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. The thought arises that Thiele could have mixed up his radula preparations, for the type of radula he figured for bisinuata is of similar style to that of both Aforia magnifica Strebel and Turridrupa jubata Hinds, both of which have a large-based, unicuspid central tooth. Measurements (mm.) — height width 31 11 Synonymy — 1903 Pleurotoma (Surcula) lobata Sowerby, Marine Invest. South Africa 2, p. 213, pi. 4, fig. 9. 1958 Tunis lobata Sowerby, Barnard, Ann. South African Mus. 44, p. 107; 1963, Ann. South African Mus. 46, p. 419. Types — The type is in the British Museum (Nat- ural History). Records — SOUTH AFRICA: Cape Natal, N. by E., 24 mi., 440 fms. (type locality); Buffalo River, N. 15 mi., 310 fms. ( Sowerby ) ; off Cape Point, 380-900 fms. ( Barnard ) . Ptychosyrinx timorensis subspecies timorensis (Tesch, 1915) (PI. 223, figs. 3, 4) Range — Pliocene of Timor Island, Indonesia. Remarks — The known examples of this species also lack the apical whorls but the adult shell, judged from Tesch’s figures, closely resembles a Becent Indonesian deep-water shell described be- low. The Timor fossil is more slender than either bisinuata or the Becent Indonesian subspecies and the peripheral sculpture is of rounded rather than the laterally compressed nodules of bisinuata. [22 - 852] March 31, 1964 INDO-PACIFIC MOLLUSCA, vol. 1, no. 5 Ptychosyrinx 291 Plate 225. Geographical distribution of Ptychosyrinx bisinu- ata (Martens), P. lobata (Sowerby), P. timorensis timor- ensis (Tesch), and P. timorensis teschi new subspecies. Measurements (mm.) — height width 32-54 - Tesch 53.0 17.5 from Tesch’s fig. 53b Synonymy — 1915 Pleurotoma (s. str. ) timorensis Tesch, Palaeont. von Timor, p. 27, pi. 77(5), figs. 52, 53. Types — The type locality is Timor Island, Plio- cene. Ptychosyrinx timorensis subspecies teschi new subspecies (PI. 223, figs. 5, 6) Range — Indonesia, Celebes, Moluccas and Bor- neo, 347 to 559 fms. Remarks — This subspecies appears to be the di- rect Recent descendant of the Pliocene timorensis. It differs from that species mainly in being propor- tionately broader, resulting in a greater spire angle, in having a shorter spire and a more capacious body whorl. Description — Adult shell 46 to 53 mm. ( about 2 inches) in height, with a tall spire but a short and flexed anterior canal. Colour uniformly dull white. Whorls 9 to 10/2, exclusive of the embryonic whorls which are missing in all specimens examined. Spire whorls sculptured with a strong subsutural cord and a weaker spiral above it, the latter forming the margin of the suture. There are 2 or 3 threads be- tween the subsutural cord and the peripheral Ca- rina. The latter is studded with closely-spaced, prominent, squarish nodules, 18 to 22 per whorl, obscurely overridden by 3 flat-topped linear-spaced spirals. Two spiral threads and one primary cord are between the peripheral carina and the lower suture. Base with 2 or 3 strong spiral cords above, and 8 or 9 spiral threads below on the neck and an- terior canal. Spire about 1.3 times the height of the aperture plus canal; spire angle 30° to 33°. Sinus broadly V-shaped, relatively shallow, with a broadly rounded apex at the peripheral sinus an- gulation. Operculum leaf-shaped with an apical nucleus. In timorensis timorensis the spire angle is 24° to 25°, and the height of the spire is almost twice that of the aperture plus canal. An example from the Gulf of Tomini, Celebes (Albatross Sta. 5601) exhibits the “bisinuate” ab- normality of a second sinuation of the outer lip caused by the fusing and fluting of the two main basal cords. Measurements (mm.) — height width 53.7 17.0 Albatross Sta. 5586; Borneo 53.0 18.7 Albatross Sta. 5648; Celebes 47.0 17.0 holotype, Albatross Sta. 5587 Types — The holotype is in the United States National Museum, Washington, D. C. The type locality is at the Albatross Station 5587, N.W. of Sipadan Id., Borneo, in 415 fms. USNM no. 239068 [22 - 853] 292 Ptychosyrinx A. W. B. Powell T urridae Records — CELEBES: off North Id., Buton Strait, 519 fms., green mud (Albatross Sta. 5647); off North Id., Buton Strait, 559 fms., green mud (Albatross Sta. 5648); Gulf of Boni, 15 mi. S.E. of Tg Olang, 484 fms., grey mud (Albatross Sta. 5656); Gulf of Tomini, (Albatross Sta. 5601). MOLUCCAS: March Id., Molucca Pass, 417 fms., grey mud (Albatross Sta. 5618). BORNEO: E. of Sipadan Id., Sibuko Bay, 347 fms., green mud (Albatross Sta. 5586); N.W. of Sipadan Id., 415 fms., green mud, sand and coral (holotype, Albatross Sta. 5587), (all USNM). Ptychosyrinx truncata ( Schepman, 1913 ) (PI. 223, figs. 7, 8) Range — Known only from the Banda Sea, In- donesia. Remarks — All four specimens mentioned by Schepman are lacking their upper whorls, and without knowledge of the protoconch the generic location of this species is conjectural. The broadly open, shallow peripheral sinus and rather short an- terior canal suggest location in Ptychosyrinx rather than in Gemmula. Sculpture consisting of strongly- waved, axial, fold-like ribs becoming weaker over the last whorl, and crossed throughout by numer- ous spiral threads. Measurements (mm.) — height width 19+ 8 Synonymy — 1913 Pleurotoma (Gemmula) truncata Schepman, Siboga Exped., Monog. 