutural cord, the form of the peripheral gemmules which are not only fused in vertical pairs but extend “comma”- like below the carina, and the prominence of three or four of the basal spirals, as well as the simplicity of the coloration which is white, except for a single spiral light-brown line on the subsutural cord.

The subspecies diomeclea, described following, indicates the relationship of cosmoi to be with con- gener rather than with kieneri.

Plate 192. Gemmula congener cosmoi (Sykes). 50 fms., off Tosa, Japan. 63 mm.

[22 - 714]

March 31, 1964

INDO-PACIFIC MOLLUSCA, vol. 1, no. 5

Gemmula 253

Plate 193. Geographical distribution of Gemmula congener and its subspecies, and Gemmula amabilis (Weinkauff), G. hawletji Iredale and G. dampierana new species.

Type Locality — Kii, Japan.

Records — JAPAN: Tosa and Kii (ANSP); off Tosa, 50 fms. (USNM); off Tosa, 100 fms. (A. W. B. Powell coll.); Sagami Bay (Thaanum coll.).

Description — Adult shell 52 to 67 mm. ( 2 to 2M inches) in height, elongate-fusiform, with a tall spire and long tapered anterior canal. Adult whorls 9, strongly angled by a broad square-faced periph- eral carina situated just below the middle of whorl height. This carina is densely studded with laterally compressed cog-like axials which extend a little distance below the carina in comma-like fashion. There is a strong but narrow crested subsutural fold and one, sometimes two, smooth primary cords between the carina and the lower suture. Shoulder with three to four fine crisp secondary spiral threads. Body whorl with three or four spiral cords much stronger than the rest, situated on the upper part of the base. Below this the spirals be- come weaker and rather crowded over the neck and canal. Colour as described above, pure white, except for a subsutural light-brown line. The whole surface crossed by rather dense but weak axial growth lines which tend to crenulate the primary basal spirals.

Measurements (mm.) —

height width

67.0 21.5

64.0 20.7

63.0 18.7

52.0 17.0

46.5 12.0 holotype

Synonymy —

1930 Tunis cosmoi Sykes, Proe. Malac. Soc. 19, p. 82, text fig.

1954 Gemmula cosmoi Sykes, Kira, Coloured Illust. Shells of Japan, pi. 35, fig. 13.

Gemmula congener subspecies diomedea new subspecies (PI. 191, figs. 5, 6)

Range — Philippines, 100 to 350 fathoms.

Remarks — This subspecies forms a link between the delicately sculptured Japanese cosmoi and the robust, heavily carinated Indian Ocean congener.

So far diomedea is known only from deep water off the Philippines. It resembles congener in having a bicarinate subsutural fold but the wide shallow shoulder area and the four extra strong basal spirals are characters more in accord with cosmoi. The subspecific name is derived from Diomedea, the genus for the wandering albatross.

In a number of instances this subspecies exhibits heavy folds and flutings at the termination of the basal primary cords and these strongly laciniate the outer lip margin.

Description — Adult shell 63 to 88 mm. ( 2/2 to 3/2 inches) in height, elongate-fusiform, with a tall spire and long considerably flexed anterior canal. Adule whorls 9, strongly angled at about middle whorl height by a broad square-cut peripheral carina which is densely studded with vertically fused gemmules. Subsutural fold composed of two closely spaced gemmate cords. Threads, 3 to 5, on the deeply concave shoulder, 2 primary cords between the peripheral carina and the lower suture with 2 to 3 weak threads in the interspaces. Body whorl including neck and anterior canal with numerous primary cords and intermediate threads.

[22 - 715]

254 Gemmula

A. W. B. Powell

Turridae

Four spiral cords on the upper part of the base much stronger than the rest and in some examples these are developed into heavy meandering and twisted folds that laciniate the margin of the outer lip. Spiral sculpture of dense, strong, axial threads which render all the cords gemmate or crenulate. Colour white except for a subsutural band of light- brown which covers both subsutural cords.

Measurements (mm.) —

height	width
88.0	27.0	Malocot Pt., Luzon Id., 198 fms.
73.0	20.5	holotype
63.3	18.2	paratype
53.6	16.5	off Panglao Id., 220 fms.

T ypes — The type locality is Albatross Station 5397, 198 fathoms, off Malocot, West Luzon Island, Philippines. The holotype is in the U. S. National Museum, no. 238878.

Records — PHILIPPINES: S.E. of Pt. Tanon, Cebu Id., 310 fms. (Albatross Sta. 5335); N.W. of Panglao Id., 220 fms. (Albatross Sta. 5198); off Dupon Bay, Leyte Id., 350 fms., green mud (Albatross Sta. 5407); off Apo Id., Negros Id., 256 fms. (Albatross Sta. 5538); off Malocot Pt., West Luzon Id., 198 fms., mud and sand (Albatross Sta. 5397); West of Siquijor Id., 254 fms., green mud (Albatross Sta. 5369); off Tayabas Lt., Luzon Id., 106 fms., black sand (Albatross Sta. 5369); off Tayabas Bay, Luzon Id., 190 fms., green mud (Albatross Sta. 5374); off Opol, Mindanao Id., 214 fms. ( Albatross Sta. 5501 ) ; off Pt. Sella, Mindanao Id., 219 fms. (Albatross Sta. 5541); off Abgao, Leyte Id., 182 fms., green mud (Albatross Sta. 5403); also the follow- ing Albatross Philippine stations - 5118, 5404, 5410, 5508, 5535 and 5589 (USNM).

Plate 194. Figs. 1, 2, Gemmula miocoronifera new name. Miocene of Java. Formerly Pleurotoma coronifer Martin, 1879, non Bellardi, 1877 (from K. Martin, 1879, pi. 11, figs. 2, 2a). Figs. 3-5, Gemmula imitatrix Martin. Lower or Mid- dle Miocene of Java, Nanggulan Beds. Figs. 3, 4, original types for PL coronifera Martin, 1884 (not 1879). Fig. 5, syntype of imitatrix Martin.

Gemmula miocoronifera new name

(PI. 194, figs. 1, 2)

Range — Miocene of Liotjitjankang, Java Island, Indonesia (type locality).

Remarks — This species is characterised by its tall, narrowly tapered spire and double-submargin- ing of the suture, the top spiral cord being the smaller. It seems more closely related to kieneri (Doumet) than to congener, and is certainly not a synonym of Vredenburg’s mekranica.

The name Pleurotoma coronifer Martin, 1879, must be considered as a homonym of Pleurotoma coronifera Bellardi, 1877, for there is no proper Latin form “coronifer,” only coronifera, a fact which Martin realized in 1884 when he corrected his spelling to coronifera. I hereby rename Martin’s homonym Gemmula miocoronifera new name. Mar- tin did not give any measurements in his original description.

Synonymy —

1879 Pleurotoma coronifer Martin, Tertiarschichten auf Java, p. 61, pi. 2, fig. 2 (non Bellardi, 1877).

1884 Pleurotoma coronifera Martin, Samml. Geolog. Reichs- Mus. Leiden, Ser. 1, 3, p. 58 (in part, synonymy only).

Gemmula imitatrix (Martin, 1916)

(PI. 194, figs. 3-5)

Range — Miocene of Djokdjokarta (Nanggulan), Java Island, Indonesia (type locality).

Remarks — This species is a new name for the same author’s coronifera of 1884, not coronifer, again of the same author, 1879. The species has a wider spire angle and the gemmate keel is less prominent than in the original coronifer.

Measurements (mm.) —

height width

19.0

Synonymy —

1884 Pleurotoma coronifera Martin, Samml. Geol. Reichs- Mus., Leiden, Ser. 1, 3, p. 58, pi. 4, fig. 58 (not of Martin, 1879).

1916 Pleurotoma (Hemipleurotoma) imitatrix Martin, Die altmiocene Fauna des West-Progogebirges auf Java, Samml. Geol. Reichs-Mus., Leiden, 2, p. 229, pi. 1, figs. 13, 13a.

1938 Tunis (Gemmula) imitatrix Martin, Oostingh, Die Moll. Plioc. Sud. Bantam in Java, De Ingen. Ned.- Indie, Gast. 1, p. 28.

[22 - 716]

March 31, 1964

A. W. B. Powell

Gemmula 255

Gemmula sindiensis (Vredenburg, 1925)

(PI. 195, figs. 1-3)

Range — Post-Eocene of India.

Remarks — This species, which its author “re- garded in all probability as a premutation of Pleu- rotoma congener,” is in one important respect dis- similar in that the subsutural fold carries only one spiral cord, which allies it instead with cosmoi. The Indian fossil, however, is uniformly more strongly spirally sculptured than the Japanese Recent spe- cies, yet does not show a selective development of the three or four upper base spirals so charac- teristic of cosmoi.

Measurements (mm.) —

height width height of spire height of body whorl 27 10 16 16

Synonymy —

1925 Pleurotoma (Gemmula) sindiensis Vredenburg, Mem. Geol. Surv. India 50(1), p. 58, pi. 5, figs. 13, 14.

Records — N.W. INDIA (post-Eocene): Gaj of Kachh, near Warsar, north of lakao.

Plate 195. Figs. 1-3, Gemmula sindiensis (Vredenburg). Gaj of Kachh, India. Post-Eocene. Figs. 4, 5, Gemmula pul- chella Shuto. Lower Pliocene, Takanabe member, fapan ( both holotypes from the original figures ) .

Gemmula pulchella Shuto, 1961

(PI. 195, figs. 4, 5)

Range — Lower Pliocene of Japan.

Remarks — This shell was described as a sub- species of granosa Helbling (i.e. kieneri Doumet) but from figures and the description the alliance seems to be with cosmoi Sykes. It is described as having a very small protoconch, 22 gemmules on the penultimate whorl and 29 on the body whorl, and an anal sinus that is broader than the carina, but Shuto s figure ( text fig. 4, no. 3 ) shows a sinus restricted to, if not narrower than, the carina. In cosmoi the gemmules range from 25 to 32 and in kieneri, 30 to 35 per whorl.

Although the type of pulchella is a small shell, only 10.2 mm. in height, it has six adult whorls and is evidently approaching full size. It can be assessed as a forerunner of cosmoi characterised by a dwarf size and fewer, stronger gemmules.

Measurements (mm.) —

height width

10.2 5.6

Synonymy —

1961 Gemmula granosa pulchella Shuto, Mem. Fac. Sci. Kyushu Univ. Ser. D, Geol. 11(2), p. 80, pi. 10, figs. 1, 2; text figs. 3, 4.

Records — IAPAN (middle part of Takanabe member, Lower Pliocene ) : roadside small cliff at Nihonmatsu, Taka- nabe machi, Koyu gun, Miyazaki Prefecture.

[22 - 717]

256 Gemmula

A. W. B. Powell

Turridae

[These occasional blank areas occur between genera and subgenera to permit the insertion of new material and future sections in their proper systematic sequence.]

[22 - 718]

March 31, 1964

INDO-PACIFIC MOLLUSCA, vol. 1, no. 5

Gemmula 257

The Martini Series

Synonymy

Gemmula martini (Tesch, 1915)

(PI. 196, figs. 1-4)

Range — Pliocene of Timor; Recent, off East Africa and off Borneo in deep water.

Remarks — This species along with aethiopica Thiele, sibogae Schepman and sibukoensis form a group characterised by the peripheral carina set low on the spire whorls, almost at the lower suture, which results in a rather flat-sided profile to each whorl in the spire.

Description — Adult shell 46 to 77 mm. ( about 2 to 3 inches) in height, solid, broadly fusiform, uniformly dull-white, without colour markings. Adult whorls 11, outlines straight except for a slightly projecting, broadly rounded sinus-rib set just above the lower suture. Sculpture simple, of smooth narrowly rounded but relatively strong spiral cords. Above the peripheral carina the inter- spaces are without interstitial spirals but below the carina, over the base, neck and canal, all the cords have one or two spiral threads between them. The peripheral carina is composed of 2 or 3 closely spaced spiral cords which are densely studded with vertically and partially fused gemmules, 38 to 50 per whorl. Spire equal to height of aperture plus canal, angle 35° to 37°.

Measurements (mm.) —

height	width
77.0	25.0	East Africa, 638 metres ( type of valdiviae )
68.0	22.5	Sibuko Bay, Borneo, 310 fms.
49.0	16.0	Timor, Noil Aintie (Pliocene)
48.0	16.5	type of martini (fig. 48b of Tesch)

1903 Pleurotoma (Gemmula) carinata Gray, Martens, Gast. Deutsch. Tiefsee-Exped., p. 76 (non Griffith & Pidgeon, 1834).

1915 Pleurotoma ( s.str. ) martini Tesch, Palaeont. von Timor, p. 26, pi. 77(5), figs. 48, 49.

1925 Pleurotoma (Gemmula) valdiviae Thiele, Wissenschaft. Ergebn. Deutschen Tiefsee-Exped., 17, Gastr. 2, p. 208, pi. 23, fig. 1.

Types — The type locality for martini (Tesch) is in a Pliocene deposit, station W III, Noil Noni, be- tween Kapan and Niki-Niki, Timor, Indonesia.

Records — TIMOR: (Pliocene) Noil Noni (type locality of martini ); Noil Aintie, Kolo, Pliocene (Brit. Mus. ). EAST AFRICA: off Somaliland, 0° 27.4' S., 42° 47.3' E., 638 metres (type locality of valdiviae)- off Somaliland, 977 and 1134 metres (Thiele, 1925). SUMATRA: off north coast, 6° 56.3' N., 93° 32.7' E., 362 metres; off Nias Id., 470, 614 and 660 metres (Thiele, 1925). BORNEO: S.E. of Mabul Id., Sibuko Bay, grey mud and sand ( Albatross Sta. 5590, USNM).

Gemmula aethiopica (Thiele, 1925)

(PI. 196, fig. 5)

Range — Off Somalia, East Africa, 638 metres.

Remarks — From its figure and description this species appears to be closely allied to martini Tesch but has sufficient differences to warrant specific separation from that species. Compared with mar- tini, the shoulder is not so straight in profile but is inwardly subangled medially by a groove that delimits a broad but flat subsutural collar. The gemmules are fused into oval shapes taller than wide and set across the tricordate peripheral ca- rina. These gemmules are much stronger but less numerous than in martini. None of the spiral cords, not even those on the base, appear to have inter- mediate spirals.

Plate 196

Figs. 1-3 Gemmula martini (Tesch). Pliocene of Timor.

Tesch ’s original figures, pi. 77.

4 Gemmula martini (Tesch). Holotype of Pleuro-

toma (Gemmula) valdiviae (Thiele), off East Africa (from Thiele, 1925, pi. 23, fig. 1).

5 Gemmula aethiopica (Thiele). Holotype. 638

meters, off East Africa (from Thiele, 1925, pi.

34, fig. 25).

6, 7 Gemmula sibogae Schepman. Holotype. 472

meters, between Makjan and Halmahera Ids., Indonesia (from Schepman, 1913, pi. 26, fig. 2).

8, 9 Gemmula sibukoensis new species. 310 fms., S.E. of Mabul Id., Sibuko Bay, Borneo. Fig. 8 is holotype, 46.2 mm.

10 Gemmula vagata (E. A. Smith). Holotype. 200- 350 fms., off Trincomalee, Ceylon (from E. A. Smith, 1895, pi. 1, fig. 3).

[22 - 729]

258 Gemmula

A. W. B. Powell

Turridae

Measurements (mm.) —

height width

47 19

Synonymy —

1925 Pleurotoma (Gemmula) aethiopica Thiele, Wissen- schaft. Ergebn. Deutschen Tiefsee-Expech, 17, Gastr. 2, p. 208(174), pi. 34(22), fig. 25.

T ypes — The type locality is off East Africa, Valdivia Sta. 253, 0° 27.4' S„ 42° 47.3' E„ 638 metres, off southern Somalia.

Gemmula sibogae (Schepman, 1913)

(PI. 196, figs. 6, 7)

Range — Indonesia, between Makjan and Hal- mahera, 472 metres.

Remarks — Compared with aethiopica, the spire angle is much less, the peripheral gemmules appear to be stouter and rather fewer per whorl, and the base more suddenly contracted. The coloration is described as whitish, faintly yellowish on the keel, between the nodules. The spire angle judged from figures is 42° in aethiopica but only 33° in sibogae.

Measurements (mm.) —

height width

27.5 11.5

Synonymy —

1913 Pleurotoma (Gemmula) sibogae Schepman, Siboga Exped., Pt. 5, 49e, p. 404, pi. 26, fig. 2.

Types — The type locality is Indonesia, Siboga Sta. 137, channel between Makjan and Halmahera, fine dark muddy sand, 472 metres.

Gemmula sibukoensis new species (PI. 196, figs. 8, 9)

Range — Borneo, Moluccas and Philippines, 50 to 484 fathoms.