49(1 )e, p. 404, pi. 26, fig. 1. Records — INDONESIA: Banda Sea, 2798 metres (type locality ) . [22 - 854] March 31, 1964 INDO-PACIFIC MOLLUSCA, vol. 1, no. 5 Ptychosyrinx 293 Subgenus Kuroshioturris Shuto, 1961 Type: Gemmula (Kuroshioturris) hyugaensis Shuto, 1961 When Shuto proposed this subgenus he associ- ated it with Gemmula because of its similar sculp- ture and sinus. However, the smooth, globose paucispiral protoconch of Kuroshioturris suggests that it is more closely related to Ptychosyrinx, rather than to Gemmula which has a multispiral, axially costate protoconch. The species from the Upper Miocene-Lower Pliocene of Japan described as Gemmula (Ptycho- syrinx) nipponicus Shuto, 1961, seems to fit better into his subgenus Kuroshioturris. The protoconch is described as rather large and composed of a de- pressed, rounded and smooth first volution and a convex and inflated second one, which is smooth except for the last quarter whorl with brephic axials. His figure of the protoconch, however, gives the impression that the embryo terminates abruptly with the commencement of adult sculpture. Synonymy — 1961 Kuroshioturris Shuto, Mem. Faculty Sci., Kyushu Univ., Series D, Geol. 11(2), p. 83. Subgenus of Gemmula; type by original designation: G. (K.) hyugaensis Shuto, 1961. Ptychosyrinx nipponica ( Shuto, 1961 ) (PI. 226, figs. 3, 4) Range — Japan, near Yamaji (Upper Miocene- Lowest Pliocene). Remarks — This shell closely resembles typical Ptychosyrinx in its adult facies, recalling timorensis teschi new subspecies, but with greater simplicity of sculpture, having a peripheral row of stout but more distant nodules and 2 very prominent spiral cords on the upper base. The only other spiral sculpture consists of weak threads, several sub- margining the suture and a number on the neck and anterior canal. There is a tendency toward “bisinuation” of the outer lip at the termination of the upper basal cords. Measurements (mm.) — height width 12.9 7.2 12.2 7.5 11.1 6.3 Synonymy — 1961 Gemmula (Ptychosyrinx) nipponicus Shuto, Mem. Fac. Sci. Kyushu Univ. Ser. D, Geol. 11(2), p. 81, pi. 3, figs. 7, 8, 13, 19; pi. 8 (not 7), fig. 14, text figs. 3, 4. Records — JAPAN : entrance cutting to tunnel, 400 metres north of Yamaji, Mino mura, Koyu gun, Miyazaki Prefecture ( Kawabaru member, lower Takanabe member, lower Upper Miocene to Lowest Pliocene) (type locality). Ptychosyrinx hyugaensis ( Shuto, 1961 ) (PI. 226, figs. 1, 2) Range — Japan, Miyazaki Prefecture (Lowest Pliocene ) . Remarks — The early whorls have a peripheral carina studded with rounded nodules but these be- come obsolete over the body whorl. The basal cords are also obsolete to subobsolete but are represented by an angulation. A blunt biangulation of the body whorl results. Plate 226. Figs, 1, 2, Ptychosyrinx (Kuroshioturris) hyuga- ensis (Shuto). Lower Pliocene of Miyazaki Pref., Japan (from Shuto, 1961, pi. 3, figs. 2, 3). Figs. 3, 4, Ptychosyrinx nipponica (Shuto). Lowest Pliocene of Miyazaki Pref., Ja- pan (from Shuto, 1961, pi. 3, figs. 7, 8). [22 - 865] 294 Ptychosyrinx A. W. B. Powell Turridae Measurements (mm.) — height width 13.5 7.8 12.9 7.6 Synonymy — 1961 Gemmula (Kuroshioturris) hyugaensis Shuto, Mem. Fac. Sci. Kyushu Univ. Ser. D, Geol. 11(2), p. 83, pi. 3, figs. 2, 3, 4; text figs. 3, 4. Records — JAPAN : roadside cutting at Hagenoshita, Miya- zaki Prefecture (lower part of Takanabe member, Lowest Pliocene), (type locality). Ptychosyrinx totomiensis (Makiyama, 1931) Range — Japan, Kakegawa and Miyazaki groups (Lower Pliocene). Remarks — This species closely resembles a Pty- chosyrinx timorensis teschi of optimum sculptural development. The subsutural ridge is strong and sharply projecting, the peripheral nodules are strong and closely spaced and all of the upper basal spirals are strong cords. Except for the relatively large, paucispiral protoconch, without brephic ax- ials, the species would fit into Ptychosyrinx typical. Measurements (mm.) — height width 12.0 6.7 Hagenoshita Synonymy — 1931 Tunis (Gemmula) totomiensis Makiyama, Mem. Coll. Sci. Kyoto Imp. Univ. Ser. B, 7(1), p. 46, pi. 1, figs. 10, 11. 1961 Gemmula (Kuroshioturris) totomiensis Shuto, Mem. Fac. Sci. Kyushu Univ., Ser. D, Geol. 11(2), p. 84, pi. 6, figs. 1, 2, 3, 10; text figs. 3, 4. Records — JAPAN: roadside cutting at Hagenoshita, Miyazake Prefecture (lower part of Takanabe member, Lowest Pliocene) (type locality). Ptychosyrinx asukana (Yokoyama, 1926) Range — Japan (Pliocene) and Okinawa (Mio- cene-Pliocene ) . Remarks — MacNeil ( 1960, p. 103 ) writes of this species: “There seems to be little doubt that the specimen Yokoyama referred to this species in 1928 is a true Gemmula. However, doubt has been raised over the identity of the species described under this name in 1926. Hatai and Nisiyama ( 1952, p. 198) place it in Clavatula. Yokoyama’s figure is poor but nevertheless he says in the description that ‘the angle is the place where the old sinus band is present, on which the periodic ends of the sinus remain as tubercles.’ If this description is ac- curate it would almost certainly place the shell in the Turrinae and probably in the genus Gemmula.” However, MacNeil overlooked that Yokoyama stated in his original description “Whorls about seven of which one and a half are nuclear, smooth and rounded.” This rules out the species as a Gem- mula and also renders inapplicable MacNeil’s com- parison of the species with my longwoodensis (Powell, 1942) from the Duntroonian Upper Oligo- cene of Southland, New Zealand, not Australia, as stated by him. Description — Adult shell about 40 mm. ( about VA inches) in height, slender and rather simple in sculptural characters. Angulate below the middle of whorl height, the periphery not carinated or flanged but studded with round to slightly pointed tubercles. In the Okinawan specimen there is a subsutural fold traversed by 3 weak spiral cords and 3 rather massive plain spirals encircle the up- per part of the base, after which the spiral cords become weaker and more closely spaced. The sur- face, including the peripheral angle and its tuber- cles, as well as the interstices of the 3 main basal spirals, is crowded with spiral threads. This species is likely to prove rather closely allied to nipponica Shuto, 1961. The measurements for the holotype below are estimated. Measurements (mm.) — height width 40.0 6.7 holotype 12.5 4.9 Okinawa Synonymy — 1926 PDrillia asukana Yokoyama, Journ. Tokyo. Imp. Univ. Fac. Sci., Sec. 2, 1(9), p. 331, pi. 38, fig. 18 (not Pleurotoma asukana Yokoyama, 1928). 1960 Gemmula aff. asukana Yokoyama, MacNeil, U. S. Geol. Surv. Prof. Paper 339, p. 103, pi. 5, fig. 12. Records — JAPAN : Asuka ( Satsuka beds. Pliocene ) (type locality). OKINAWA: (Yonabaru clay member, Miocene). ( [22 - 866] March 31, 1964 INDO-PACIFIC MOLLUSCA, vol. 1, no. 5 Optoturris 295 Genus Optoturris Powell, 1944 Type: Pleurotoma optata Harris, 1897 This is an Australian Miocene genus which is also recorded from the Lower Pliocene of Japan. Pro- toconch paucispiral of 1/2 smooth whorls plus a half whorl of curved axials, the entire protoconch peg-like in shape but with the tip small and asym- metric. The sinus is broadly U-shaped, not very deep and extends over more than half the shoulder, with its lower edge situated at the weak periph- eral carina. The genus seems to have no living rep- resentatives. Synonymy — 1944 Optoturris Powell, Records Auckland Inst, and Mu- seum 3(1), p. 12. Type by original designation: Pleurotoma optata Harris, 1897. Optoturris optata (Harris, 1897) Range — Janjukian and Balcombian Miocene of Victoria, Australia. Description — Adult shell 15 to 17 mm. (less than /I inch) in height, elongate-fusiform, with a tall spire and a moderately long canal. Whorls slightly convex, except for a weak peripheral angulation at about one third of the whorl height. Sculpture of dense thread-like spirals crossed by numerous axial growth lines which thicken at the rest stages, espe- cially in the troughs of the sinus, imparting a rather dense comma-shaped sculptural pattern to the sinus area. Interior of outer lip deeply fluted within. Measurements (mm.) — height width 16.9 6.0 Balcombe Bay 16.5 6.0 holotype 15.0 5.3 Balcombe Bay Synonymy — 1897 Pleurotoma optata Harris, Cat. Tert. Moll. Brit. Mus. 1, p. 44, pi. 3, figs. 4, a, b. 1944 Optoturris optatus Harris, Powell, Rec. Auck. Inst. Mus. 3(1), p. 12. Types — The type is in the British Museum (Nat- ural History) in London, England. Records — AUSTRALIA: Hobson’s Bay, Victoria, “Eo- cene” = Balcombian, Miocene ( type locality ) ; Muddy Creek, lower beds, Victoria (Balcombian) (Auck. Mus.). Optoturris edita (Powell, 1944) ( PI. 227 ) Range — Balcombian Miocene of Victoria, Aus- tralia. Remarks — Related to optata but larger, 25 mm. ( 1 inch ) in height, narrower, with taller spire and longer canal. Sinus and sculpture similar to that of optata. Measurements (mm.) — height width 25.0 6.5 holotype Synonymy — 1944 Optoturris editus Powell, Rec. Auck. Inst. Museum 3(1), p. 12, pi. 7, fig. 3. Types — The type locality is Grice’s Creek, Vic- toria (Balcombian, Miocene). The holotype is in the Auckland Museum, New Zealand. Plate 227. Left fig., Optoturris edita Powell, Balcombian Miocene of Grice’s Creek, Victoria, Australia (from Powell, 1944, pi. 7, fig. 3). 25 mm. Right fig., Optoturris kyushuen- sis Shuto. Lower Pliocene of Hagenoshita, Japan (from Shuto, 1961, pi. 4, fig. 12). [22 - 871] 296 Optoturris A. W. B. Powell Tmridae Optoturris paracantha (Tenison-Woods, 1877) Range — Janjukian Miocene of Tasmania and Victoria, Australia. Remarks — Shell larger, broader and of more solid build than the other two Australian species; similarly sculptured but more strongly and with a broad deeply concave sulcus between the suture and the sinus area. Also, the thickened growth- lines, where they cross the sinus area, resolve into a regular gemmate series, reminiscent of Gemmula. Measurements (mm.) — height width 25.6 10.0 Table Cape, Tasmania Synonymy — 1877 Pleurotoma paracantha Tenison-Woods, Proc. Roy. Soc. Tasm., for 1876, p. 105. 1898 Bathytoma paracantha T.-Woods, Tate, Proc. Roy. Soc. N.S.W. 31, p. 398. 1944 Optoturris paracanthus T.