Remarks — This is another member of the group with the peripheral carina set low down on the spire whorls. It appears nearest to sibogae Schep- man, but differs in having a heavier peripheral ca- rina a little above the lower suture, allowing the emergence of the uppermost basal spiral over the later whorls, and the shape of the body whorl, which is but slowly contracted over the base to a shorter and flexed, not straight canal.

Description — Adult shell 41 to 48 mm. (about 2 inches) in height. Fusiform, with a tall spire, about 1 1/5 times height of aperture, plus canal. Whorls 10, plus a small polygyrate conic proto- conch of 3 whorls or more, the last, at least, axially costate. Protoconch eroded and the tip missing in

all available specimens. Spire whorls turreted by a strong rather broadly rounded peripheral sinus- rib, which is set low on the whorls, almost at the lower suture, but with a narrow space beneath it, which allows the emergence, over the last few whorls, of the uppermost basal spiral. Peripheral sinus-rib densely sculptured with laterally com- pressed, concavely arcuate axials, 27 to 31 per whorl, overridden by 2, occasionally 3, weak spirals. Spiral sculpture consisting of rather strong but narrow cords, 3 to 5 on the wide, steeply descend- ing shoulder and 16 to 18 on the base; 3 on the upper part of the base rather stronger than the rest, which diminish in size gradually to the end of the anterior canal. Labial sinus of moderate depth, V to U-shaped, its apex wider than the crest of the sinus-rib. Colour creamy white. Surface crossed by dense sharp axial growth lines.

Measurements (mm.) — height width

47.8 16.2 figured paratype

46.2 16.2 holotype

T ypes — The type locality is Albatross station 5590, S.E. of Mabul Island, Sibuko Bay, Borneo, in 310 fathoms. The holotype is in the United States National Museum, no. 239111.

Records — BORNEO: S.E. of Mabul Id., Sibuko Bay, 310 fms. (holotype) (Albatross Sta. 5590); off Mabul Id., Sibuko Bay, 260 fms., sandy mud (Albatross Sta. 5589); off Silungan Id., Sibuko Bay, 305 fms., green mud and sand (Albatross Sta. 5592). CELEBES: Gulf of Boni, 484 fms., grey mud (Albatross Sta. 5656); MOLUCCAS: off Kayoa Id., 265 fms., grey mud and fine sand (Albatross Sta. 5626); off Makyan Id., Molucca Pass, 288 fms., fine sandy mud (Albatross Sta. 5624). PHILIPPINES: off Liangan River, Mindanao Id., 410 fms., grey mud and sand (Alba- tross Sta. 5511); N. of Biliran Id., 50 fms., mud and sand (Albatross Sta. 5210) (all USNM).

Gemmula vagata (E. A. Smith, 1895)

(PI. 196, fig. 10)

Range — Indian Ocean from Aden to the Anda- man Islands, in deep water.

Remarks — This very distinctive species is at once recognised by its almost vertical-sided whorls be- tween a strongly projecting flange-like peripheral keel.

Description — Adult shell 60 to 65 mm. ( about 2 inches ) in height, rather heavily built and strongly sculptured. Spire tall and narrow with rather straight and almost vertical sides, interrupted just below the middle by a prominently projecting, heavy, flange-like, gemmate keel. Spire 1.2 to 1.3 times height of aperture plus anterior canal. The sculpture of the spire whorls consists of a moder-

[22 - 730]

March 31, 1964

INDO-PACIFIC MOLLUSCA, vol. 1, no. 5

Gemmula 259

Plate 197. Geographical distribution of Gemmula aethio- pica (Thiele), G. martini (Tesch), G. sibogae (Schepman) and G. sibukoensis new species.

ately strong subsutural square-cut fold, bearing 2 spiral cords, with 2 or 3 fine spiral threads between them, followed by a deeply concave shoulder, bear- ing 5 to 9 weak spiral threads between the sub- sutural fold and the strongly gemmate cog-like peripheral sinus-rib. About 24 gemmules on the penultimate. Below the peripheral sinus-rib, 2 of the basal series of plain narrow spirals are emer- gent. About 16 to 18 basal spirals present, those above wide-spaced and with several fine threads in each interspace. Colour (Smith, 1904) “The infra- sutural keel is generally somewhat reddish and the central carina is spotted with the same colour be- tween the tubercles.”

The species seems to be nearest allied to gem- mulina Martens, a smaller and less solid shell from the Archibenthal of Indonesia and the South China Sea.

Measurements (mm.) —

height	width
65.0	23.0	Ceylon, 200 to 350 fms. (type)
61.0	18.0	Chagos Archipelago, 494 metres
53.0	17.0	Chagos Archipelago, 494 metres
46.0	15.0	Gulf of Aden, 1022 metres

Synonymy —

1895 Pleurotoma vagata E. A. Smith, Ann. Mag. Nat. Hist., Ser. 6, 16, p. 3, pi. 1, fig. 3; 1904, E. A. Smith, Ann. Mag. Nat. Hist., Ser. 7, 13, p. 456.

1909 Pleurotoma vagata Smith, Annandale & Stewart, Illustr. Zool. Investigator, Calcutta, Moll., 6, pi. 14, figs. 3, 3a.

Types — Indian Museum, Calcutta.

Gemmula praesignis (E. A. Smith, 1895)

(PI. 198, fig. 1)

Range — Off Colombo, Ceylon, 675 fathoms.

Remarks — Smith’s species, based presumably upon a single imperfect shell minus the protoconch as well as the tip of the anterior canal, does not look very different from that author’s vagata , de- scribed in the same paper from approximately the same area; deep water off Ceylon. From the figure, praesignis would appear to differ from vagata in having a heavier peripheral carina with more prominent and less numerous nodules, a very strong sharply crested subsutural spiral and fewer sec- ondary spirals in the shoulder concavity.

Description (E. A. Smith)— “The prominent row of tubercles around the middle of the whorls, the keel beneath the suture, and the broad sinus in the labium are the principal features of this species. The apex of the spire being broken away makes it impossible to state with certainty the exact number of whorls, but they would probably amount to eleven or twelve. The entire surface exhibits fine flexuous lines of growth.”

Measurements (mm.) — height width

42.0 15.0 holotype

Synonymy —

1895 Pleurotoma praesignis E. A. Smith, Ann Mag. Nat.

Hist., Ser. 6, 16, p. 4, pi. 1, fig. 4.

Types — Indian Museum, Calcutta.

Records — CEYLON : off Colombo, 675 fms. Schepman, 1913, Siboga Exped., Moll., p. 39, recorded this species from the Flores Sea in 794 metres and Halmahera Sea in 411 metres, but both records are suspect since they were based upon “very young” specimens.

Records — CEYLON : off Trincomalee, 200-350 fms. (type locality). INDIA: off Travancore coast, 360 fms. (Smith, 1904); off Malabar coast, 295-360 fms. (USNM); GULF OF ADEN: 732 metres, 1022 metres and 1061 metres. 4° 58' 42” S., 73° 16' 24" E., north of the CHAGOS AR- CHIPELAGO, 494 metres ( John Murray Exped. ) ( Brit. Mus.). ANDAMAN ISLANDS: 185 fms. (Smith, 1904).

Gemmula ducalis (Thiele, 1925)

(PI. 198, fig. 2)

Range — Off East Africa, 1134 metres.

Remarks — This juvenile shell is only 6 mm. in

[22 - 7311

260 Gemmula

INDO-PACIFIC MOLLUSCA, vol. 1, no. 5

Turridae

Plate 198. Fig. 1, Gemmula praesignis (E. A. Smith). Holotype. 675 fms., off Colombo, Ceylon (from E. A. Smith, 1895, pi. 1, fig. 4). Fig. 2, Gemmula ducalis (Thiele). Holo- type. 1,134 meters, off Somaliland, East Africa (from Thiele, 1925, pi. 23, fig. 2). Fig. 3, Gemmula rotatilis (Martens). Holotype. 1,134 meters, off Somaliland, East Africa (from Martens, 1903, pi. 1, fig. 3).

height, and consists of a typical multispiral, axially costate protoconch and three post-nuclear whorls. It is impossible to say at this stage of our knowl- edge of the genus if this is really a distinct species or merely a juvenile of one already described.

Synonymy —

1925 Pleurotoma (Gemmula) ducalis Thiele, Wissenschaft.

Ergebn. Deutschen Tiefsee-Exped., 17, Gastr. 2, p.

209, pi. 23, fig. 2.

Records — Off Somaliland, East Africa, 1° 49' N., 45° 29.5' E., 1134 metres.

Gemmula gemmulina (Martens, 1902)

(PI. 200, fig. 2; pi. 201, figs. 1, 2)

Range — Sumatra to China Sea and the Philip- pines, 35 to 505 fathoms.

Remarks — The Albatross Expedition material re- corded below is claimed with some confidence to represent von Marten’s species, the type of which came from off the west coast of Sumatra in 677 metres.

Nothing quite matching gemmulina in sculptural detail was found although the Albatross material exhibits a considerable range of sculptural varia- tion. It is possible, of course, that von Marten’s fig- ure is over-simplified so far as secondary sculpture is concerned.

A characteristic feature, however, is common to both von Marten’s figures and the Albatross speci- mens and that is the strong crisp nature of the bi- carinate subsutural fold.

The discrepancy noted above concerns the sub- sidiary spirals on the concave shoulder between the subsutural fold and the peripheral carina. In von Marten’s figure only one, quite distinct, spiral appears in the shoulder concavity but in the Al- batross material these spirals number from 3 to 5,

of which never more than 3 are relatively strong. Since there is such variation in a series of 7 Alba- tross specimens from 5 localities it is likely that a wider series would include such an example as Marten’s type within the range of variation admis- sible for the species.

The species shows relationship with vagata (Smith), which differs in having almost vertical- sided whorls, a longer canal, a weaker bicarinate subsutural fold and weaker secondary sculpture, as well as attaining a larger adult size.

Description (based upon the figured Molucca Passage shell) — Adult shell 29 mm. ( 1 1/8 inches ) in height, solid, white, covered by a thin shining light creamy-buff periostracum. Pagodiform, with a tall spire and relatively short anterior canal. Spire about 1.3 times height of aperture plus canal. Adult whorls 10 (protoconch worn in all specimens but indicated as tall, polygyrate and axially costate). Sculptured with two closely spaced crisp spiral cords forming the subsutural fold, a prominent gemmate bicarinate peripheral keel below the mid- dle of whorl height and subsidiary spiral cords, 3 between the subsutural fold and the peripheral keel and 2 to 3 below it. Base with about 17 sub- equal primary cords and a single weak thread in each interspace over the region of the neck. The peripheral gemmules are vertically fused into lat- erally compressed nodes giving a cog-wheel effect. The nodes number 20 on the penultimate. Sinus openly U-shaped, not very deep, its apex rather wider than the peripheral keel. The whole surface of the shell crossed by regular thin axial growth threads.

Plate 199. Geographical distribution of Gemmula vagata (E. A. Smith) and G. gemmulina (Martens).

[22 - 732]

March 31, 1964

INDO-PACIFIC MOLLUSCA, vol. 1, no. 5

Gemmula 261

An extreme variant from 90 fathoms off Luzon Island has a very prominent bicarinate subsutural fold and 5 spiral cords in the shoulder concavity.

Measurements (mm.) —

height	width
37.5	12.0	Pratas Id., China Sea, 122 fms.
35.7	11.7	Luzon Id., 90 fms.
29.0	10.2	Molucca Passage, 275 fms.
27.0	9.2	Molucca Passage, 230 fms.
23.2	8.0	Molucca Passage, 230 fms.
20.5	6.3	type

Synonymy —

1902 Fleurotoma (Gemmula) gemmulina Martens, Sitzungsb. d. Gesell. naturf. Freunde, Berlin, p. 238; 1903 Martens, Gast. der Deutsch. Tiefsee-Exped., 1898- 1899, 7, p. 77, pi. 1, fig. 2.

Records — SUMATRA: off west coast, 677 metres (type); off Siberut Id., 750 metres (von Martens). MOLUCCAS: off Makyan Id., Molucca Passage, 275 fms. grey mud, Sta. 5622; off Gillolo Id., Molucca Passage, 230 fms., Sta. 5625. CELEBES: S.E. of Tikola Peninsula, Buton Strait, 37 fms., grey mud, Sta. 5642. BORNEO: S. of Silungan Id., Sibuko Bay, 305 fms., Sta. 5592. CHINA SEA: off Pratas Id., 122 fms. PHILIPPINES: off Cape Santiago, Luzon Id., 280 fms., Sta. 5283; off Pitogo, Luzon Id., 90 fms., grey mud and sand, Sta. 5376; N. of Ambil Id., 102 fms., fine sand and mud, Sta. 5278; Iligan Bay, N. Mindanao Id., 505 fms, grey mud and fine sand, Sta. 5513 (all Albatross stations, USNM).

Plate 200. Fig. 1, Gemmula amabilis (Weinkauff). 732 meters, in the Gulf of Aden. 18 mm. Fig. 2, Gemmula gem- mulina Martens. 275 fms., off Makyan Id., Molucca Passage, Albatross station 5622. 29 mm. Fig. 3, Gemmula graeffei (Weinkauff). Momi Bay, south of Nabilo Light, Fiji. 22 mm. Fig. 4, Gemmula hombroni Hedley. Cebu id., Philip- pines, 26 mm.

Gemmula amabilis (Weinkauff, 1875)

(PI. 200, fig. 1; pi. 201, figs. 3-7)

Range — Red Sea.

Remarks — So far as can be judged from Wein- kauff’s very sketchy figures of his species and the more detailed figures of Sturany, 1903, presumably the same species, the deep-water John Murray Ex- pedition series of specimens from 732 metres off Aden may at present be considered conspecific. The John Murray specimens are more elongate and when compared with authentic amabilis material, may prove to be a bathymetric form of that species, which was probably from shallow water.

Description (Weinkauff, 1875, p. 29) — “Testa fusiformis, solidiuscula, pallide flavefusca, spiraliter cingulata, interstitiis sublente striis incrementi sculpta, cingula una valida, distincte nodosa, cari- nam distinctam efflciens, noduli albide picti: spira elato-conica, apice acuto, anfractibus 11, unicari- natis, sutura evanescente obliqua divisi, ultimus convexus, canali longo productus; apertura verti- calis piriformis, intus sublente costata, margine ex- terno basi producto (Jickeli). Long 18 diam. maj. 6, apert. 4 mm.’’ (See our pi. 201, figs. 3-5).

Description (based upon Aden, 732 metres ex- ample) — Adult shell 20 to 26 mm. (% to 1 inch) in height, elongate-fusiform with a tall spire whose angle is 26° to 27°, and with a moderately long- flexed anterior canal. Whorls 9, plus a tall narrowly conic protoconch of 3/2 whorls, first 2 whorls smooth, remainder strongly axially costate. Post- nuclear whorls sculptured with a relatively strong tricordate subsutural fold, the central cord strong- est and forming a sharp keel. Shoulder concavity with three crisp spiral threads, followed by a prom- inent bicarinate gemmate flange which is located a little below the middle of the whorl. Gemmules vertically fused into laterally compressed nodes, 18 on the penultimate. A single smooth cord and a weak thread are above it between the peripheral carina and the lower suture. Body whorl with three strong smooth widely spaced cords above, below which are 13 finer, closely spaced cords and a sim- ilar weaker number over the anterior end. Sinus a broadly open shallow “U,” its rounded apex occupy- ing the full width of the peripheral carina. Colour uniformly white.

Measurements (mm.) —

height	width
24.0	7.6	off Aden, 732	metres
18.0	6.0	off Aden, 732	metres
18.0	6.0	type, Massana,	, Red Sea

[22 - 733]

262 Gemmula

A. W. B. Powell

Turridae

Synonymy —

1875 Pleurotoma amabilis (Jickeli ms.) Weinkauff, Conch.

Cab. 4(3), p. 29, pi. 6, figs. 4, 6.

1875 Pleurotoma (Gemmula) amabilis (Jickeli ms.) Wein- kauff, Jahrb. Mai. Ges. 2, p. 291, pi. 9, figs. 6, 8. 1903 Pleurotoma ( Gemmula ) amabilis Weinkauff, Sturany, Gast. des Rothen Meeres, Exped. S. M. Schiff “Pola,” pi. 3, fig. 3 a-c.

Records — RED SEA: Massana (type locality); Gulf of Aden, 732 metres (John Murray Exped. Sta. 176, Brit. Mus. ).

Gemmula hombroni (Hedley, 1922)

(PI. 200, fig. 4; pi. 201, fig. 8)

Range — Andaman Islands and Indonesia to the Philippines, and Japan to northern Australia.

Remarks — This small, slender, uniformly reddish- brown shell is one of the most distinctive of the genus. It has long been recognised under the pre- occupied name of fusca Hombron & Jacquinot, but some confusion has occurred through Tryon’s (1884) action in erroneously uniting it with gem- mata Hinds, the Lower California-Panamic type of the genus.