-Woods, Powell, Rec. Auck. Inst. Mus. 3(1), p. 12 Records — AUSTRALIA: Table Cape, Tasmania (type locality); Torquay, Victoria (Janjukian, Miocene). Optoturris kyushuensis Shuto, 1961 (PI. 227) Range — Takanabe Lower Pliocene of Japan. Remarks — Shell small, 16.9 mm. (about M inch) in height. It appears from the description and fig- ure to be undoubtedly of the above genus, and closely resembles the Australian Balcombian edita, from which it is distinguished chiefly by both stronger primary spirals on the base and more defi- nite granules or gemmules on the sinus rib. Measurements (mm.) — height width 16.9 5.1 Synonymy — 1961 Optoturris kyushuensis Shuto, Mem. Fac. Sci. Kyushu Univ. Ser. D, Geol. 11(2), p. 75, pi. 4, fig. 12. Records — JAPAN: Hagenoshita, unconsolidated grey fine sandstone (Takanabe member, Lower Pliocene) (type lo- cality ) . [22 - 872] March 31, 1964 INDO-PACIFIC MOLLUSCA, vol. 1, no. 5 Epidirella 297 Genus Epidirella Iredale, 1931 Type: Pleurotoma xanthophaes Watson, 1886 This genus appears to be more nearly allied to Ptychosyrinx than to Gemmula, but examination of the radula will be necessary to determine its true relationship. In Gemmula the radula is of the same type as in Turns, that is, a pair of “wishbone- shaped” marginals, whereas in Ptychosyrinx Thiele it is with a large unicuspid central tooth in addition to modified “wishbone” marginals but no laterals. The sinus in Epidirella resembles that of Pty- chosyrinx in being a peripheral, shallow, broadly- open “V,” but the sculpture is prominently bicari- nate-gemmate; that of Ptychosyrinx is gemmate only on the peripheral keel. There is, in addition to the two gemmate keels, a closely spaced pair of subsutural cords and these also are gemmate, the sculptural effect on the spire whorls being an entire covering of gemmules. The protoconch is tall, with three whorls, the first smooth and narrowly dome-shaped, followed by weak, oblique axials, crossed on the last whorl by an incipient bicarination. Operculum leaf- shaped, with apical nucleus. The shell is white, covered by a light horny pe- riostracum, and this also is true of Ptychosyrinx. The known distribution of Epidirella is the east Australian Continental Shelf, Tasmania and New South Wales, but it is likely that the genus may be found to extend into the Indo-Pacific in deep water. A fossil species, sayceana Chapman, 1912, from the Kalimnan, Lower Pliocene of Victoria, is tentatively referred to this genus. Synonymy — 1931 Epidirella Iredale, Records Australian Mus. 18(4), p. 226. Type by original designation: Hemipleurotoma tasmanica May, 1911 = Pleurotoma xanthophaes Watson, 1886. Epidirella xanthophaes (Watson, 1886) (PI. 228) Range — Southeast Australia and Tasmania. Remarks — Hedley ( 1922, p. 231 ) referred this species to his genus Epideira and cited Hemipleu- rotoma tasmanica May, 1911, as a synonym. Iredale (1931, p. 225) could not reconcile May’s species with Watson’s description and figure of xantho- phaes, and in erecting a new genus, Epidirella, cited tasmanica as type. Laseron (1954, p. 7), ob- viously unaware of Iredale’s genus, provided an- other new genus, Austro gemmula, for May’s species and at the same time remarked that Watson’s spe- cies has never been recognized since its first re- cording. A photograph of the type of xanthophaes Wat- son, here reproduced, shows the sculpture to be stronger and more noticeably gemmate than it ap- pears in Watson’s figure, but not so pronounced as in most examples attributed to tasmanica that the writer has seen. Nevertheless, the strength of the sculpture seems to be rather variable, and so upon the available material separation from Watson’s species appears unwarranted. Measurements (mm.) — height width 26.5 9.5 Twofold Bay 25.0 — Crookhaven ( Laseron ) 24.0 8.4 holotype of xanthophaes 20.0 7.0 Disaster Bay Synonymy — 1886 Pleurotoma xanthophaes Watson, Challenger Zool. 15, p. 282, pi. 26, fig. 1. 1911 Hemipleurotoma tasmanica May, Proc. Roy. Soe. Tasm., 1910, p. 391, pi. 13, fig. 16. 1922 Epideira xanthophaes Watson, Hedley, Rec. Aust. Mus. 13(6), p. 231. 1931 Epidirella tasmanica May, Iredale, Rec. Aust. Mus. 18(4), p. 226. 1954 Austrogemmula tasmanica May, Laseron, N.S.W. Turridae, Handb. Roy. Zool. Soc. N.S.W., p. 7, pi. 1, figs. 8, 9. Plate 228. Epidirella xanthophaes (Watson). Holotype. 30-35 fms., off Port Jackson, New South Wales, Australia. 24.0 mm. [22 - 875] 298 Epidirella A. W. B. Powell Turridae Types — The holotype of Watson’s xanthophaes is in the British Museum (Natural History), Lon- don. Records — NEW SOUTH WALES: 30-35 fms., off Port Jackson ( type locality for xanthophaes ) ; Disaster Bay, 5 fms.; off Crookhaven, 30-35 fms. (Laseron, 1954); off Cape Everard, 70-80 fms. (Aust. Mus. ); off Twofold Bay, 5-8 fms. (T. Gerrard). TASMANIA: off Schouten Id., 40 fms. (type locality for tasmanica) . VICTORIA: Lakes Entrance, 20 fms. (Macpherson & Gabriel). Epidirella sayceana (Chapman, 1912) Range — Lower Pliocene of Victoria, Australia. Synonymy — 1912 Pleurotoma sayceana Chapman, Proc. Roy. Soc. Viet. 25(n.s.), p. 191, pi. 12, fig. 7. 1944 ? Epidirella sayceana Chapman, Powell, Rec. Auck. Inst. Mus. 3(1), p. 16. Records — VICTORIA: Lakes Entrance (type locality); Jemmy’s Point (Kalimnan, Lower Pliocene). March 31, 1964 INDO-PACIFIC MOLLUSCA, vol. 1, no. 5 Epidirona 299 Genus Epidirona Iredale, 1931 Type: Epidirona hedleyi Iredale, 1931 The genus Epidirona, despite its resemblance to the Clavinae, belongs in the Turrinae as shown both by the peripheral location of the sinus and by the radula which is a modified “wishbone” type. These shells have a tall acuminate spire but a trun- cated body whorl; the anterior canal is not pro- duced but ends in a broad shallow notch at the termination of a well-marked fasciole. The anal sinus is a moderately deep, narrowly rounded “U,” situated at the periphery. The protoconch is small and of two smooth turbinate whorls. The post-nu- clear sculpture is typically of fine spiral cords and threads, crossed obliquely by weak axials which are strengthened into elongate, laterally-compressed nodes over both the subsutural fold and the periph- eral