Plate 201. Figs. 1, 2, Gemmula gemmulina (Martens) (also see pi. 200, fig. 2), 677 meters, off the west coast of Sumatra (from Martens, 1902, pi. 2, figs. 2, 2a). Figs. 3-5, Gemmula amabilis (Weinkauff), Massana, Red Sea (from Weinkauff, 1875, pi. 9, figs. 6, 8). Figs. 6, 7, G. amabilis (Weinkauff), Red Sea, (from Sturany, 1903, pi. 3, fig. 3). Fig. 8, Gemmula hombroni (Hedley), Padang, Sumatra ( holotype of Pleurotoma padangensis Thiele, original pi. 23, fig. 5).

Description — Adult shell 25 to 27 mm. (about 1 inch) in height, elongate-fusiform, with a tall slen- der spire and long slightly-flexed anterior canal. Whorls 8 to 8)2, exclusive of a tall, narrowly conic, polygyrate protoconch of 3/2 to 4 whorls, all but the tip sculptured with concavely-arcuate, stout axials. Adult whorls very strongly and definitely sculp- tured, firstly by a heavy, smooth, narrowly crested subsutural cord, followed by the flange-like, gem- mate peripheral sinus-rib, set below the middle of whorl-height, and finally a single heavy, smooth spiral cord, uppermost of the basal series and emer- gent between the peripheral carina and the lower suture. Base with about 9 rather wide-spaced, strong, flat-topped spiral cords to about the middle of the neck, after which the spirals rapidly diminish in size and become more closely spaced. The con- cave shoulder area is encircled by 3 to 5 spiral threads. Spire a little taller than height of aperture plus canal, angle 22° to 25°. Gemmules confined to the peripheral carina, 18 to 22 per whorl, smooth, laterally compressed and cog-like. Sinus at periph- ery moderately deep, U-shaped. Colour uniform, yellowish brown to reddish brown.

Measurements (mm.) —

height	width
28.0	8.5	Cavite, Manila Bay, Philippines
27.0	7.5	Cavite, Manila Bay, Philippines
25.5	7.0	Cavite, Manila Bay, Philippines
11.5	2.4	type of padangensis

Synonymy —

1853 Pleurotoma fusca Hombron & Jacquinot, Voy. Pole sud., Zook, 4, p. Ill, pi. 25, figs. 19, 20 (non PL fusca C. B. Adams 1845).

1884 Pleurotoma gemmata Hinds, Tryon, Manual of Conch. 6, p. 173 (in part).

1913 Pleurotoma (Gemmula) fusca H. & J., Schepman, Siboga Exped. 49e, p. 402.

1922 Gemmula hombroni H. and J., Hedley, Rec. Aust. Mus. 13(6), p. 218, nom. nov. for PL fusca Hom- bron & Jacquinot, 1853, non C. B. Adams 1845.

1925 Pleurotoma (Gemmula) padangensis Thiele, Wissen- schaft. Ergebn. Deutschen Tiefsee-Exped., 17, Gastr. 2, p. 335, pi. 23, fig. 5.

Records — AUSTRALIA: Torres Strait (type of fusca). PHILIPPINES: Mariveles, Bataan, Luzon Id.; % mi. off Balibatikan, Papapas Bay, Batangas Province, Luzon Id., 4 fms.; 1 mi. E. of Limey, Bataan, Luzon Id., 9-10 fms.; 3 mi. E.N.E. of San Nicolas Shoals Light, Manila Bay, Luzon Id., 10 fms.; /2 mi. off Matabunkay Beach, Batangas Province, Luzon Id., 3-4 fms.; I2 mi. E. of Cabcaben, S.E. Bataan, Luzon Id., 2 fms. ( duPont-Acad. Exped., 1958, ANSP ) ; N. of Corregidor Id., 37 fms. (Albatross Sta. 5109); Opol, Mindanao Id., (USNM); near Cavite, Manila Bay ( MCZ). JAPAN: Yenoshima (USNM). ANDAMAN IS- LANDS: (USNM). SUMATRA: Padang (type of padan- gensis). Schepman (1913) records the species from 20 Indonesian stations, 9-522 metres, and off western New Guinea, 32 metres.

[22 - 734]

March 31, 1964

INDO-PACIFIC MOLLUSCA, vol. 1, no. 5

Gemmula 263

Plate 202. Geographical distribution of Gemmula hom- hroni (Hedley) in solid dots, and Gemmula graeffei (Wein- kauff) in open circles.

Gemmula graeffei (Weinkauff, 1875)

(PI. 200, fig. 3)

Range — Fiji, Queensland and Philippines.

Remarks — This species stands nearest to monili- fera, from which it differs in the spire’s height being approximately equal to that of the aperture plus canal, the more slowly contracted base, heav- ier tricarinate subsutural fold and absence of a zoned colour pattern.

Description (based upon a topotypic specimen) — Adult shell 19 to 22 mm. (about % inch) in height, elongate-fusiform with a tall spire and long, slightly flexed anterior canal. Whorls 8, exclusive of a tall, conical, polygyrate protoconch of 3 to 3/2 axially costate whorls. Spire equal to height of aperture plus canal; angle about 30°. Spire whorls with a massive subsutural fold crossed by 3 spiral cords, the central one forming a narrowly crested keel. A moderately deep concavity bearing 2 spiral threads separates the subsutural fold from the pe- ripheral keel which is flange-like and composed of 2 strong gemmate cords. The gemmules are fused vertically into laterally compressed nodules. These nodules, which are cog-like, number 25 on the penultimate whorl. A single strong spiral cord be- tween the peripheral carina and the lower suture with a second cord subemergent toward the termi- nation of the last whorl. Base long and very slowly contracted, crossed above by 4 strong smooth spiral cords, the second of which is in line with or a little above the top of the aperture. Wider-spaced pri- mary cords over the neck, with 2 threads in each interspace and about 16 gradually reduced threads over the anterior end. Entire surface between the

spirals with dense, crisp axial threads. Sinus deep, narrow, U-shaped, its apex occupying the full width of the peripheral carina. Colour uniformly light- buff.

Measurements (mm.) —

height	width
22.0	7.0	Momi Bay, Fiji
21.0	7.0	Pagapas Bay, Philippines
20.0	6.0	type
19.5 Synonymy	5.0	Pagapas Bay, Philippines

1875 Pleurotoma (Gemmula) graeffei Weinkauff, Conch. Cab. 4(3), p. 71, pi. 3, figs. 9, 10; 1875, Jahrb. deut. malak. Ges. 2, p. 290, pi. 9, figs. 9, 10.

1884 Pleurotoma (Gemmula) graeffei Weinkauff, Tryon, Manual of Conch. 6, p. 173.

Records — Fl]l: (type locality); Momi Bay, S. of Nabilo Light, 13-15 fms. (W. Jennings). QUEENSLAND: Low Isles, off Port Douglas; Cairns Reef, 5-8 fms. ( Austr. Mus. ) . PHILIPPINES: /2 mi. off Balibatikan, Pagapas Bay, Batangas Province, Luzon Id., 4 fms.; 1 mi. E. of San Nicolas Shoals Light, Manila Bay, Luzon Id., 10 fms.; E. side Jagoliao Id., N.W. end Bohol Id., 12-24 ft.; E. end of Sisiman Bay, Ba- taan, Luzon Id., 8 fms.; E. end of Corregidor Id., Manila Bay, 4 fms. ( duPont-Acad. Exped., 1958, ANSP).

The species has been recorded from a wide range of localities but all require personal re-examination.

Gemmula hawleyi (Iredale, 1931)

(PI. 203)

Range — New South Wales, Australia.

Remarks — This is the only known temperate wa- ter occurrence of this otherwise tropical genus. Ire- dale (1931, p. 226) introduced this species as the type of a new genus, Eugemmula, claiming that it differed from the West American type of Gemmula, which was stated to have a longer canal and a different apex. These alleged differences, however, prove to be too slight to warrant even subgeneric recognition.

Plate 203. Gemmula hawleyi (Iredale). Port Jackson, New South Wales, Australia (from Laseron, 1954, pi. 1, figs. 5-7).

[22 - 735]

264 Gemmula

A. W. B. Powell

Turridae

The species hawleyi belongs to the group of mo- nilifera, graeffei and hombroni and is in fact doubt- fully distinct from the first mentioned species.

Too few examples, all dead shells, are known to afford a critical comparison at present, but if a topotype in the writer’s collection proves a constant criterion for the species, then it is distinguished from monilifera by having two cords between the periphery and the lower suture, with a third sub- emergent over the last half whorl. Although all ex- amples known to the writer are dead shells there are traces of a zoned colour pattern typical of that found in monilifera.

Measurements ( mm.) —

height	width
35.0	12.0	type of hawleyi Iredale
21.2 Synonymy —	7.5	Port Jackson, Sydney

1931 Eugemmula hawleyi Iredale, Rec. Aust. Mus. 18(4), p. 226, pi. 25, figs. 11, 14.

1954 Eugemmula hawleyi Iredale, Laseron, The N. S. W. Turridae, Handb. Roy. Zool. Soc. N.S.W., p. 7, pi. 1, figs. 5-7.

Type — Australian Museum, Sydney.

Records — NEW SOUTH WALES: Sow and Pigs Reef, Port Jackson (“Triton” dredgings, type locality); Crowdy Head, dredged; off Port Stephens ( Laseron, 1954).

Gemmula rotatilis (Martens, 1902)

(PI. 198, fig. 3)

Range — Off Somalia, East Africa, 1134 metres.

Remarks — This shell was described from a not fully mature shell as evidenced by the size of the protoconch in relation to the rest of the shell. The protoconch is polygyrate, narrowly conic, and closely and radially ribbed. The general appear- ance of the shell suggests alliance with the Dunt- roonian. Upper Oligocene longwoodensis Powell, 1942, from Southland, New Zealand. The East Afri- can shell, compared with the New Zealand fossil, has larger, peripheral, pointed nodes on the sinus area; and the single subsutural spiral cord is stronger, narrowly crested and more prominently raised. Both species are strongly angulated at the periphery but are not carinate-flanged.

Measurements (mm.) — height width

11.5 5.0 from von Martens

Synonymy —

1902 Pleurotoma (Gemmula) rotatilis Martens, Sitzungber. der Gesellsch. nat. Freunde, Berlin, p. 239; 1903, Gast. Deutsch. Tiefsee-Exped., 1898-1899, 7, p. 78, pi. 1, fig. 3.

Types — The type locality is off Mogadiscio, So- malia, East Africa, 1° 49' N„ 45° 29' E., in 1134 metres.

Gemmula iris (Vredenburg, 1921)

(PI. 204, figs. 1, 2)

Range — Burma, Tertiary.

Remarks — This species has a very tall and nar- row spire but a somewhat truncated anterior canal. The whorls are evenly and lightly convex, the gem- mate double sinus rib projecting less than the sub- sutural fold. Protoconch tall and conical, first VA whorls smooth, the remaining 1/2 whorls strongly axially costate. Spire height almost twice the height of aperture plus canal.

Measurements (mm.) —

height width

19.0 6.0

Synonymy —

1921 Pleurotoma (Hemipleurotoma) iris Vredenburg, Rec.

Geol. Surv. India 53(2), p. 98, pi. 12, figs. 14, 15.

Records — BURMA: Kyaungon, Thanga (Tertiary).

Gemmula thyrsus (Vredenburg, 1921)

(PI. 204, fig. 4)

Range — Burma, Tertiary.

Remarks — This narrowly fusiform shell is char- acterised by a strong, sharply crested subsutural fold, a strong gemmate peripheral carina and a prominent smooth cord between the periphery and the lower suture. Spire about 1.1 times height of aperture plus canal.

Measurements (mm.) —

height width

13.3 4.1 estimated from figure

Synonymy —

1921 Pleurotoma (Gemmula) thyrsus Vredenburg, Rec. Geol.

Surv. India 53(2), p. 103, pi. 12, figs. 11.

Records — BURMA: Kyaungon, Myaukmigon, Thanga

(Tertiary).

Gemmula birmanica (Vredenburg, 1921)

(PI. 204, fig. 3)

Range — Burma, Tertiary.

Remarks — The author of this species remarked that it is probably a premutation of Pleurotoma ca- rinata woodwardi Martin, a fossil which occurs abundantly in the Pliocene formations of Java. “The Burmese shell appears to be distinguished prin- cipally by its smaller size, its decidedly smaller pro- toconch, its somewhat better developed circum- sutural rim, and its generally more elongate spire.”

March 31, 1964

INDO-PACIFIC MOLLUSCA, vol. 1, no. 5

Gemmula 265

It has a relatively short canal as in iris but is dis- tinguished from that species in the greater promi- nence of the peripheral carina, which angulates the whorls. Height of spire about 1.3 times height of aperture plus canal.

Measurements (mm.) —

height width

21.0 8.0 estimated from figure

Synonymy —

1921 Pleurotoma (Gemmula) birmanica Vredenburg, Ree.

Geol. Surv. India 53(2), p. 102, pi. 12, fig. 12.

Records — BURMA : Dalabe, Kyaungon, Myaukmigon,

Thanga (Tertiary).

Gemmula pakistanica (Eames, 1952)

(PI. 204, fig. 8)

Range — Eocene of Pakistan, Zinda Pir section (Upper Chocolate Clays).

Remarks — Compared by its author with Pleuro- toma (Gemmula) sindiensis Vredenburg from the Lower Nari of Bhagothoro Hill, Sind, which has a higher spire, Pleurotoma (H emipleurotoma) bonneti bhagothorensis Vredenburg from the same locality and Pleurotoma tricincta Martin, 1935, from the Neogene of Buton, which is more broadly conic.

Measurements (mm.) —

height	width
6.7 +	3.4	holotype
Synonymy —

1952 Tunis (Gemmula) pakistanica Eames, Phil. Trans. Roy.

Soc. London, Ser. B., No. 631, vol. 236, p. 124, pi.

5, fig. 116.

Gemmula soriensis (Eames, 1952)

(PI. 204, fig. 5)

Range — Eocene of Pakistan, Zinda Pir section (Ghazij Shales).

Remarks — Compared by its author with Pleuro- toma (Gemmula) sindiensis Vredenburg from the Lower Nari of Bhagothoro Hill, Sind, and the fore- going species, pakistanica Eames, the former hav- ing a higher spire and more numerous anterior spirals and the latter a more broadly conic spire and narrower, more projecting peripheral carina than in soriensis.

Measurements (mm.) —

height width

7.9 3.0 holotype

Synonymy —

1952 Tunis (Gemmula) soriensis Eames, Phil. Trans. Roy.

Soc., London, Ser. B, No. 631, vol. 236, p. 126, pi.

5, fig. 114.

Gemmula karangensis (Martin, 1895)

(PI. 204, figs. 6, 7)

Range — Lower Miocene, Karang, Java.

Remarks — This species has a bicarinate moder- ately projecting keel, distantly crenulated rather than gemmate. From the small and not very dis- tinct figures the subsidiary spirals appear thin and sharply raised. The surface is crossed by thin axial threads which do not appear to render the sub- sidiary spirals gemmate. The specimen is probably immature. The author compares the species with coronifera Martin and carinata Gray.

Synonymy —

1895 Pleurotoma ( s. str. ) karangensis Martin, Samml. geol.

Reichs-Mus., Leiden, 1, p. 36, pi. 6, figs. 90, 90a.

Gemmula kotorai (Nomura & Zinbo, 1935)

Range — Japan, Yanagawa shell-beds in Hukusima Basin, N.E. Honshu, Miocene.

Plate 204. Figs. 1, 2, Gemmula iris (Vredenburg). Holo- type. Tertiary: Thanga; Kyaungon, Burma (from Vreden- burg, 1921, pi. 12, figs. 14, 15). Fig. 3, Gemmula birmanica (Vredenburg). Holotype. Tertiary: Thanga and Dalabe, Burma (from Vredenburg, 1921, pi. 12, fig. 12). Fig. 4, Gemmula thyrsus (Vredenburg). Holotype. Tertiary: Thanga, Burma (from Verdenburg, 1921, pi. 12, fig. 11). Fig. 5, Gemmula soriensis (Eames). Holotype. Eocene: Ghazig Shales, Zinda Pir section, Pakistan ( from Eames, 1952, pi. 5, fig. 114). Figs. 6, 7, Gemmula karangensis Mar- tin. Lower Miocene: Karang, Java (from Martin, 1895, pi. 6, figs. 90, 90a). Fig. 8, Gemmula pakistanica (Eames). Holotype. Eocene: Zinda Pir section, Pakistan (from Eames, 1952, pi. 5, fig. 116).

[22 - 737]

266 Gemmula

A. W. B. Powell

Turridae

Remarks — It is impossible to tell from the figure what this unique fossil is like. Its authors state that “This species resembles T. ( Gemmula ) granosa (Helbling), a well-known living and fossil species in Japan and the Malayan Archipelago, but the present shell is smaller with fewer revolving cords and less granular sculpture.”