carina. The outer lip is thin, without a varix, and there is no parietal tubercle. Interior of aper- ture spirally fluted. For radula, see pi. 229. The Recent geographical range of the genus is mostly southern Australia to Tasmania and New South Wales but there is evidence that it occurs much farther afield as instanced by E. A. Smith’s multiseriata which has been recorded from Ceylon, Persian Gulf and China Seas. Four Tertiary species are known from southern Australian horizons, extending from the Janjukian Miocene to the Adelaidean Lower mid-Pliocene. The type species of this genus, a well-known New South Wales shell, has been the subject of much confusion both at the generic and the specific level. When Hedley (1918, p. M79) proposed Epideira he cited Clavatula striata Gray, 1827, as type, but later (1922, p. 230, pi. 43, figs. 18-20) showed that he had in mind the New South Wales shell previously known as either striata Gray (but not Gray of 1827) or owenii Reeve, 1843. Iredale (1931, p. 225) rejected Epideira in con- nection with the striata of Hedley, stating that Gray’s description did not fit the New South Wales shell and that Gray’s species, at present indeter- minate, probably came from Western Australia. Watson (1886, p. 312) used Pleurotoma owenii (Gray ms.) Reeve, 1843, for the New South Wales shell, citing Clavatula striata Gray as a synonym, but Reeve’s species was stated to be from the East Coast of Africa. The specimen used by Reeve to illustrate his owenii certainly resembles the New South Wales shell but he based the name upon a specimen in Gray’s cabinet and remarked that it “is of unusually large size, at least one-third longer than that selected for illustration.” The types of both striata and owenii seem to be lost and this induced Iredale to clarify the situa- tion by providing a new genus and species name for the New South Wales shell. Synonymy — 1931 Epidirona Iredale, Records Australian Mus. 18(4), p. 225. Type by original designation: Epidirona hed- leyi Iredale, 1931. A full treatment of Epidirona is planned for a future paper, and for the present I am considering in detail only the sole, strictly Indo-Pacific species, multiseriata (E. A. Smith, 1877). The others, mainly from southeast Australia, are listed here alphabetically by species, but in their original ge- neric combinations. Epidirona multiseriata (E. A. Smith, 1877) (PI. 230, fig. 3) Range — Persian Gulf to the South China Sea. Remarks — This species closely resembles the type of Epidirona in most essentials; the sculpture is much stronger but of similar style, the form of the sinus, anterior canal features, thin outer lip, presence of fluting within the aperture and lack of a parietal tubercle all match very closely. The only deviation from typical Epidirona is in the proto- conch which is carinate on whorls two and three. Description — Adult shell, 15 to 19 mm. (M to % inches) in height, solid, drilliaform, with tall acu- minate spire but truncated body whorl. Whorls eight, plus a small depressed-turbinate protoconch of three whorls, the first whorl smooth, the second carinated below the middle and the third crossed by regular arcuate axials in addition to the carina. Sculpture strong, consisting of spiral gemmate cords crossed by weaker axials. On the spire there are two massive gemmate folds, one subsutural and the other forming the sinus ridge; both are com- Plate 229. Radula of Epidirona hedleyi Iredale. From specimen from ANSP, Port Jackson, New South Wales, Australia (courtesy of V. Orr). [22 - 879] 300 Epidirona A. W. B. Powell Turridae posed of two fused cords. The gemmules also are fused to form tall, narrowly rectangular nodes, the subsutural series slightly retractive and the sinus series slightly pro tractive; they number about twenty per whorl. Between the two folds is a deep sulcus bearing one or two spiral threads. On the base below the periphery there are eight strong spiral gemmate cords and a further series of six plain spiral cords on the anterior end. The body whorl is truncated and has a very short anterior canal with a wide shallow notch at the termination of a well-defined fasciole. Outer lip thin, fluted within. Sinus moderately deep, nar- rowly U-shaped, occupying the upper portion of the broad peripheral carina. Colour dull creamy white. Measurements (mm.) — height width 19.0 7.0 Mekran Coast (Brit. Mus.) 15.0 5.0 Ceylon (holotype) 14.2 5.6 Singapore Synonymy — 1877 Pleurotoma multiseriata E. A. Smith, Ann. Mag. Nat. Hist. (ser. 4) 19, p. 491. 1917 Turris (Gemmula) multiseriata E. A. Smith, Melvill, Proc. Malac. Soc. 12, p. 145, pi. 8, fig. 3. Types — The type locality is Ceylon. The holo- type is in the British Museum (Natural History), London. Records — CEYLON : (type locality); INDIA: Madras (Winckworth coll., Brit. Mus. ); PERSIAN GULF; Mekran Coast (Townsend coll., Brit. Mus.); MALAYA: Johore Strait (Winckworth coll., Brit. Mus.); Singapore (Powell coll.). THAILAND: 4 mi. E. of Phuket Harbour, 80 ft. ( R. Tucker Abbott, 1963 ) . List of Recent Epidirona species beachportensis (Cotton & Godfrey, 1938), Epi- deira, Rec. S. Aust. Mus. 6(2), p. 205, pi. 17, fig. 4 (150 fms. off Beachport, South Aus- tralia ) . Candida Laseron, 1954, Epidirona, N.S.W. Turridae, Handb. Roy. Zool. Soc. N.S.W., p. 10, pi. 2, fig. 30 (70-80 fms. off Cape Everard, N.S.W., Australia ) . See pi. 230, fig. 9. Plate 230 Figs. 1, 2 Epidirona hedleyi Iredale. Port Jackson, New South Wales, Australia. 