Measurements (mm.) —

height width

9 4 holotype

Synonymy —

1935 Tunis (Gemmula) kotorai Nomura & Zinbo, Saito Ho-on Kai Mus., Res. Bull. No. 6, p. 170, pi. 15, fig. 18.

Gemmula kishimaensis Shuto & Ueda, 1963

Range — Japan, Upper Eocene to Middle Oligo- cene.

Remarks — From the fairly detailed description and figures of this species it would seem to be a true Gemmula, not a member of the subgenus Hemipleurotoma, as claimed by its authors. Cer- tainly it is described as having a paucispiral proto- conch, but this statement is qualified by the remark “but the details are unknown.” However, Glibert (1960, p. 6) considered Hemipleurotoma to be a synonym of Gemmula.

Description — Adult shell 11 to 15 mm. (about /2 inch ) . Shell narrowly fusiform with tall spire and presumed long canal. Median carina heavy, strongly gemmate. Shoulder steeply descending. Subsutural fold broad but not prominent and bearing weak spiral lirations. Four primary spirals below the periphery to the basal subangulation.

Measurements (mm.) —

height width

15.1 9.0

11.5

Synonymy —

1963 Gemmula (Hemipleurotoma) kishimaensis Shuto and

Ueda, Japanese Journal of Geology and Geography 34,

no. 1, p. 4, pi. 1, figs. 6, 11.

Types — The holotype is in the Kyushu Univer- sity, Japan.

Records — JAPAN : Obo, Saga Prefecture, Mazean Stage ( Middle Oligocene ) ( type locality ) .

New Zealand Tertiary Gemmula

The genus Gemmula is not found living in New Zealand waters but it was well represented during

the warmer periods of the Eocene, Oligocene and Miocene and seems finally to have become extinct here in the lower Pliocene. Following is a list of the New Zealand species.

Gemmula bimarginata (Suter, 1917)

Locality — N.Z.G.S. loc. 630, Teaneraki, Enfield, near Oamaru ( probably McCullough’s Bridge, South Canterbury; Bortonian, Eocene). The type is in the New Zealand Geological Survey, Welling- ton.

Synonymy —

1917 Tunis bimarginatus Suter, N. Z. Geol. Surv. Pal. Bull. 5, p. 44, pi. 5, fig. 13.

1942 Gemmula bimarginata Suter, Powell, Bull. 2, Auck. Inst. Mus., p. 49.

Gemmula clifdenensis Powell, 1942

( PI. 205, fig. 1 )

Locality — Clifden (6c.), Southland (Altonian, Miocene). The type is in the Auckland Museum.

Synonymy —

1942 Gemmula clifdenensis Powell, Bull. 2, Auck. Inst. Mus., p. 49, pi. 13, fig. 14.

Gemmula disjuncta Laws, 1936

Locality — Kaawa Creek, south of Port Waikato (Opoitian, Lower Pliocene). The type is in the Auckland Museum.

Synonymy —

1928 Tunis cf. duplex Suter, Bartrum & Powell, Trans. N. Z. Inst. 59, p. 151.

1936 Gemmula disjuncta Laws, Trans. Roy Soc. N. Z. 66, p. 120, pi. 17, fig. 82.

Gemmula duplex (Suter, 1917)

Locality — McCullough’s Bridge, South Canter- bury (Bortonian, Eocene). The type is in the New Zealand Geological Survey, Wellington.

Synonymy —

1917 Tunis duplex Suter, N. Z. Geol. Surv. Pal. Bull. 5,

p. 45, pi. 5, fig. 15.

Gemmula kaiparaensis (Marshall, 1918) Locality — Pakaurangi Point, Kaipara ( Altonian, Miocene ) .

Synonymy —

1918 Tunis kaiparaensis Marshall, Trans. N. Z. Inst. 50,

p. 268, pi. 18, figs. 9, 9a.

Gemmula lawsi Powell, 1942

(PI. 205, fig. 2)

Locality — Pakaurangi Point, Kaipara ( Altonian, Miocene ) .

[22 - 738]

March 31, 1964

INDO-PACIFIC MOLLUSCA, vol. 1, no. 5

Gemmula 267

Synonymy —

1942 Gemmula lawsi Powell, Bull. 2, Auck. Inst. Mus., p. 50, pi. 13, fig. 12.

Gemmula longwoodensis Powell, 1942

(PI. 205, fig. 3)

Locality — N.Z.G.S. loc. 2563, Longwood, S. D., Orepuki, Southland ( Duntroonian, Upper Oligo- cene). The type is in the New Zealand Geological Survey, Wellington.

Synonymy —

1942 Gemmula longwoodensis Powell, Bull. 2, Auck. Inst. Mus., p. 49, pi. 13, fig. 13.

Gemmula margaritata (Marshall, 1919) Locality — Hampden, Otago ( Bortonian, Eo- cene ) .

Synonymy —

1919 Turris margaritatus Marshall, Trans. N. Z. Inst. 51, p. 230, pi. 17, fig. 2.

Gemmula orba Marwick, 1931

(PI. 206)

Locality — N.Z.G.S., loc. 1322, Ormond Series, Gisborne (Upper Miocene). The type is in the New Zealand Geological Survey, Wellington.

Synonymy —

1931 Gemmula orba Marwick, N. Z. Geol. Surv. Pal. Bull. 13, p. 133, pi. 15, fig. 277.

Gemmula ornata (Marshall, 1918)

Locality — Pakaurangi Point, Kaipara ( Altonian, Miocene ) .

Synonymy —

1918 Turris ornatus Marshall, Trans. N. Z. Inst. 50, p. 268,

pi. 18, figs. 8, 8a.

Gemmula peraspera Marwick, 1931

(PI. 206)

Locality — N.Z.G.S., loc. 1322, Ormond Series, Gisborne (Upper Miocene). The type is in the New Zealand Geological Survey, Wellington.

Synonymy —

1931 Gemmula peraspera Marwick, N. Z. Geol. Surv. Pal. Bull. 13, p. 133, pi. 15, fig. 276.

Gemmula polita (Marshall, 1919)

Locality — Hampden, Otago ( Bortonian, Mio- cene ) .

Synonymy —

1919 Turris politus Marshall, 1919, Trans. N. Z. Inst. 51,

p. 230, pi. 17, fig. 9.

Plate 205. Tertiary New Zealand Gemmula. Fig. 1, G. clifdenensis Powell. Altonian, Miocene: Clifden, Southland. Fig. 2, G. lawsi Powell. Altonian, Miocene: Pakaurangi Point, Kaipara. Fig. 3, G. longwoodensis Powell. Duntroon- ian. Upper Oligocene: Orepuki, Southland (all from Powell, 1942, pi. 13, figs. 14, 12 and 13 respectively).

Plate 206. Gemmula peraspera Marwick (left fig.) and G. orba Marwick (right fig.). Upper Miocene: Gisborne, Ormond Series, New Zealand (from Marwick, 1931, pi. 15, figs. 276, 277).

[22 - 739]

268 Gemmula

A. W. B. Powell

Turridae

Gemmula reticulata (Marshall, 1919)

Locality — Hampden, Otago ( Bortonian, Eo- cene ) .

Synonymy —

1919 Tunis reticulatus Marshall, Trans. N. Z. Inst. 51, p.

231, pi. 17, fig. 8.

Gemmula waihaoensis Finlay, 1924

Locality — McCullough’s Bridge, South Canter- bury ( Bortonian, Eocene ) .

Synonymy —

1917 Tunis regius Suter, N. Z. Geol. Surv. Pal. Bull. 5, p.

46, pi. 12, fig. 14 (non Roding, 1798).

1924 Gemmula waihaoensis Finlay, Proc. Malac. Soc. 16, p.

103.

Australian Tertiary Gemmula Gemmula gellibrandensis Chappie, 1934

Remarks — This species is atypical, if Chappie’s original description is as stated: “Protoconch of two convex whorls; initial portion slightly inflated, ob- lique; anterior whorl costate.” The type is in the National Museum in Melbourne, Australia.

Locality — Princetown, % mi. W. of Gellibrand Biver, Victoria (Balcombian, Miocene).

Synonymy —

1934 Gemmula gellibrandensis Chappie, Mem. Nat. Mus.

Melb. 8, p. 163, pi. 19, figs. 3, 3a.

Gemmula samueli (Tenison-Woods, 1879)

Remarks — This species is not a typical Gem- mula; the protoconch, although multispiral, devel- ops axials only over the last 2 whorls, the initial 2M being smooth and polished. The double periph- eral carina is axially costate rather than gemmate and the broad U-shaped peripheral sinus is not so deep as in most species of Gemmula. The subsid- iary spirals, however, especially those on the base, are truly gemmate where crossed by the relatively weak axials.

In its slender fusiform shape, its tubercular rather than gemmate peripheral sculpture and its open U-shaped sinus, the species approaches Lucerapex. Members of the latter genus, however, have a smooth paucispiral protoconch.

The species samueli could represent a trend in Gemmula from which the prototype of Lucerapex could have been derived, or perhaps more likely, Lucerapex could be considered as a derivation from Fusiturris.

The Victorian Pleurotoma murrayana Pritchard, 1904, from the same horizon as samueli , appears to

be a true Lucerapex. It is described as having a protoconch of 2/2 whorls, with a blunt apex, smooth and inclined to be angled medially. The whorls are nodosely keeled and there is a broad deep sinus on the shoulder.

Measurements (mm.) —

height width

15.5 4.7 Altona Shaft

Locality — Muddy Creek, lower beds, Victoria (Balcombian, Miocene) (type locality); Altona Shaft.

Synonymy —

1879 Pleurotoma samueli Tenison-Woods, Proc. Linn. Soc.

N. S. W. 3, p. 226, pi. 20, fig. 3.

1944 Gemmula samueli Tenison-Woods, Powell, Rec. Auck. Inst. Mus. 3(1), p. 13.

Plate 207. Gemmula husamaru Nomura. Off Tiba Prefec- ture, Japan. An indeterminate species (from Nomura, 1940, pi. 1, figs. 4a, b).

[22 - 740]

March 31, 1964

INDO-PACIFIC MOLLUSCA, vol. 1, no. 5

Gemmula 269

Subgenus Unedogemmula MacNeil, 1960

Type: Tunis unedo Kiener, 1839-40

Originally proposed as a genus, Unedogemmula is here reduced to subgeneric status. It is Lophio- toma-like in its adult stage, having a plain periph- eral keel but the early whorls show distinct gem- mulations on the keel. Occasional examples have the gemmules persisting over most of the keel. It could be a Gemmula tending towards obsolescence of the gemmules or a Lophiotoma exhibiting ves- tigial evidence of the origin of that group from Gemmula. At most, Unedogemmula would appear to merit no more than subgeneric status.

Synonymy —

1960 Unedogemmula MacNeil, U. S. Geol. Survey Prof. Paper 339, p. 101 (type by original designation: Tunis unedo Kiener).

Gemmula unedo (Kiener, 1839-40)

(PI. 175, figs. 1, 6; pi. 208, figs. 1, 2)

Range — Japan, to the East Indies and the Per- sian Gulf.

Remarks — This large attractive Gemmula is more common in Japan than elsewhere in its rather wide range. It is characterized by its light yellow- ish colour over which are numerous, small flecks of darker brown, by sharply indented suture, by its deep, rather large, U-shaped sinus, by the con- cave upper shoulder, and by the numerous, rough spiral threads on the lower three-fourths of the last whorl. The peripheral carina is weakly gem- mate in the first few post-nuclear whorls, although in some specimens the beading may persist to the penultimate whorl. G. deshayesii, also from south- east Asia, differs in being an almost solid tan with a few axial, zigzag flames of cream, in being smoother, and in having a proportionately smaller and shallower sinus.

A large series of specimens from the Persian Gulf from the Townsend collection in the British Museum confirm the suspicion that Melvill’s Turns invicta are young unedo.

Description — Adult shell, 75-105 mm. (3-4 inches) in height, solid, fusiform, with a tall spire and long anterior canal. Spire a little more than the height of the aperture plus canal; spire angle 30-35. Sculptured firstly with two narrow, sharply raised, spiral cords submargining the suture, fol- lowed by a bimarginate sinus rib which forms the

peripheral angle, at a little below median whorl height, then 2-3 primary spiral cords emergent between the sinus rib and the lower suture. On the base and neck there are about seventeen primary spirals. On the concave shoulder area there are from 4 to 7 crisp secondary spirals. All the inter- spaces carry from one to three spiral threads. Whorls 12-13, exclusive of a multispiral narrowly conic, smooth protoconch of 3-3/2 whorls, terminat- ing in a half whorl of brephic axials. The early post-nuclear whorls bear distinct gemmules on the peripheral carina and in a few instances these per- sist over most of the remaining whorls. Colour pattern consisting of small reddish-brown dots, diffused into slightly larger maculations around the subsutural collar and the sinus rib, the whole loosely and irregularly connected axially by flex- uous pale reddish-brown streaks that follow the successive growth lines. Ground colour and interior of aperture white. In occasional specimens from Japan, the sinus rib carries a third but weaker spiral than the two margining ones. This feature is clearly shown in Kiener’s illustration of the spe- cies. Operculum leaf-shaped, with an apical nu- cleus.

Measurements (mm.) —

height	width
105.7	31.0	Tayabas, 190 fathoms, Philippines
94.0	26.5	Tosa, 50 fathoms, Japan
93.5	26.3	Tosa, 50 fathoms, Japan
88.5	27.0	Persian Gulf

Plate 208. Fig. 1, Gemmula (Unedogemmula) unedo (Kiener). “Mers de Unde” (from Reeve, 1843, pi. 2, fig. 12). Fig. 2, holotype of invicta Melvill, a synonym of unedo (from Melvill, 1910, pi. 2. fig. 27). Figs. 3, 4, G. (U.) binda (Garrard). Holotype. 75 fms., off Broken Bay, New South Wales, Australia. 77 mm. (photo by D. McMichael).

[22 - 761]

270 Unedogemmula

A. W. B. Powell

Turridae

87.0 28.0 Sagami Bay, Japan

81.4 24.5 Tosa, 50 fathoms, Japan

72.0 24.0 Kii, 40 fathoms, Japan

31.7 12.0 holotype of invicta Melvill

Synonymy —

1839-40 Pleurotoma unedo Kiener, Coquilles Vivantes, Pleu- rotome 5, p. 19, pi. 14, fig. 1.

1843 Pleurotoma unedo Kiener, Reeve, Conch. Iconica 1, pi. 2, fig. 12.

1884 Pleurotoma unedo Kiener, Tryon, Manual of Conch. 6, p. 165, pi. 3, fig. 20.

1910 Tunis invicta Melvill, Ann. Mag. Nat. Hist., Ser. 8, 6, p. 15, pi. 2, fig. 27.

1951 Tunis unedo Kiener, Hirase & Taki, Handb., Illustr. Shells Japan, pi. 115, fig. 2.

1954 Gemmula unedo Kiener, Kira, Coloured Illustr. Shells Japan, pi. 35, fig. 17.

1956 Gemmula unedo Kiener, Kaicher, Indo-Pacific Sea Shells, pi. 1, fig. 11.

1960 Unedogemmula unedo Kiener, MacNeil, U. S. Geol. Surv. Prof. Paper 339, p. 101.

Types — Kiener ’s type locality is “mers de l’lnde.” Melvill’s type of invicta is in the British Museum (Natural History).

Records — JAPAN : Tosa, 40-50 fathoms; Sagami Bay; Awa, Nushima, Awaji; Wakayama; Nagasaki (ANSP); Kii, 40 fathoms (A.W.B.P. coll). PHILIPPINES: Tayabas Bay, Luzon, green mud, 190 fathoms; off Lauis Point, E. Cebu, grey mud and sand, 159 fathoms; S. of Tayabas Lt., Luzon, black sand, 106 fathoms; off Point Talin, Luzon, mud, shell and coarse sand, 114 fathoms; off Pt. Dumureg, Masbate, green mud, 153 fathoms; W. of Balabac Island, 148 fath- oms; off Davao, Mindanao, 135 fathoms; S.W. of Corregidor Lt., Luzon, mud, shell and coarse sand, 118 fathoms (Alba- tross, USNM ). CHINA SEA: S. of Pratas Island, 153 fathoms (USNM). MOLUCCAS: Molucca Pass, off Makyan Island, grey mud, 275 fathoms (USNM). PERSIAN GULF (Townsend collection; type locality of invicta)-. Gulf of Oman, 73 metres (John Murray Exped., Sta. 72).

Gemmula binda (Garrard, 1961)

(PI. 208, figs. 3, 4)

Range — New South Wales, 75 fathoms.