27 mm. 3 Epidirona multiseriata (E. A. Smith). Holotype. Ceylon. 15 mm. Type in Brit. Mus. (Nat. Hist. ) 4 Epidirona carinata Laseron. Holotype. 40-50 fms., off New South Wales, Australia. 17 mm. 5 Epidirona nodulosa Laseron. Holotype. 50 fms., off New South Wales, Australia. 19.5 mm. 6 Epidirona gabensis (Hedley). Holotype. 80 fms., off Gabo Id., Victoria, Australia. 7 Epidirona torquata (Hedley). Holotype. Port Arthur, Tasmania. 8 Epidirona molleri Laseron. Holotype. 50 fms., off New South Wales, Australia. 16 mm. 9 Epidirona Candida Laseron. Holotype. 70-80 fms., off New South Wales, Australia. 15 mm. 10 Epidirona tuberculata Laseron. Holotype. 40-50 fms., off New South Wales, Australia. 15.5 mm. 11 Epidirona costifera Laseron. Holotype. 40-50 fms., off New South Wales, Australia. 16 mm. ( all types except fig. 3 in the Australian Museum, Sydney ) [22 - 880] March 31, 1964 INDO-PACIFIC MOLLUSCA, vol. 1, no. 5 Epidirona 301 carinata Laseron, 1954, Epidirona, N.S.W. Turridae, Handb. Roy. Zool. Soc. N.S.W., p. 9, pi. 2, figs. 24-26 (40-50 fms. off Manning R„ N.S.W., Aus- tralia). See pi. 230, fig. 4. costifera Laseron, 1954, Epidirona, N.S.W. Turri- dae, Handb. Roy. Zool. Soc. N.S.W., p. 10, pi. 2, fig. 29 (40-50 fms. off Manning R., N.S.W., Australia). See pi. 230, fig. 11. flindersi (Cotton & Godfrey, 1938), Epideira, Rec. 5. Aust. Mus. 6(2), p. 205, pi. 17, fig. 1 (80 mi. west of Eucla, 75 fms.. South Australia). gabensis ( Hedley, 1922 ) Epideira, Records Austral- ian Mus. 13(6), p. 228, pi. 53, fig. 16 (80 fms., Gabo Id., Victoria, Australia). See pi. 230, fig. 6. hedleyi Iredale, 1931, Epidirona, Records Austral- ian Mus. 18(4), p. 225 (Port Jackson, New South Wales ) . See pi. 230, figs. 1, 2. jaffaensis (Verco, 1909), Drillia, Trans. Roy. Soe. South Australia 33, p. 298, pi. 26, figs. 7-9 ( 130 fms. off Cape Jaffa, South Australia). molleri Laseron, 1954, Epidirona, N.S.W. Turridae, Handb. Roy. Zool. Soc. N.S.W., p. 11, pi. 2, figs. 31, 32 (50 fms. off Crowdy Head, N.S.W., Australia ) . See pi. 230, fig. 8. monile (Kiener, 1839-40), Pleurotoma, Coq. Viv. Pleurot. 5, p. 52, pi. 15, fig. 3 (non Rrocchi, 1814; see quoyi Desmoulins, 1842). nodulosa Laseron, 1954, Epidirona, N.S.W. Tur- ridae, Handb. Roy. Zool. Soc. N.S.W., p. 9, pi. 1, figs. 21-23 ( 50 fms. off Crowdy Head, N.S.W., Australia ) . See pi. 230, fig. 5. perksi (Verco, 1896), Surcula, Trans. Roy. Soc. South Australia 20, p. 224, pi. 7, fig. 3, a-c ( 15 fms. off Thistle Id., South Australia). philipineri ( Tenison-Woods, 1877), Pleurotoma, Proc. Roy. Soc. Tasmania, p. 136 (N.W. coast of Tasmania). quoyi (Desmoulins, 1842), Pleurotoma, Actes Soc. Linn. Rordeaux 12, p. 167, nom. nov. for Pi. monile Kiener, 1839-40 (non PI. monile Broc- chi, 1814). (Western Port, Victoria). schoutanica (May, 1911), Drillia, Proc. Roy. Soc. Tasmania (1910), p. 391, pi. 14, fig. 17 (80 fms., off Schouten Id., Tasmania). striata Gray, Hedley, 1922, Epideira, (not of Gray, 1827), Records Australian Mus. 13(6), p. 230, pi. 43, fig. 18-20 ( is now hedleyi Iredale, 1931 ) (New South Wales). torquata (Hedley, 1922), Epideira, Records Aus- tralian Mus. 13(6), p. 230, pi. 43, fig. 21 (Port Arthur, Tasmania). See pi. 230, fig. 7. tuberculata Laseron, 1954, Epidirona, N.S.W. Tur- ridae, Handb. Roy. Zool. Soc. N.S.W., p. 10, pi. 2, figs. 27, 28 ( 30-35 fms., off Port Stephens, New South Wales, Australia). See pi. 230, fig. 10. List of fossil Epidirona species adelaidensis (Ludbrook, 1941), Bathytoma, Trans. Roy. Soc. South Australia 65(1), p. 97, pi. 5, fig. 17 (Abattoirs Bore, 400-500 ft., Adelaide, South Australia; Pliocene). laevis Pritchard, 1904, var. of Pleurotoma selwyni, Proc. Roy. Soc. Victoria 17, p. 328 (non laevis Bell, 1890; see suppressa Finlay, 1927). powelli Ludbrook, 1957, Epidirona, Trans. Roy. Soc. South Australia 81, p. 86, pi. 5, fig. 3 (Weymouth’s Bore, 310-530 ft., Adelaide, South Australia; Lower mid-Pliocene ) . suppressa (Finlay, 1927), var. of Epideira selwyni, Trans. N.Z. Inst. 57, p. 516, nom. nov. for laevis Pritchard, 1904 (non Bell, 1890) (Vic- toria, Australia; Miocene). vardoni (Tate, 1899), Surcula, Trans. Roy. Soc. South Australia 23, p. 108, pi. 1, fig. 3, a-b (Murray Desert, Victoria, Australia (Janjuk- ian?, Miocene). [22-881] 302 Epidirona 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 - 882] March 31, 1964 INDO-PACIFIC MOLLUSCA, vol. 1, no. 5 Lophiotoma 303 Genus Lophiotoma Casey, 1904 Type: Pleurotoma acuta Perry, 1811. This is a well-known group of Indo-Pacific tur- rids closely resembling Tunis but with the sinus always on the peripheral keel, not on a subsidiary ridge above the periphery, as in true Tunis. For shells identical with Lophiotoma in adult features, but with a blunt paucispiral protoconch, see the subgenus Lophioturris, following. The counterparts of this genus in the tropical Eastern Pacific, the Caribbean and the Tertiary of the South Eastern United States are the genera Pleuroliria and Polystira, the former Eocene and Oligocene and the latter Miocene to Recent. A more shallow and broadly V-shaped sinus, and protoconch differences, separate these two from Lophiotoma, but there was probably a common an- cestry. Protoconch light reddish brown, tall, polygyrate, narrowly conical of 3/2 to 4 whorls, strongly sculp- tured with numerous crisp flexuous axials over the last 2/2 whorls, crossed by numerous faint spiral threads, which are visible only on the axials, not in the interstices. The first 1/2 whorls are smooth, prob- ably due to erosion, for no very well preserved apices were found. A non-eroded protoconch is probably axially costate throughout, as in Xenuro- turris. The weak nodulation or undulation of the peripheral sinus rib on the first post-embryonic whorl, shown in the figure, does not persist over subsequent whorls, nor is it a common feature in other material examined. The protoconch figure is from a Persian Gulf specimen. Synonymy — 1904 Lophiotoma Casey, Trans. Academy Sci. St. Louis 14(5), p. 130. 1928 Lophiotoma Casey, Woodring, Publ. no. 385, Carnegie Inst. Washington, pt. 2, p. 146. Type by subsequent designation: Pleurotoma tigrina Lamarck, = acuta Perry, 1811. Plate 232. Nuclear whorls of Lophiotoma s.s. and the sub- genus Lophioturris. Left fig., Lophiotoma (Lophiotoma) acuta (Perry) from the Persian Gulf. Right fig., Lophiotoma ( Lophioturris ) indica (Roding) from Yeppoon, Queensland, Australia. Lophiotoma acuta (Perry, 1811) (Color pi. 180; pi. 233; pi. 234) Range — Recent: Red Sea and East Africa to Ja- pan and Melanesia. Fossil: Pliocene and Pleisto- cene of Indonesia. Remarks — This species, better known as tigrina Lamarck, is the most widespread of the turrids. It is also most variable in shape, sculpture and in colour pattern. The more stable of these variants conform vaguely to geographic patterns over mar- ginal areas but in no hard and fast manner, for not only do forms integrate and coalesce at the cross- roads between the Indian and Pacific Oceans but odd forms are likely to crop up in an otherwise con- stant assemblage for an area. This is true of jickelii Weinkauff, the type local- ity for which is the Red Sea. The jickelii form tends to coarser sculpture than in acuta typical, the trend being a reduction in strength of the peripheral carina and increased prominence of the primary cords. See pi. 180, figs. 14, 19. The writer has seen only two Red Sea examples so is not competent to state what the overall char- acteristics of the Red Sea populations may be. The few Persian Gulf specimens examined show slight carinal reduction but are nevertheless nearer to typical acuta. Forms resembling jickelii were noted also from Zanzibar, Philippines and Buchan’s Point, Cairns, Queensland, but in all cases either in asso- Plate 233. Figs. 1-3, Lophiotoma acuta (Perry) form jickelii (Weinkauff). Cebu Id., Philippines. Figs. 4, 5, Lophiotoma acuta (Perry). Cotype of L. microsticta Casey, 1904. Cebu Id., Philippines (U.S. Nat. Mus.). [22 - 9131 304 Lophiotoma A. W. B. Powell Turridae ciation with or in close proximity to a population of typical acuta. If the Red Sea populations should prove to be predominantly jickelii type then there may be a case for considering that population a geographical subspecies which tends to infiltrate eastward to Zan- zibar, the South China Sea and north Queensland, but the evidence at present is against such an assumption. What little is known of the ecological conditions under which the respective forms live shows this factor to be of importance both in respect to the proportions of the shell and the coloration. The elongated, pale, sparsely-spotted form, microsticta Casey from the Philippines, for instance, is the com- mon and apparently exclusive form in the Marshall Islands where it occurs in shallow water on white coral sand. Similar shells range northward to Japan. A narrow but profusely-maculated form is found in East Indies waters and these closely resemble Lamarck’s Encyclopedie methodique figure (pi. 439, fig. 6) of his marmorata (non marmorata La- marck, 1822). It is significant that shells of this form from Koeroedoi Island, Dutch New Guinea, were found living on black sand and silt. A broad heavily-maculated shell is the common form at Zanzibar, East Africa and Madagascar, and this form spreads across the Indian Ocean and in- filtrates strongly throughout the East Indies, the Philippines and Queensland. Sowerby’s Pleurotoma notata from Hong Kong resembles the jickelii form of acuta, but in this case suppression of the peripheral carina is due to shell injury at the commencement of the antepenultimate whorl. See pi. 234. Another synonym of acuta is WeinkaufFs pictu- rata which fits the pale, sparsely patterned shells with a relatively short canal, found on sand in many shallow-water locations along the north Queensland coast. These are nearest to acuta typi- cal, but the broad heavily maculated form of acuta, the narrow heavily maculated form (maculata) and the coarsely sculptured jickelii form also occur along the Queensland coast and on the offshore is- lands. All occur in sandy locations, but the macu- lata form seems to favour deeper water on a shelly bottom. In Fiji, the typical, the maculata and the pictu- rata forms all occur. Specimens, with field notes, from the collection of Mr. W. Jennings show that the typical form occurs in sand at 2 or 3 feet below low water, the picturata form from 4 or 5 fathoms and the maculata form from 4 or 5 feet below low tide off the edge of coral reefs. Still another probable synonym of acuta is Dun- ker’s Pleurotoma peaseana, the type of which is supposed to be in the Godeffroy Museum, Ham- burg. It was referred to the synonymy of marmo- rata (= acuta) by Hedley (1922, p. 216). Specimens which approach