Remarks — This species is based upon a single specimen which may yet prove to be identical with the wide-ranging unedo. The New South Wales shell appears to differ from unedo only in having a more obtusely rounded periphery with a weaker sinus rib and a shallower shoulder concavity. Also the basal spirals are more numerous and therefore more closely spaced. Examination of the type speci- men shows the sinus to be moderately deep, V- shaped, on the peripheral carina. At least six of the early post-nuclear whorls are gemmate.

Description ( original ) — “Shell large, heavily built, with tall slender spire, eleven main whorls, apical whorls missing; outer lip, although damaged, appears to have been sharp, without any folding or thickening, and sinus on peripheral keel; four- teen well developed raised revolving threads visible within the aperture; anterior canal fairly long and

straight, small portion missing; whorls sharply angled at periphery, with a broad concave shoulder and well rounded base; sculpture consists of num- erous well defined revolving lirae; interspersed with finer threads, crossed by oblique growth lines veering to right above periphery and to left below it, where sculpture has a general granulated ap- pearance; one well defined thread prominent just below suture, two other fairly smooth prominent threads form the peripheral keel, whilst sculpture generally tends to become smoother towards anter- ior end of shell; colour consists of chestnut spots on prominent thread below suture and on peri- phery, with more general blotching on lower part of main whorl; background and interior of aper- ture white.”

Measurements (mm.) —

height width

77.0 25.0 holotype

Synonymy —

1961 Tunis binda Garrard, Joum. Malac. Soc. Aust., No. 5, p. 32, pi. 1, fig. 7.

Records — NEW SOUTH WALES: off Broken Bay, 75 fathoms (type locality). The holotype is in the Australian Museum, Sydney.

Gemmula deshayesii (Doumet, 1839)

(PI. 175, figs. 7, 8; pi. 210, figs. 1, 2)

Range — Japan, China and Hongkong.

Remarks — This moderately common, east Asian species is characterized by an unusually solid shell, by its almost uniform, tawny to olive-brown colour which, however, has flexuous, axial whitish streaks. It differs from unedo (Kiener) in having a propor- tionately shorter anterior canal, in having smoother spiral cords, and in lacking numerous, small brown flecks.

Both Kiener ( 1839-40 ) and Reeve ( 1843 ) erro- neously identified this species as indica Deshayes, 1832. The latter species is larger, broader, less solid, with strong flecks and spots of brown, and known only from a single Ceylonese specimen. Tryon ( 1884 ) erroneously considered deshayesii to be a synonym of indica Deshayes because of Kiener’s and Reeve’s earlier errors. Some Japanese authors have erroneously appied the name of poly- tropa (Helbling) to this species, evidently on von Marten’s assumption in 1869 (Malakozool. Blatter 16, p. 235).

Description — Adult shell 65-70 mm. (2/2-21 inches) in height, solid, fusiform with a tall spire and long anterior canal. Spire a little more than the height of the aperture plus canal; spire angle

[22 - 762]

March 31, 1964

INDO-PACIFIC MOLLUSCA, vol. 1, no. 5

Gemmula 271

27° -30°. Sculpture rather uniformly and closely lirate except for one smooth primary cord below the suture and a not very prominent bicarinate sinus rib, situated just below the middle of whorl height. Below this and over the base there is a closely packed, rather indefinite assortment of pri- mary and secondary cords and interstitial threads. There are 5 or 6 crisp threads on the shoulder area; and 4 to 6 of the post-nuclear whorls are dis- tinctly gemmate. Whorls of rather straight outline, shoulder weakly concave and sinus rib not promi- nently projecting. Number of whorls 10-11, exclu- sive of a polygyrate, narrowly conic, smooth pro- toconch of approximately four whorls, the last axially costate. None of the available protoconchs are sufficiently well preserved for more accurate description. Colour uniformly olive or yellowish olive with a few white, flexuous, axial streaks which follow the trend of the growth lines and appear to coincide with rest stages of growth. Interior of aperture, columella and anterior canal white.

Measurements (mm.) —

height	width
64.0	18.5	Hongkong
61.0	18.0	Hongkong
60.0	18.25	Hongkong
54.0	15.5	Kii, Japan

Synonymy —

1839 Pleurotoma deshayesii Doumet, Rev. Zool. ( Soc. Cuv. )

2, p. 325 (description only).

1840 Pleurotoma deshayesii Doumet, Guerin’s Mag. Zool.,

Anat. Comp. & Zool., pi. 11 (title page dated 1844, plate 1840). “Mers de la Chine.”

1843 Pleurotoma deshayesii Doumet, Reeve, Conch. Ieonica 1, pi. 3, fig. 19 (China).

1876 Pleurotoma deshayesii Doumet, Weinkauff, Conch.

Cab. 4(3), p. 26, sp. 22, pi. 5, fig. 7.

1884 Pleurotoma indica Deshayes (in part), Tryon, Manual of Conch. 6, p. 168, pi. 6, fig. 80 only.

1886 Pleurotoma deshayesii Doumet, Watson, Challenger Zool. 15, p. 284.

1960 Tunis polytropa Helbhng, Kira, Coloured Illustr. Shells Japan, pi. 35, fig. 16.

Records — JAPAN : Fukura, Awaji (USNM); Minoshima, Wakayama-ken; Kii (ANSP and A. W. B. Powell coll.). CHINA: Foochow; Amoy; Pei-tai-ho, N. Chihli coast (USNM). TAIWAN: (ANSP). HONG KONG: (USNM; A. W. B. Powell coll.); 20-30 fms., Hong Kong (ANSP); Aap Li Chaau, Hong Kong ( R. D. Purchon, ANSP ) .

Gemmula indica (Deshayes, 1832)

(PI. 209)

Range — Known only from Ceylon.

Remarks — This large attractive species is evi- dently known from only one specimen which was collected by Ch. Belanger in Ceylon. I have not had an opportunity to examine the type, and have provisionally placed it in the subgenus Unedo- gemmida. It resembles unedo (Kiener), but ap-

Plate 209. “Pleurotoma” indica Deshayes. Ceylon (from Belanger, 1832, pi. 2, figs. 9, 10).

pears to differ in having a proportionately broader body whorl, a shorter anterior canal, more rounded whorls in the spire and in having a much darker brown coloration. G. indica (Deshayes) is umbili- cate, but this feature may be due to an abnormality which is known to occur in other species. Should this species be re-discovered and prove to be a true Turris or a Lophiotoma it will have to be re-named because of Lophiotoma indica (Roding, 1798).

Description — Shell 80 mm. (about 3 inches) in length, fusiform, moderately solid, with a high spire, a broad body whorl, and a rather short an-

Plate 210. Figs. 1, 2, Gemmula (Unedogemmula) de- shayesii (Doumet). Awaji, Japan. 60 mm. Fig. 3, G. (U.) ina MacNeil. Miocene of Okinawa Id., Ryukyu Ids. 40.2 mm. (from MacNeil, 1960, pi. 5, fig. 7). Fig. 4, G. (U.) kool- hoveni Oostingh. Pliocene of South Bantam, Java Id., Indo- nesia (from Oostingh, 1938, pi. 1, fig. 22).

[22 - 763]

272 Unedogemmula

A. W. B. Powell

Turridae

terior canal which is umbilicate. Sinus deep and U-shaped. Whorls in spire convex and only slightly angulate. Sinus scar appears to be flattish and axially striate. Color of shell yellowish brown with darker brown axial, narrow streaks on the body whorl, and with dark-brown spots and spiral streaks on the sinus scar and on the numerous spiral subsutural threads.

Measurements (mm.) —

height width

80 26 from the type figure

Synonymy —

1832 Pleurotoma indica Deshayes, in Belanger’s Voyage aux Indes-Orientales, Pt. 2, Zoologie, p. 421, Mollusques pi. 2, figs. 9, 10 (December).

Types — The type locality is the “cotes de Cey- lan.” I do not know the present location of the type.

Records — Known only from Ceylon (Deshayes, 1832).

Gemmula hastula (Reeve, 1843)

(PI. 211)

Range — Persian Gulf and Arabian Sea.

Remarks — This species is based upon a single well-preserved specimen of unknown locality in

Plate 211. Gemmula (Unedogemmula) hastula (Reeve, 1843). Holotype from the British Museum (Natural His- tory). Locality unknown, but probably the Persian Gulf. 38.5 mm.

the British Museum. However, a series of speci- mens in the John Murray Expedition material is from the Persian Gulf, and a further locality is added by an undoubted synonym, Pleurotoma try- panodes Melvill, 1904, from the Arabian Sea.

This species is certainly not a synonym of indica Roding, 1798 ( = marmorata Lamarck, 1822 ) as claimed by Tryon (1884). From indica it is dis- tinguished by more prominent subsutural and less prominent peripheral spirals, resulting in a regu- larly conic not pagodiform spire. Further, the pro- toconch of four whorls, two smooth and two axially costate, followed by weak gemmules on the peri- pheral carina of the first post-nuclear whorl, make the reference of the species to the subgenus Unedo- gemmula positive.

Melvill’s Pleurotoma trypanodes is based upon a bleached and somewhat worn example of hastula. Melvill’s illustration gives the impression of rather profuse granulation of the spiral cords, but except for weak granules on the carina of the first 2 or 3 post-nuclear whorls, subsequently it is not the cords that are granulate but a false impression of granu- lation resulting from dense axial threads between the cords and these are strongest above the peri- pheral carina.

Description — Shell rather small and slender, 21- 32 mm. (about 1 inch) in height, with a tall, nar- rowly conic spire and a long, almost straight anter- ior canal. Spire very slightly less than the height of the aperture plus canal. Adult sculpture of reg- ular, smooth, spiral cords, the most prominent being a closely spaced, subsutural pair and a single peripheral one which terminates in a deep, narrow sinus. Three closely spaced spiral threads present between the subsutural and the peripheral carinae, and two pairs of threads, alternately strong and weak, are between the peripheral carina and the lower suture. Base rather densely sculptured with relatively strong and weaker intermediate spirals. Colour whitish, obscurely axially marked and dif- fused with yellowish-brown.

Measurements (mm.) —

height	width
21.0	7.0	holotype of trypanodes Melvill
28.0	8.0	Persian Gulf, 73 metres
29.0	9.0	Arabian Sea, 40 fathoms
32.5 Synonymy	9.0	holotype of hastula Reeve

1843 Pleurotoma hastula Reeve, Conch. Iconica 1, pi. 17, fig. 139 (locality unknown).

1904 Pleurotoma trypanodes Melvill, Proc. Malac. Soc. 6, p. 57, pi. 5, fig. 2.

1917 Tunis (Tomopleura) trypanodes Melvill, Proc. Malac. Soc. 12, p. 148.

[22 - 764]

March 31, 1964

INDO-PACIFIC MOLLUSCA, vol. 1, no. 5

Gemmula 273

Plate 212. Geographical distribution of Gemmula (Unedo- gemmula) unedo (Kiener), deshayesii (Doumet), hastula (Reeve), and binda (Garrard).

Types — The types of both hastula Reeve and trypanodes Melvill are in the British Museum (Natural History).

Records — ARABIAN SEA: 18° 58' N; 71° 45' E., 40 fathoms (type locality of trypanodes) ; 100 miles W. of Bombay, on cable ( Brit. Mus. ) . PERSIAN GULF : 25° 38' 18" N.; 56° 26' 36" E., 73 metres (John Murray Exped. Sta. 72; Brit. Mus.).

Gemmula bemmeleni (Oostingh, 1941)

Remarks — The author of this species remarks that it is very close to his koolhoveni but it has more numerous spirals. He compares it with “polytropa Helbling” (i.e., Lophioturris leucotropis Adams & Reeve, 1850), but bemmeleni by its gemmate early whorls is clearly a Unedogemmula.

Measurements (mm.) —

height width spire angle

30 18 35

angle more prominent and situated at about the lower third of whorl height, which gives a pagodi- form profile to the spire. Also the shoulder is straight, not deeply concave as in these other Re- cent species. MacNeil compared his species with indica Roding, but that species has a paucispiral protoconch, lacks gemmules on the early whorls and is therefore more satisfactorily located in Lophioturris.

Description ( original ) — “Shell of medium size, fusiform. Protoconch not preserved on type. Aper- ture about half as long as shell, produced anter- iorly to form a straight canal. Anal sinus as deter- mined by growth lines openly V-shaped, slightly broader at the inner end than the peripheral keel. Parietal callus thin. Whorls angulate with a promi- nent peripheral keel. Sculpture consisting of mod- erately strong revolving lines on the lower part of the body-whorl with secondary and tertiary threads and less prominent revolving lines on the sub- sutural slope and columella. Subsutural collar per- sistent in adult stage, very strong in young gem- mulate stage.”

Synonymy —

1941 Tunis (Gemmula) bemmeleni Oostingh, De Ingenieur in Ned. -Indie, No. 7, p. 64, fig. 2.

Records — Central JAVA, Semarang (Pliocene). Gemmula ina MacNeil, 1960 (PI. 210, fig. 3)

Range — Okinawa ( Miocene ) .

Remarks — This fossil species differs from both unedo and deshayesii in having the peripheral

Measurements (mm.) —

height W idth

40.2 13.0 holotype

Synonymy —

1960 Unedogemmula ina MacNeil, U. S. Geol. Surv. Prof. Paper 339, p. 102, pi. 5, fig. 7.

Type locality — Okinawa (Yonabaru clay, Mio- cene ) .

[22 - 765]

274 Unedogemmula

A. W. B. Powell

Turridae

Gemmula koolhoveni (Oostingh, 1938)

(PL 210, fig. 4)

Remarks — This species, described as a Turris (Gemmula), has gemmules only on the upper spire whorls and in this and in its general likeness to unedo Kiener suggests location in the subgenus Unedogemmula. From unedo, the Javanese Plio- cene species differs mainly in its stronger spiral sculpture.

Measurements (mm.) —

height width

39 12 holotype

50.5 - ( Oostingh )

Synonymy —

1938 Turris (Gemmula) koolhoveni Oostingh, Die Moll. Plioc. Sud. Bantam in Java, De Ingen. Ned.-Indie, Gast. I, p. 28, pi. 1, fig. 22.

Records — JAVA: South Bantam (Pliocene).

Gemmula sondeiana (Martin, 1895)

(PI. 213, figs. 4-6)

Range — Sonde, Java Island, Indonesia, Pliocene.

Remarks — This is by far the most slender-spired member in the subgenus Unedogemmula. The spire is very tall and narrow, but the anterior canal rela- tively short, and the sculpture is of closely-spaced, smooth spirals, except for the not very prominent peripheral carina which is gemmate over the early whorls. Martin did not give any size for this spe- cies.

Plate 213. Figs. 1-3, Gemmula ( Unedogemmula ) hay deni ( Vredenburg). Post-Eocene, Mekran beds, India (from Vredenburg, 1925, pi. 9, figs. 4, 5). Figs. 4-6, G. (U.) son- deiana (Martin). Pliocene of Sonde, Java Id., Indonesia (from K. Martin, 1895, pi. 6, figs. 89, a, b).

Synonymy —

1895 Pleurotoma sondeiana Martin, Die Fossilien von Java. Geol. Reichs-Mus., Leiden, 1, p. 35, pi. 6, figs. 89a, b.

Records — JAVA: Sonde (Pliocene). SUMATRA: Plio- cene (K. Martin, 1928, p. 12).

Gemmula haydeni (Vredenburg, 1925)

(PI. 213, figs. 1-3)

Range — India, post-Eocene.

Remarks — The material upon which this spe- cies is based is not very well preserved, for all the figured specimens are without the protoconch and most of the anterior canal. Vredenburg compared his species both with the Javanese Pliocene son- deiana Martin, 1895, and with the living species deshayesii Doumet, 1839. Vredenburg’s remark that “the sinus band is more distinctly crenulated at early stages of growth in Pleurotoma sondeiana than in Pleurotoma haydeni” suggests that both are in Unedogemmula.

The species haydeni, judging from the illustra- tions and very full description, differs from the Recent deshayesii mainly in the increased strength of the secondary spiral sculpture which renders the sinus rib less conspicuous.

Measurements (mm.) —

height width

59+ 21

Synonymy —

1925 Pleurotoma haydeni Vredenburg, Mem. Geol. Surv.

India 50(1), p. 44, pi. 9, figs. 3-5.

Records — INDIA: west of Gharh Hill (Mekran beds, post-Eocene ) .

Gemmula ickei (Martin, 1906)

(PI. 214)

Range — Lower Miocene of Java.

Description (from Vredenburg, 1925, p. 40) — Shell rather large, slender, with elongate conical spire, measuring nearly three-fifths of the total height, and with a rather large body whorl, pos- teriorly globose, anteriorly rather abruptly con- tracted into a narrow, rather elongate stem, either straight or else steeply tortuous toward its extrem- ity. The protoconch, slightly oblique to the axis of the spire proper, is conoidal and consists of a minute nucleus followed by four convex whorls, the first two of which are smooth, while the two next whorls are decorated with numerous, some- what oblique ribs stretching from suture to suture.

March 31, 1964

INDO-PACIFIC MOLLUSCA, vol. 1, no. 5

Gemmula 275

Synonymy —

1906 Pleurotoma ickei Martin, Samml. Geol. Reichs-Mus., Leiden, n.s., 1, p. 293. pi. 43, fig. 703.

1925 Pleurotoma ickei Martin, Vredenburg, Mem. Geol. Surv. India 50(1), p. 40.

Records — JAVA: Tjadasngampar, Rembang beds. Lower Miocene, type locality.

Plate 214. Gemmula (U nedogemmula) ickei (Martin). Lower Miocene of Java Id., Indonesia (from K. Martin, 1906, pi. 43, fig. 703).

Gemmula ickei subspecies virginoides (Vredenburg, 1925)

Range — India, post-Eocene.

Remarks — This subspecific name was intro- duced in a curious way as follows: “Should it therefore be considered necessary to distinguish the Indian specimens, they might be taken to rep- resent a variety virginoides.” The comparison was with the typical species from the Miocene of Java.

Vredenburg’s figures are very indistinct but his remarks, quoted below, indicate an U nedogem- mula: “Compared with the specimens of ickei Martin from the Miocene beds of Tjadasngampar on the Tji Longan in Java the beautiful shell above described agrees in every essential character, with the exception of the crenulations of the sinus ridge which disappear at an earlier stage in the Indian specimens.”

The Indian shell is very slender and the spire whorls are strongly sculptured with three spiral ridges. The protoconch is described as conoidal with a minute nucleus, followed by four convex whorls, the first two smooth and the rest axially costate.

Measurements (mm.) —

height width

46+ 15

Synonymy —

1925 Pleurotoma ickei virginoides Vredenburg, Mem. Geol.

Surv. India 50(1), p. 44, pi. 1, figs. 8, 9.

Records — INDIA: Gaj of Kach, Teyra River near Ram- pur (post-Eocene).

[22 - 767]

276 Unedogemmula

A. W. B. Powell

Turridae

[These occasional blank areas occur between genera and subgenera to permit the insertion of new material and future sections in their proper systematic sequence.]

[22 - 768]

March 31, 1964

INDO-PACIFIC MOLLUSCA, vol. 1, no. 5

Pinguigemmula 277

Genus Pinguigemmula MacNeil, 1960

Type: Pinguigemmula okinavensis MacNeil, 1960

This genus resembles Gemrnula but has a broadly conic spire, a strongly contracted base and a moderately long, straight anterior canal. In gen- eral, all of the spirals at and above the periphery are strong and beaded, while those below the peri- phery are smooth and much weaker. The proto- conch unfortunately is not known, but from frag- ments remaining is judged to be multispiral. It is something more than a bizarre-shaped Gemmula, for the sinus is of very different form from the deep narrow peripheral V-shape of that genus. In Pin- guigemmula the sinus is very shallowly, broadly open, descending straight at about 30° to the axis, over the shoulder area, then sweeping forward tangentially from the peripheral carina, almost at right angles to the upper arm. All of the Miocene- Pliocene specimens of the type species and most of the Recent shells examined exhibit a curious ab- normality of growth in the form of twisted flutings of the outer lip, comparable with those observed in some specimens of Gemmula congener diomedea (new subspecies) and in two species of Ptychosy- rinx.

Speculating on the possible reason for this ab- normality, MacNeil ( 1960 ) suggested that it may represent a response to an oxygen-poor environ- ment in which one or more incurrent siphons are developed. Whatever the cause, it seems to be a gerontic condition which suddenly develops after

full size has been achieved. MacNeil’s explanation, however, seems unlikely, since most of the flutings are not hollow.

Synonymy —

1960 Pinguigemmula MacNeil, U. S. Geol. Survey Prof. Paper 339, p. 103. Type by original designation: P. okinavensis MacNeil, 1960.

Pinguigemmula okinavensis MacNeil, 1960

(PI. 215, figs. 1, 2)

Range — Okinawa ( Miocene or Pliocene ) .

Description — Adult shell 41.5 mm. (about 1M inches ) in height, stout, with a broadly conic spire, less than height of aperture plus anterior canal, which is moderately long and straight. Protoconch unknown. Subsutural slope bearing 4 strong beaded spirals with deeply channeled interspaces, the lowest one bearing 2 rows of beads and form- ing the peripheral carina. Sculpture below the periphery consisting of rounded unbeaded spirals which are coarsest below the periphery and be- come less coarse on the base of the whorl and colu- mella, those on the columella commonly inter- spaced with secondary spirals. Suture closed and hidden in a deep groove. Anal sinus terminating on the peripheral carina, asymmetrically V-shaped with the lower limb nearly horizontal and the up- per limb inclined nearly 45 degrees. (From Mac- Neil’s original description. )

Measurements (mm.) —

height width spire angle

41.5 20.2 45° to 60°

Synonymy —

1960 Pinguigemmula okinavensis MacNeil, U. S. Geol. Surv. Prof. Paper 339, p. 104, pi. 9, figs. 12-14.

Plate 215

Figs. 1, 2 Pinguigemmula okinavensis MacNeil. Holotype, fig. 2. Miocene-Pliocene, Okinawa Id., Ryukyu Ids. 41.5 mm.

3, 4 Pinguigemmula luzonica new species. Holotype.

178 fms., off Menor Id., Luzon Id., Philippines.

35.0 mm.

5, 6 Pinguigemmula philippinensis new species. Holo- type. 280 fms., off Santiago, Luzon Id., Philip- pines. 50.2 mm.

7 Pinguigemmula thielei (Finlay). Holotype of Pleurotoma (Gemmula) fusiformis Thiele, 1925, non Sowerby, 1823. 614 meters off west coast of Sumatra Id., Indonesia.

[22 - 789]

278 Pinguigemmula

A. W. B. Powell

Turridae

Types — The type locality is Okinawa ( Shinzato tuff, Miocene or Pliocene).

Pinguigemmula luzonica new species (PL 215, figs. 3, 4)

Range — Philippine Islands, 178 to 297 fathoms.

Remarks — This species closely resembles the Miocene-Pliocene type of the genus, from which it is distinguished chiefly by its smaller angle of spire, by its slightly longer anterior canal, and in having only two gemmate cords on the spire whorls above the peripheral carina.

Description — Adult shell 35 mm. ( about 1/2 inches) in height, broadly fusiform, with a squat spire and deeply contracted base, produced into a moderately long tapered and perfectly straight an- terior canal. Spire three-fourths of the height of the aperture plus canal. Whorls 9, exclusive of the protoconch of which only the eroded last whorl remains in the holotype. Spire broadly conical, outlines at first slightly concave then lightly convex over the last 4 whorls. Sculpture of spire whorls consisting of 3 very heavy, revolving, flat-topped keels with deeply channelled, narrow, almost linear, interspaces, one subsutural, one peripheral and one intermediate. The subsutural keel bears 3 weak spiral threads; the intermediate keel bears 2 threads; and the third or sinus keel bears 3 threads. All three keels are regularly and very closely gem- mate, those of the sinus rib in the form of 3 gem- mules fused vertically. Base, including the anterior canal, with 18 smooth primary cords and an inter- mediate thread in each interspace over the anterior half of the canal. Sinus as described for the genus, with its upper limb descending almost straight, then projecting tangentially forward, but in the case of the holotype, which exhibits the gerontic state mentioned above, the lower edge of the sinus is bent sharply downward along with a fluted and thickened extension of the second basal spiral. Colour uniformly very pale-buff.

Measurements (mm.) —

height width spire angle 35.0 14.0 45° holotype

Types — The holotype is in United States Na- tional Museum, Washington, no. 237784.

Records — PHILIPPINES: off Hermana, Menor Id., Lu- zon Id., 178 fins., sand, shell and mud (Albatross Sta. 5331) (type locality); off Hermana, Menor Id., 297 fms., green mud (Albatross Sta. 5438, USNM).

Pinguigemmula philippinensis new species (PI. 215, figs. 5, 6)

Range — Off Santiago, west Luzon Id., Philip- pines, in 280 fathoms.

Remarks — This species is larger and more slen- der than either okinavensis or luzonica, but it is nearer in shape and sculptural details to thielei Finlay from which it differs in having three strong spiral keels instead of two, additional to the peri- pheral carina, which projects flange-like, in thielei and philippinensis, but not in either okinavensis or luzonica.

Description — Adult shell 43 to 50 mm. ( about 1/2 to 2 inches) in height, fusiform, with a spire tall yet broadly conic, with straight outlines, tabulated by the peripheral, broadly rounded, flange-like keel which is situated almost at the lower suture. Base rapidly contracted to a long-tapered, almost straight, anterior canal. Whorls 8 to 9, exclusive of protoconch, which is missing in all examples. Sculp- ture of spire whorls consists of 4 strong, flat-topped spiral keels with deeply channeled interspaces. The peripheral sinus rib projects as a flange beyond the other three. All four keels gemmate, those on the sinus keel stronger and more regular. Base, including the anterior canal, with 25 to 27 smooth spirals, those on the base rather strong, especially the upper two, but becoming weaker and more closely-spaced over the anterior canal. Spire three- fourths of the height of the aperture plus canal. Sinus broadly and shallowly V-shaped, descending in a straight slope at 30 degrees to the axis, then

Plate 216. Geographical distribution of Pinguigemmula okinavensis MacNeil, P. philippinensis new species, P. luzon- ica new species, and P. thielei (Finlay).

(

(

(

[22 - 790]

March 31, 1964

INDO-PACIFIC MOLLUSCA, vol. 1, no. 5

Pinguigemmula 279

produced tangentially forward at right angles. A paratype exhibits a massive tubular fluting in the form of a coalescence of the upper two basal spi- rals. It develops suddenly within 5 mm. of the outer lip edge and projects downward for 5 mm. as a partially closed tube. Colour pale yellowish buff. Parietal callus and interior of aperture white.

Measurements (mm.) —

height width spire angle 50.2 18.7 40° holotype

43.5 17.2 42°

Types — The holotype is in United States Na- tional Museum, Washington, no. 237563.

Records — PHILIPPINES: off Santiago, west Luzon Id., Philippines, in 280 fms. (Albatross Sta. 5283, USNM, type locality ) .

Pinguigemmula thielei (Finlay, 1930)

(PI. 215, fig. 7)

Range — Off west coast of Sumatra, Indonesia in 614 metres.

Remarks — This species and philippinensis are obviously closely allied, but unfortunately no ex- amples of thielei are available to the writer. As- suming that Thiele’s illustration is accurate, thielei differs from philippinensis in having two instead of three spiral keels in addition to the peripheral flange-like keel. Also, in Thiele’s figure, only the peripheral keel is shown to be gemmate, whereas in philippinensis all four spiral keels are gemmate to some extent.

Measurements (mm.) —

height width spire angle 44.0 18.0 45° holotype

Synonymy —

1925 Pleurotoma (Gemmula) fusiformis Thiele, Wissen- schaft. Ergebn. Deutschen Tiefsee-Exped. 17, Gastr. 2, p. 210, pi. 22, fig. 24 (non Pleurotoma fusiformis I. de C. Sowerby, 1823, non Anton, 1839, non C. B. Adams, 1850).

1930 Gemmula thielei Finlay, Trans. N. Z. Inst. 61, p. 47 (nom. nov. for PI. (Gemmula) fusiformis Thiele, 1925).

Types — The type locality is off the west coast of Sumatra, “Valdivia” Sta. 194; 0° 15.2' N.; 98° 8.8' E., 614 metres.

Genus Cryptogemma Dali, 1918

(PI. 217, fig. a)

Type: Gemmula benthina Dali, 1908

Shells of this genus are rather broadly fusiform but with a short twisted anterior canal. Weak axial ribs are produced into rounded nodules on a con- spicuous peripheral keel. The sinus is at the peri- phery, rather slight and square-cut at the apex of a broadly open V. Apical whorls unknown. The outer shell is covered by a greenish grey periostra- cum. Operculum oval, with an apical nucleus. Ra- dula of the “wishbone type.” Recent, deep water, 1200 to 1300 fathoms, Gulf of Panama to Ecuador. Height, 10 to 30 mm.

This genus may later come into use for some at present undescribed species from deep water off Hawaii.

Synonymy —

1918 Cryptogemma Dali, Proc. U. S. National Mus. 54 (2238), p. 325. Type by monotypy: Gemmula benthina Dali, 1908.

Characteristic Species — C. benthina Dali, 1908; quentinensis Dali, 1919; serilla Dali, 1908.

Plate 217. Radulae of (a) Cryptogemma benthina (Dali, 1908) and (b) Carinoturris adrastia (Dali, 1919). Both drawings prepared by J. P. E. Morrison, U.S. National Mu- seum.

[22 - 7911

280 Pinguigemmula

A. W. B. Powell

Turridae

[These occasional blank areas occur between genera and subgenera to permit the insertion of new material and future sections in their proper systematic sequence.]

[22 - 792]

March 31, 1964

INDO-PACIFIC MOLLUSCA, vol. 1, no. 5

Carinoturris 281

Genus Carinoturris Bartsch, 1944

(PI. 217, fig. b)

Type: Cryptogemma adrastia Dali, 1919 Very similar to Cryptogemma in shape but with a plain narrowly rounded peripheral keel. Anal sinus at periphery, wide and shallow. Shell white, covered with a pale-olive periostracum. Protoconch of one smooth rounded whorl. Operculum small, broadly ovate with an apical nucleus. Radula of “wishbone” type. Height, 14 to 16 mm. Recent, deep-water, 300 to 600 fathoms off California.

Synonymy —

1944 Carinoturris Bartsch, Proc. Biol. Soc. Washington 57, p. 60. Type by original designation: Cryptogemma adrastia Dali, 1919.

Characteristic Species — C. adrastia Dali, 1919; fortis Bartsch, 1944; polycaste Dali, 1919.

Genus Hemipleurotoma Cossmann, 1889

Type: Pleurotoma archimedis Bellardi, 1878 This name is based upon a very rare fossil from the Helvetian middle Miocene of Italy, but in 1896, Cossmann invalidly proposed denticula Basterot, 1825, as neotype of his genus, but the original ci- tation must stand. Based upon this original type designation, about the only distinguishing charac- teristic of Hemipleurotoma is the very broadly open V-shaped sinus, otherwise it conforms with Gemmula. Glibert (1960, p. 4) includes Hemi- pleurotoma in the synonymy of Gemmula. The genus has been erroneously applied to the Recent Tunis cryptorrhaphe (Sowerby, 1825).

Synonymy —

1889 Hemipleurotoma Cossmann, Ann. Soc. Malac. Belgique 24, p. 264. Type by original designation: Pleuro- toma archimedis Bellardi, 1878; 1896, Essais de Paleoconch. Comp. 2, p. 78. Type by subsequent designation [invalid]: Pleurotoma denticula Bast- erot, 1825.

Genus Coronia Gregorio, 1890

Type: Pleurotoma childreni Lea, 1833 This is usually considered a synonym of Gem- mula, but Gardner, 1945 (Memoir no. 11, Geol. Soc. of America, p. 240) gave it full generic status upon the form of the protoconch, which is of two or more smooth, rapidly enlarging whorls, suc- ceeded by one or more convex whorls sculptured with numerous obliquely arcuate costae. Smooth initial whorls, to a varying degree, however, occur among the Indo-Pacific Recent species of Gem- mula, and these minor deviations would appear to have no special significance. Claiborne Eocene of the southern United States and northern Mexico.

Synonymy —

1890 Coronia Gregorio, Ann. de Geol. et de Paleontologie, Palermo, 7, p. 23. Type by subsequent designation by Cossmann, 1896: Pleurotoma acutirostra Conrad = (fide Gardner, 1945, p. 240) childreni Lea, 1833. [Coronia Ehrenberg, 1840, Bericht Verh. Akad. Wiss. Berlin, p. 206, no. 106, is a Diatom and does not preoccupy Coronia Gregorio].

Characteristic species — childreni Lea, 1833; genitiva Casey, 1904; margaritosa Casey, 1904.

Genus Trypanotoma Cossmann, 1893

Type: Pleurotoma terebriformis Meyer, 1886 A small shell with a very tall spire but trun- cated base. Protoconch blunt, of 2M smooth whorls. Adult whorls almost flat in outline but with a med- ian placed noduliferous peripheral weak carina. Sinus weak, peripheral. Claiborne and Jackson Eocene of Alabama and Louisiana.

Synonymy —

1893 Trypanotoma Cossmann, Essais de Paleont. Comp. 2, p. 109. Type by original designation: Pleurotoma terebriformis Meyer, 1886. Eocene.

Genus Sinistrella Meyer, 1887

Type: Triforis americanus Aldrich, 1885 This genus is usually considered a sinistral form of Trypanotoma but Harris & Palmer ( 1947, p. 423 ) give it full generic status, but in association with Coronia, Trypanotoma and Infracoronia. Known apparently only from the type species, which is always sinistral and comes from the Jackson Eocene of Mississippi.

Synonymy —

1887 Sinistrella O. Meyer, Ber. Senckenb. naturf. Ges., for 1887, p. 18. Type by subsequent designation (Coss- mann, 1893, p. 110): Triforis americanus Aldrich, 1885.

Subgenus Infracoronia Palmer, 1947

Type: Gemmula indoviciana (Vaughan) Harris, 1896 This is compared with Coronia from which it is stated to differ in having the dentate carina at the base of each whorl, just above the suture, and the absence of a subsutural band. It was proposed, however, as a subgenus of Sinistrella. Lower Clai- borne and Jackson Eocene of Louisiana, southern United States.

Synonymy —

1947 Infracoronia Palmer, in Harris and Palmer, Bull. Amer. Paleont. Inst., Ithaca, 30(117), Sec. 3, p. 423. Type by original designation: Gemmula indo- viciana (Vaughan) Harris, 1896. Subgenus of Gem- mula.

[22 - 823]

282 Campylacrum

A. W. B. Powell

Turridae

Genus Hesperiturris Gardner, 1945

Type: Turns nodocarinata Gabb, 1860

This is another member of the Gemmula complex which was diagnosed primarily upon the form of the protoconch which was described as of 5 to 512 volutions, the initial turn minute and largely im- mersed, the 3 to 3/2 succeeding whorls also smooth and shining, broadly rounded, and increasing rather rapidly in diameter, the final whorl in whole or in part axially costate. The adult shell has a tall nar- row spire but a truncated anterior canal. The sculp- ture is of nodules on a peripheral carina just above the lower suture. Sinus broadly U-shaped, moder- ately deep, the axis running closer to the periphery than to the posterior suture. The genus appears to be confined to the lower Claiborne Eocene of the southern United States and northern Mexico.

Synonymy —

1945 Hesperiturris Gardner, Memoirs no. 11, Geol. Soc. America, p. 237. Type by original designation: Tunis nodocarinata Gabb, 1860.

Characteristic Species — H. amichel Gardner, 1945; nodocarinata Gabb, 1860; zacatensis Gard- ner, 1945.

Genus Campylacrum Finlay & Marwick, 1937

Type: Campylacrum sanum Finlay & Marwick, 1937

This is another member of the Gemmula assem- blage, nearest to Eoturris, but of smaller size, dif- ferent apex and a twisted canal. So far, it is known only from the Upper Cretaceous (Wangaloan) of New Zealand. There is apparent a relationship also with the Parisian Eocene Oxyacrum Cossmann; both have a small, conic, polygyrate protoconch, and a wide, moderately deep, bluntly triangular sinus, situated on the peripheral keel, with the apex of the sinus not encircling the peripheral nodules, but occurring immediately above them.

Synonymy —

1937 Campylacrum Finlay and Marwick, N. Z. Geological Survey Paleo. Bull. 15, p. 84. Type by original des- ignation: C. sanum Finlay and Marwick, 1937.

Campylacrum debile Finlay & Marwick, 1937

Range — Wangaloa, South Island, New Zealand (Wangaloan, Upper Cretaceous).

Synonymy —

1937 Campylacrum debile Finlay & Marwick, N. Z. Geol. Surv. Paleo. Bull. 15, p. 86, pi. 12, figs. 1, 2.

Types — The holotype is in the Auckland Mu- seum.

Plate 218. Figs, a and b, Campylacrum sanum Finlay and Marwick. Upper Cretaceous of Wangaloan, New Zealand (from the original, 1937, pi. 12, figs. 3, 4). Fig. c, Eoturris multicinctus (Marshall, 1917). Eocene: Bortonian of Castle Hill Shaft (?). Otago, New Zealand (from Finlay and Marwick, 1937, pi. 16, fig. 2).

Campylacrum sanum Finlay & Marwick, 1937

(PI. 218, figs, a, b)

Range — Wangaloa, South Island, New Zealand (Wangaloan, Upper Cretaceous).

Synonymy —

1937 Campylacrum sanum Finlay & Marwick, N. Z. Geol. Surv. Paleo. Bull. 15, p. 86, pi. 12, figs. 3, 4.

Types — The holotype is in the Auckland Mu- seum.

Genus Eopleurotoma Cossmann, 1889

Type: Pleurotoma multicostata Deshayes, 1834

This is a Paleocene-Eocene relative of Gemmula in which the axial sculpture is in the form of long, narrow flexuous ribs which thicken at the peri- phery to form a row of laterally compressed oblique nodules.

The shell is 15 to 25 mm. in height, elongate- fusiform, but with a relatively short anterior canal. Protoconch small, smooth and paucispiral. Anal sinus broadly V-shaped, its apex extending a little above the peripheral nodules. It occurs in Europe, south western United States and Peru and is re- corded also from the Tertiary of southern and south eastern Asia and Japan, but the species re- quire re-evaluation.

Synonymy —

1889 Eopleurotoma Cossmann, Ann. Soc. Malac. Belgique 24, p. 269. Type by original designation: Pleuro- toma multicostata Deshayes, 1834; 1906, Cossmann, Essais de Paleoconch. Comp. 7, pi. 14, fig. 16.

1904 Eodrillia Casey, Trans. St. Louis Academy 14, p. 159, Type by subsequent designation (Cossmann, 1904, p. 237): Pleurotoma depygis Conrad.

March 31, 1964

INDO-PACIFIC MOLLUSCA, vol. 1, no. 5

Eoturris 283

Characteristic Species - EUROPEAN EOCENE: bernayi Boury, 1899; bezanconi Boury, 1899; bica- tenata Lamarck, 1804; cedilla Edwards, 1861; cur- vicosta Lamarck, 1804; distans Deshayes, 1865; distanticosta Cossmann & Pissarro, 1900; expedita Deshayes, 1865; flexicosta Boury, 1899; fluctuosa Deshayes, 1865; francisci Raincourt, 1876; fresvillen- sis Cossmann & Pissarro, 1900; granifera Deshayes, 1834; inculta Sowerby, 1850; insueta Boury, 1899; lajonkairei Deshayes, 1834; larteti Deshayes, 1865; lima Edwards, 1861; multicostata Deshayes, 1834; multinoda Lamarck, 1804; oligocolpa Cossmann, 1889; plicaria Deshayes, 1834; pourcyensis Coss- mann, 1901; propinqua Deshayes, 1865; rotella Ed- wards, 1861; rudiuscula Cossmann, 1889; scalarata Edwards, 1861; specialis Boury, 1899; spreta Desh- ayes, 1865; undata Lamarck, 1804.

NORTH AMERICAN EOCENE: adolescens Harris, 1937; albirupsis Harris, 1947; cainei Harris, 1937; carya Harris, 1937; cochlea Harris, 1937; depygis Conrad, 1833; desnoyersii Lea, 1833; gem- mavia Harris, 1937; hoeninghausii Lea, 1833; lis- boncola Harris, 1937; nupera Conrad, 1833; orange- bur gensis Harris, 1937; ouachitensis Harris, 1937; plumbella Harris, 1937; politico Harris, 1937; ruga- tina Harris, 1937; rugosa Lea, 1833; sabinaria Har- ris, 1937; sayi Lea, 1833; thyroidifera Harris, 1937.

PERU: paytensis Olsson, 1930; wiedeyi Olsson, 1930.

EUROPEAN PALEOCENE: selandica Koenen, 1885.

Subgenus Oxyacrum Cossmann, 1889

Type: Pleurotoma ohliterata Deshayes, 1834

Members of this subgenus are very similar to Eopleurotoma but have a tall, narrowly conical protoconch of 4M smooth whorls, plus a whorl of brephic axials. It occurs in the Eocene of Europe and England.

Synonymy —

1889 Oxyacrum Cossmann, Ann. Soc. Malac. Belgique 24, p. 274. Type by original designation: Pleurotoma ohliterata Deshayes, 1834; 1896, Cossmann, Essais de Paleoconch Comp. 2, p. 82.

Characteristic Species — E. constricta Edwards, 1861; contabulata Deshayes, inflexa Lamarck, 1804; lepta Edwards, 1861; ohliterata Deshayes, 1834.

Genus Eoturris Finlay & Marwick, 1937

Type: Turns complicata Suter, 1917 This is a New Zealand genus ranging from the Eocene (Bortonian) to the Upper Oligocene (Wai- takian). It belongs to the Gemmula series but has a considerably broader sinus, spreading above the peripheral keel on to the shoulder as in Eopleuro- toma from the European and North American Eocene. However, Eopleurotoma is distinct in hav- ing a paucispiral protoconch, a short, gently twisted neck and usually a distinct fasciole. The proto- conch in Eoturris is narrow, elongate-conic, with the axials restricted to the last quarter whorl, in- stead of being numerous and spread over the last two or three whorls as in Gemmula.

Synonymy —

1937 Eoturris Finlay and Marwick, New Zealand Geol. Survey Paleo. Bull. 15, p. 114. Type by original designation: Tunis complicatus Suter, 1917.

Eoturris complicata (Suter, 1917)

Range — New Zealand Bortonian, Eocene. Synonymy —

1917 Turris complicatus Suter, N. Z. Geol. Surv. Paleo. Bull. 5, p. 45, pi. 5, fig. 14.

1917 Surcula mordax Suter, N. Z. Geol. Surv. Paleo. Bull. 5, p. 51 (non Martin, 1915).

1942 Eoturris complicatus Suter, Powell, Bull. no. 2, Auck- land Inst. Mus., p. 46.

T types — The type locality is McCullough’s Bridge, South Canterbury, New Zealand. The holo- type is in the New Zealand Geological Survey, Wellington, New Zealand.

Eoturris multicincta (Marshall, 1917)

(PI. 218, fig. c)

Range — New Zealand Bortonian, Eocene. Synonymy —

1917 Turris multicinctus Marshall, Trans. N. Z. Inst. 49, p. 456, pi. 36, fig. 34.

1942 Eoturris multicinctus Marshall, Powell, Bull. no. 2, Auckland Inst. Mus., p. 46.

Types — The type locality is “Wangaloa,” prob- ably Castle Hill Shaft. The holotype is in the Otago University Museum, Dunedin, New Zealand.

Eoturris neglecta (Suter, 1917)

Range — New Zealand Bortonian, Eocene. Synonymy —

1917 Turris neglectus Suter, N.Z. Geol. Surv. Pal. Bull. 5, p. 46, pi. 6, fig. 1.

1924 Turris insensus Finlay, Proc. Mai. Soc., 16, p. 103 (nom. nov. for neglectus Suter, 1917, non Pleuro- toma neglecta Reeve, 1842; new name unneces- sary).

[22 - 825]

284 Epalxis

A. W. B. Powell

Turridae

1942 Eoturris neglectus Suter, Powell, Bull, no. 2, Auck. Inst. Mus., p. 46.

Types — The type locality is N.Z.G.S. loc. 630 probably McCullough’s Bridge, South Canterbury. The holotype is in the N.Z. Geological Survey.

Eoturris uttleyi (Suter, 1917)

Range — New Zealand Waitakian, Upper Oligo- cene.

Synonymy —

1917 Turris uttleyi Suter, N. Z. Geol. Surv. Paleo. Bull. 5, p. 47, pi. 6, fig. 2.

1942 Eoturris uttleyi Suter, Powell, Bull. no. 2, Auckland Inst. Mus., p. 47.

Types — The type locality is Otiake, Waitaki Valley, North Otago, New Zealand.

Genus Epalxis Cossmann, 1889

Type: Pleurotoma crenulata Lamarck, 1803 This European Eocene genus was compared with Bathytoma by its author, and, later, Glibert ( 1960 ) preserved the association by making Bathytoma a subgenus of Epalxis. Nevertheless, Epalxis seems

to fall more naturally into the series of Eopleuro- toma, Eoturris and Oxyacrum. The genus Bathy- toma by reason of its characteristic columellar cal- losity seems to have more in common with the Borsoniinae. Also the radula of Bathytoma and Micantapex do not have the characteristic “wish- bone” marginals of the typical Turrinae but slender awl-shaped ones of toxoglossate style. Epalxis closely resembles Eopleurotoma in adult features and in having a paucispiral protoconch. The chief diagnostic features are the very broadly open anal sinus which occupies most of the spire whorls from suture to suture, and its broadly rounded apex which is wider than the broadly rounded peri- phery and which bears strong laterally compressed crescentic nodules.

Synonymy —

1889 Epalxis Cossmann, Ann. Soc. Malac. Belgique 24, p. 254. Type by original designation: Pleurotoma cre- nulata Lamarck, 1803; 1896 Cossmann, Essais de Paleoconch. Comp., 2, p. 103.

Characteristic Species — E. bilirata Boury, 1899; crenulata Lamarck, 1803; lemoinei Boury, 1899; multigyrata Deshayes, 1865; ventricosa Lamarck, 1804. '

March 31, 1964

INDO-PACIFIC MOLLUSCA, vol. 1, no. 5

Lucerapex 285

Genus Lucerapex Iredale, 1936

Type: Pleurotoma casearia Hedley & Petterd, 1906

Members of this genus are rather small elongate- fusiform shells, with a very tall spire and moder- ately long canal, of light build, and rather simple sculpture of a peripheral row of scale-like to tuber- cular nodes on an angulation or a raised keel. The sinus is peripheral, broadly open, rather shallow, U-shaped and defined over the whole shell in the form of distinct axial growth lines. The protoconch is small, smooth, paucispiral and globular, ending in up to a quarter whorl of protractively arcuate thin axials.

This genus has a wide Indian Ocean distribu- tion in deep water extending from the Gulf of Aden down the East African Coast, through Indonesian waters to the Philippines, and then southward to off the coast of New South Wales. It is represented also in the Quaternary of Timor and the Miocene of South Australia.

This genus seems to be most nearly related to Fusiturris which has a Recent range extending from the Mediterranean to West Africa and has European fossil relatives reaching back to as early as the Paleocene.

Both Fusiturris and Lucerapex are of elongate- fusiform shape, have the same peripheral nodula- tion, as well as a rather shallow, broadly open, U-shaped sinus which is defined over all post-em- bryonic whorls by crisp axial growth lines. The protoconch in Fusiturris is, however, narrowly conic and of about three whorls, and the shell is, for the most part, larger and of more solid build.

It is confidently assumed that Lucerapex is an Indo-Pacific product developed from Fusiturris, in isolation, after the closing of that great equatorial waterway of the past, the Tethys Sea.

Synonymy —

1936 Lucerapex Iredale, Records Australian Mus. 19, p. 337. Type by original designation: Pleurotoma casearia Hedley and Petterd, 1906.

Lucerapex casearia ( Hedley & Petterd, 1906 )

(PI. 219)

Range — Known only from 300 fathoms and 800 fathoms off Sydney, New South Wales, Australia.

Description — Shell small, 13 mm. (M inch) in height, thin, elongate-fusiform, with a tall spire and a long, slightly flexed, spout-like anterior canal.

Protoconch paucispiral with glassy, rounded, smooth whorls. Spire whorls prominently angled at about the lower third by a narrowly rounded keel, produced into regular blunt tubercles, 18 on the penultimate whorl. Sinus on the peripheral keel, of moderate depth, narrowly rounded at the base of a broad U-shaped area. Surface smooth, except for fine axial growth lines which diverge acutely both above and below the keel. Colour varying from pearl-grey to pale-orange (probably due to staining).

Measurements (mm.) — height width

13.0 5.0 holotype

Synonymy —

1906 Pleurotoma casearia Hedley & Petterd, Rec. Aust. Mus. 6, p. 220, pi. 37, fig. 5.

1936 Lucerapex casearia H. & P., Iredale, Rec. Aust. Mus. 19(5), p. 320.

1954 Lucerapex casearia H. & P., Laseron, The N. S. W. Turridae, Roy. Zool. Soc. N. S. W. Handb., p. 8, pi. 1, fig. 15.

Types — The holotype is in the Australian Mu- seum, Sydney.

Records — NEW SOUTH WALES: 300 fms. (type lo- cality ) and 800 fms. off Sydney.

Lucerapex casearia subspecies regilla Iredale, 1936

Remarks — Iredale separated shells from 110 fathoms off Sydney, stating that they were much larger than typical 300 fathom shells, and exhibited wavy lines below the periphery, a feature absent in the typical species. This may be merely an eco- logic form.

Plate 219. Lucerapex casearia (Hedley and Petterd). Hol- otype. 300 fms., off New South Wales, Australia (from Hed- ley and Petterd, 1906, pi. 35, fig. 5).

[22 - 837]

286 Lucerapex

A. W. B. Powell

Turridae

Measurements (mm.) —

height width

21 - holotype

Synonymy —

1936 Lucerapex casearia regilla Iredale, Rec. Aust. Mus. 19(5), p. 320.

Types — The holotype is in the Australian Mu- seum, Sydney.

Records — NEW SOUTH WALES: 110 fms. off Sydney (type locality).

Lucerapex denticulata (Thiele, 1925)

(PI. 220, fig. 2)

Range — Off Somaliland, East Africa and the Gulf of Aden, 732 to 1270 metres.

Remarks — This species is characterised by the simplicity of its sculpture which consists of a row of stout pointed tubercles on a sharp peripheral angle set just below the middle whorl height. The rest of the shell is smooth. It is closer to carola Thiele, 1925, than to the type of the genus, casearia Hedley & Petterd, 1906, from which it differs in having fewer tubercles on the peripheral keel.

There is a prior Pleurotoma (Mangelia) denti- culata Smith, 1884, but Thiele’s species is not re- named since it is doubtfully distinct from carola of the same author.

Measurements (mm.) —

height width

19.2 + 7.0 Gulf of Aden, 1270 metres

13.0 4.5 holotype

Plate 220. Fig. 1, Lucerapex carola (Thiele). Fig. 2, L. denticulata (Thiele). Both holotypes. Both deep-water, off East Africa (from Thiele, 1925, pi. 24). Figs. 3, 4, Lucer- apex molengraaffi (Tesch). Pliocene of Timor Id., Indonesia (from Tesch, 1915, pi. 77).

Synonymy —

1925 Pleurotoma denticulata Thiele, Wissenschaft. Ergebn. Deutschen Tiefsee-Exped. 17, Gastr. 2, p. 216, pi. 24, fig. 2.

Records — SOMALILAND: 1° 49' N„ 45° 29.5' E„ 1134 metres (type locality). GULF OF ADEN: 14° 36' 06" N., 51° 00’ 18” E„ 1270 metres, Sta. 184; 12° 04' 06" N., 50° 38' 36" E., 732 metres, Sta. 176 (John Murray Exped., Brit. Mus.).

Lucerapex carola (Thiele, 1925)

(PI. 220, fig. 1)

Range — Off Somaliland, East Africa, 693 metres.

Remarks — This species is known to the writer only from the original figure and description. It seems to be very closely allied to, if not identical with, the same author’s denticulata. From the fig- ure, the only apparent difference observed is in the form of the peripheral tubercles, which are rounded in carola but square-cut in denticulata. However, this small difference in sculpture could be resultant from the fact that carola appears to be not fully grown.

One refrains from uniting carola and denticulata without seeing the type material, for there may be matters of minor sculpture and texture that are not revealed in either the descriptions or the fig- ures.

Measurements (mm.) —

height width

9.0 3.2 holotype

Synonymy —

1925 Pleurotoma carola Thiele, Wissenschaft. Ergebn. Deutschen Tiefsee-Exped. “Valdivia” 17, Gastr. 2, p. 216, pi. 24, fig. 3.

Lucerapex adenica new species (PI. 221, fig. 3)

Range — Gulf of Aden, 274 to 1080 metres.

Remarks — This species has a more slender spire than molengraaffi, larger peripheral nodules than in any of the other species of the genus, an unusual feature in the presence of weak subsutural gem- mules, and numerous weak spirals on the base and anterior end. The peripheral nodules spread across and beyond a broadly rounded peripheral keel. In molengraaffi the nodules are restricted to a nar- row, sharp-edged, peripheral keel, and to a less degree this is the case with casearia. In denticulata and carola the nodules are in the form of pointed tubercles on a simple peripheral angle.

Description — Shell 25 to 32 mm. ( 1 to 1& inches) in height, narrowly fusiform, with a very tall slender spire, and a moderately long slightly twisted anterior canal. Spire 1.3 times the height

March 31, 1964

INDO-PACIFIC MOLLUSCA, vol. 1, no. 5

Lucerapex 287

of the aperture plus canal; spire angle 20°. Whorls 11.5, including a small paucispiral protoconch of 1.5 whorls, tip asymmetric and inrolled, last % whorl of arcuate, brephic, axial threads. Spire whorls with a massive, rounded, median peripheral keel which occupies more than one-third of the whorl height; sculptured with heavy crescentic axials, 15 on the penultimate whorl. A row of weak subsutural gem- mules is present. Base with 4 widely spaced, weak spirals on the upper part of the base, with only the uppermost emergent over the spire whorls; re- mainder of base with linear spaced, weak spirals. Sinus peripheral, broadly U-shaped, its scar clearly marked over all the post-nuclear whorls by strong axial growth lines. Colour pure white.

Measurements (mm.) —

height width

30.8 8.0 holotype

29.7 8.0

23.7 7.0

Types — The holotype is in the British Museum (Natural History).

Records — GULF OF ADEN: 13° 05' 36" N., 46° 24' 42" E., 1022 metres, Sta. 34 (type locality); 13° 06' 12" N., 46° 24' 30" E., 1061-1080 metres, Sta. 193; 13° 46' 30" N., 47° 48' 54" E., 274 metres, Sta. 191 (John Murray Exped., Brit. Mus. ).

Lucerapex indagatoris (Finlay, 1927)

(PI. 221, figs. 4, 5)

Range — Off South India, 360 to 430 fathoms. Remarks — This species is difficult to place ge- nerically without having seen specimens. It would seem, however, on the basis of the published de- scription and figures, to have more claim for in- clusion in Lucerapex than in Gemmula. The nature of the peripheral nodulation and in particular the subsutural series of weak gemmules indicate a shell fairly close to adenica new species, but differ- ing chiefly in the low-set position of the peripheral carina. Unfortunately, the apical whorls are un- known.

Measurements (mm.) —

height width

36.0 10.0

Synonymy —

1899 Pleurotoma optata E. A. Smith, Ann. Mag. Nat. Hist., Ser. 7, 4, p. 238.

1909 Pleurotoma optata E. A. Smith, Annandale & Stewart, Illust. Zool. Investigator, Moll., pi. 9, figs. 1, la.

1927 Gemmula indagatoris Finlay, Trans. N. Z. Inst. 57, p. 517, nom. nov. for PI. optata Smith, 1899 (non Pi. optata Harris, 1897).

Types — The type is in the Indian Museum, Cal- cutta.

Records — INDIA : off the south coast, 430 fms. ( type locality); off Travancore, 360 fms. (E. A. Smith).

Lucerapex molengraaffi (Tesch, 1915)

(PI. 220, figs. 3, 4; pi. 221, figs. 1, 2)

Range — Quaternary of Timor and the Recent of Borneo, Celebes and the Philippines, 254 to 559 fathoms.

Remarks — This species which was described from the Quaternary of Timor is identical with Re- cent deep-water material from “Albatross” dredg- ings taken in the vicinity of Borneo, Celebes and the Philippines.

From the other species of the genus it differs in having a square-cut, peripheral flange bearing the nodules, which later become scale-like, and in having distinct spiral threads on the base.

Description — Adult shell 30 to 33 mm. ( about 1 inch) in height, elongate-fusiform with a tall spire and a long, flexed canal with a spout-like termina- tion; light build; and prominently and medially carinated by a square-cut raised flange bearing regular, closely spaced, squarish nodules, 15 or 16 on the penultimate whorl, but becoming crowded and scale-like over the last half-whorl. Spire slightly greater than the height of the aperture plus canal; spire angle 20° to 22°. Whorls 10, including a small, smooth, globular protoconch of two whorls.

Plate 221. Figs. 1, 2, Lucerapex molengraaffi (Tesch). 254 fms., off west Siquijor Id., Philippines. 32 mm. Fig. 3, Lu- cerapex adenica new species. Holotype. 1,022 meters, Gulf of Aden. 30.8 mm. Figs. 4, 5, Lucerapex indagatoris ( Fin- lay). Holotype of Pleurotoma optata E. A. Smith, 1899, non Harris, 1897. 430 fms., off south coast of India (from An- nandale and Stewart, 1909, pi. 9, figs. 1, la).

[22 - 839]

288 Lucerapex

A. W. B. Powell

Turridae

Sinus wide, shallow, U-shaped, and on the periph- eral carina. Two, sometimes three, weak wavy spiral threads encircle the upper base. Axial growth lines strong, especially on the base.

Measurements (mm.) -

height	width
30.25	10.0
29.0	8.2	Borneo, 305 fms.
28.5	8.7	Noil Tobe, Timor, Pliocene
25.0	8.0
Synonymy	-

1915 Pleurotoma (s. str. ) Molengraaffi Tesch, Palaont. von Timor 5(9), Jungtert, und Quartare, Moll, von Timor, p. 28, pi. 77, figs. 54-56.

Records — TIMOR: Pliocene (type locality); Noil Tobe (Pliocene) (Brit. Mus.). BORNEO: off Silungan Id. 305 fms., Sta. 5592; off Sipaden Id., Sibuko Bay, 347 fms., Sta. 5586. CELEBES: Buton Strait, 559 fms., Sta. 5648. PHIL- IPPINES: west of Siquijor, 254 fms., Sta. 5537 (Albatross USNM ).

Lucerapex murrayana ( Pritchard, 1904 )

Range — Balcombian, Miocene, in Victoria, Aus- tralia.

Remarks — No specimens of this species have been examined by the writer but the description and not very distinct figure suggest that it belong to Lucerapex.

Description ( from the original) — “Shell small to medium size with a rather blunt apex, slender elongate spire and a body whorl shorter than the spire. Embryo consisting of about 2% whorls, blunt apically, smooth and inclined to be angled medially after about the first half turn; this portion is also rather more tumid and protrudes over the remain- der of the embryonic whorls; whorls strongly nodosely keeled about the middle of each whorl and forming a well marked shoulder on the body whorl. The keel marks the position of the sinus and is very regularly and acutely nodulose, nodules about 18 to 20 on the penultimate and body whorls. Outer lip thin and sharp, with a broad deep sinus at the shoulder.”

Measurements (mm.) —

height width

28.0 9.0 holotype

Synonymy —

1904 Pleurotoma murrayana Pritchard, Proc. Roy. Soc. Vic- toria 17, p. 335, pi. 19, fig. 10.

1944 Lophiotoma murrayana Pritchard, Powell, Rec. Auck. Inst. Mus. 3(1), p. 9.

Records — AUSTRALIA: River Murray Cliffs near Mor- gan, Victoria (Balcombian, Miocene).

Plate 222. Geographical distribution of Lucerapex casearia (Thiele), L. denticulata (Thiele), L. indagatoris (Finlay),

(Hedley and Petterd), L. molengraaffi (Tesch), L. carola L. adenica new species, and L. murrayana (Pritchard).

(

[22 - 840

March 31, 1964

INDO-PACIFIC MOLLUSCA, vol. 1, no. 5

Ptychosyrinx 289

Genus Ptychosyrinx Thiele, 1925

Type: Pleurotoma (Subulata) bisinuata Martens, 1901

This is a deep-water genus strongly resembling Gemmula, except for the sinus which, although situated on the gemmate peripheral carina, is not deep and narrow, but broadly V to U shaped at its apex and not very deep. Unfortunately the apical whorls, except in the type species, are eroded away in all examples so far examined. Thiele ( 1925, pi. 23, fig. 4) showed the protoconch of bisinuata as narrowly conic with four axially costate whorls.

The radula in the type species, bisinuata, figured by Thiele ( 1929, p. 359, fig. 436 ) is unlike that of Gemmula, in that a large rectangular-based central tooth is present in addition to marginals, similar to the “wishbone” type of typical Gemmula. It could be considered prototypic of the Turrinae, compar- able with the situation in the Clavinae where cer- tain genera, Drillia, Clavus and Spirotropis for example, preserve a prototypic radula of central, lateral and marginal teeth. Most of the members of the subfamily Clavinae have a toxoglossate type of dentition consisting only of a pair of slender mar- ginals.

The type species, bisinuata, as the name indi- cates, exhibits a spout-like projection of the lower outer lip as well as an anal siphonal notch. This same feature occurs in an example of timorensis teschi new subspecies from deep water in the Gulf of Tomini, Celebes, but it is not a constant feature of that subspecies. It results from a fusing and flut- ing of the two main basal cords.

The genus Ptychosyrinx is evidently of very wide distribution in the deep-sea basins, for apart from the East African and Indonesian occurrences it has been recorded recently from deep water off Ber- muda, as Bathybermudia carynae Haas, 1949. Haas’ genus was proposed under the mistaken impression that the bisinuate character of the type specimen of Ptychosyrinx was a diagnostic character of that genus.

Since this is now shown not to be the case, Bathybermudia falls as a synonym of Ptychosyrinx.

Synonymy —

1925 Ptychosyrinx Thiele, Wissenschaft, Ergebnisse Deutschen Tiefsee-Exped. 17(2), p. 210. Type by original designation (fide Powell, 1942, p. 20): Pleurotoma bisinuata Martens, 1901.

1949 Bathybermudia Haas, Bull. Inst. Catalana d’Hist. Nat., 37, p. 70. Type by monotypy: B. carynae Haas, 1949.

Ptychosyrinx bisinuata (Martens, 1901)

( PI. 223, figs. 1, 2 )

Range — Off East Africa, Somaliland to Zanzibar, 818 to 1362 metres.

Remarks — Shell tall-spired but with a rather short and recurved anterior canal. Protoconch polygyrate, narrowly conic, with 4 densely, axially costate whorls. Spire whorls sculptured with a broadly rounded, median carina studded with prominent, laterally compressed nodules. Spiral sculpture consisting of 2 or 3 threads between the upper suture and the peripheral carina, 2 or 3 threads below the carina and 3 strong, smooth cords on the upper part of the base, followed by about 9 threads over the neck and anterior canal. The uppermost of the 3 main basal spirals is emer- gent over the last whorl. The bisinuate nature of the outer lip exhibited by the type specimen has been shown above to be an abnormality. Colour pale yellowish.

Plate 223. Figs. 1, 2, Ptychosyrinx bisinuata Martens. Holo- type. 1,134 meters, off East Africa. Figs. 3, 4, Ptychosyrinx timorensis (Tesch). Pliocene of Timor Id., Indonesia (from Tesch, 1915, pi. 77, figs. 53a, b). Figs. 5, 6, Ptychosyrinx timorensis teschi new subspecies. 415 fms., northwest of Sipadan Id., Borneo. Fig. 5, holotype, 47 mm. Figs. 7, 8, Ptychosyrinx truncata (Schepman). 2,798 meters, Banda Sea, Indonesia (from Schepman, 1913, pi. 26, fig. 1). 19 mm.

[22 - 851]

290 Ptychosyrinx

A. W. B. Powell

Turridae

Measurements (mm.) —
height	width
33.0	11.5
30.0	9.0

Synonymy —

1901 Pleurotoma (Subulata) bisinuata Martens, Sitzungs- berichte Gesellsch. nat. Freunde, Berlin, p. 17.

1903 Drillia (Subulata) bisinuata Martens, Gast. Deutsch.

Tiefsee-Exped., 1898-1899, 7, p. 82, pi. 1, fig. 8. 1925 Ptychosyrinx bisinuata Martens, Thiele, Wissenschaft. Ergebn. Deutschen Tiefsee-Exped. 17, Gastr. 2, p. 210.

Records — EAST AFRICA: 1° 49' N„ 45° 29' E., 1134 metres and 2° 58' N., 46° 50' E., 1362 metres; off coast of Somaliland, 6° 18' N., 49° 32' E., 1079 metres; Pemba Channel, Zanzibar, 818 metres (Martens).

Ptychosyrinx lobata (Sowerby, 1903)

(PI. 224)

Range — Off Natal and South Africa, 300 to 900 fathoms.

Remarks — Superficially this shell looks identical with Ptychosyrinx bisinuata (Martens, 1901), a deep water East African species. Tomlin added a remark to the type tablet of lobata in the British Museum, claiming that it is synonymous with Mar- ten’s species, but one hesitates to accept this opin- ion, in view of Barnard’s (1958) remarks on the very different nature of the radula in topotypes of lobata.

Plate 224. Ptychosyrinx lobata ( Sowerby, 1903 ) . Holotype from the British Museum (Natural History). 440 fms., off Cape Natal, South Africa. 31 mm.

Thiele (1929, p. 359) figured a radula for bisinuata, which has a very large unicuspid rec- tangular-based central in addition to a pair of mar- ginals. However, in all the lobata specimens ex- amined by Barnard, the central tooth of the radula was found to be missing. The radula of an East London specimen was described as “with c. 75 pairs of teeth, no central plate, laterals broadly cuneiform, one margin sharply angular (in edge view) no appendage” (his “lateral” tooth should read “marginal”).

Unfortunately none of Barnard’s specimens had the protoconch intact, nor was it preserved in Sow- erby’s holotype. The operculum is described by Barnard as oval, with an apical nucleus, again dis- cordant with Thiele’s “eiformig konzentrisch.”

In the face of the above evidence lobata and bisinuata cannot be considered conspecific, proba- bly not even congeneric, but for the present, lobata is included in Ptychosyrinx until more is known concerning its type species.