nearest to Perry’s fig- ure in proportions, sculptural detail and coloration come from the islands of the tropical Southwest Pacific. Typical acuta is a relatively broad shell, strongly carinate and with a sparse pattern of small brown dots on a pale ground. Description of Typical Form — Adult shell 45 to 50 mm. (K to 2 inches) in height, of light build, fusiform, with a tall spire, 27° to 34°, and with a long rather straight canal. Spire slightly taller than the height of the aperture plus canal. Whorls 10 or 11, plus a tall narrowly-conic protoconch of 3M to 4 whorls, of yellowish brown colour and strongly sculptured with closely spaced, slightly oblique and arcuate axials which are crossed at right angles by faint spiral threads. Spire whorls dominated by the centrally placed sinus carina, consisting of an up- per and a lower sharp-edged keel with a smooth concavity between. The upper keel projects slightly more than the lower one. The primary cords con- sist of a strong subsutural one, and three below the carina, the second one emergent at the suture and the third below the level of the top of the aperture. The secondary sculpture forms a dense surface-cover of spiral threads of varying strength, 8 to 10 in number on the shoulder from between the subsutural cord and the sinus rib. Ground col- our, creamy-white, sparsely speckled with small, dark-brown, rectangular spots, most prominent on the subsutural cord, both edges of the sinus rib and on the three primary cords below it. Most of the secondary spirals are minutely speckled. Measurements (mm.) — ( a ) Typical sparsely speckled form height width spire angle 50.0 16.0 32° Fiji 46.0 15.5 34° Fiji 44.0 13.0 28° Low Isles, Queensland 43.5 14.0 33° Fiji 30.0 11.0 32° Samoa Spire angle 28° to 34° = average 31.8° (b) Narrow, sparsely maculated form ( microsticta ) 59.7 14.6 23° Cebu, cotype of microsticta 51.4 13.0 25° Kii, Japan 48.7 13.5 25° Ryukyu Ids. 44.0 12.3 25° Palau Ids. Spire angle 23° to 25° = = average 24.5° [22 - 914] March 31, 1964 INDO-PACIFIC MOLLUSCA, vol. 1, no. 5 Lophiotoma 305 ( c ) Broad, heavily maculated form ( tigrina Lamarck ) 66.5 17.0 28° Chwaka, Zanzibar 65.0 18.0 30° Kiwengwa, Zanzibar 60.0 16.7 30° Mozambique 58.0 17.7 29° Mozambique 57.5 16.0 29° Ko Phuket, W. Thailand 53.0 16.0 30° Mozambique 43.5 13.2 28° Low Isles, Queensland 41.5 13.0 30° King’s Reef, N. Queens- land 39.5 12.0 27° Tutuila, Samoa Spire angle 27° to 30° = average 29.1° (d) Narrow, heavily marck, 1816) maci dated form ( marmorata La- 48.0 13.0 28° all Kaipoeri, Dutch New Guinea 46.2 12.5 27° 42.5 11.0 26° 30.1 7.7 24° 25.2 7.0 26° Spire angle 24° to 28° = average 26.2° Description of jickelii form — The jickelii form is narrow shell with the sinus rib formed of two closely spaced, rounded cords and with the second- ary sculpture quite strongly developed. The sinus rib is not strongly projecting as in the typical spe- cies and the upper margining cord does not project farther than the lower one. The colour pattern is in the form of numerous dark, reddish brown, rec- tangular patches and spots roughly connected longitudinally with wavy lines, reminiscent of the crispus pattern. Adult shell 42 to 47 mm. (about 1/2 to 2 inches ) in height, fusiform, with a tall spire, 22° to 25°, and a long canal. Spire approximately equal to the height of the aperture plus canal. Spire whorls weakly carinated medially by the sinus ridge which is bordered below by a rounded cord. Subsutural fold well-developed as a broad collar, almost as strong as the sinus rib. The whole surface crossed by strong, rounded, spiral cords and crisp threads. Whorls 11 or more. Measurements (mm.) — height width spire angle 53.0 13.0 _ 49.0 14.0 25° 47.0 13.7 22° 41.7 11.6 24° ( Weinkauff) Synonymy — 1811 Pleurotoma acuta Perry, Conchology, London, pi. 54, fig. 5 (no locality). 1816 Pleurotoma marmorata Lamarck, Tableau Encycloped- ique et Methodique, Paris, pi. 439, fig. 6, Le Liste, p. 8. 1822 Pleurotoma tigrina Lamarck, Hist. Nat. Anim. sans Vert. 7, p. 95. 1829 Pleurotoma punctata Schubert & Wagner, Suppl. Conch. Cab. 12, p. 155, pi. 234, figs. 4103, a, b (no locality). 1839-40 Pleurotoma tigrina Lamarck, Kiener, Coquilles Vi- vantes 5, p. 10, pi. 8, fig. 1 (Indian Ocean and Madagascar ) . 1843 Pleurotoma tigrina Lamarck, Deshayes, Anim. sans Vert. 9, p. 352, sp. 20 (no locality). 1843 Pleurotoma tigrina Lamarck, Reeve, Conch. Iconica 1, pi. 1, fig. 3 (Philippines, sandy mud). 1871 Pleurotoma {Turris) peaseana Dunker, Malak. Blatt. 18, p. 154. 1875 Pleurotoma jickelii Weinkaulf, Conch. Cab. 4(3), p. 20, pi. 4, figs. 2, 3. 1876 Pleurotoma peaseana Dunker, Weinkauff, Conch. Cab. 4(3), p. 69, pi. 15, figs. 1-3. 1876 Pleurotoma picturata Weinkauff, Conch. Cab. 4(3), p. 66, pi. 2, fig. 10. 1884 Pleurotoma tigrina Lamarck, Tryon, Manual of Conch. 6, p. 164, pi. 2, fig. 10. 1888 Pleurotoma notata Sowerby, Proc. Zool. Soc., p. 566, pi. 28, fig. 17. 1904 Lophiotoma microsticta Casey, Trans. Acad. Sci. St. Louis 14, p. 130 (not figured; Cebu, Philippines). 1915 Pleurotoma (s.str. ) tigrina Lamarck, Tesch, Palaeon- tologie von Timor 9, Stuttgart, p. 24, pi. 76(4), fig. 36 (Pliocene, Timor). 1931 Pleurotoma tigrina Lamarck, van der Vlerk, Leidsche Geol. Meded. Rijksmus. Geol. Min. Leiden 5, p. 219 ( Sumatra, Java, Timor and Celebes. Miocene, Pliocene and Pleistocene records). 1956 Lophiotoma tigrina Lamarck, Kaicher, Indo-Pacific Sea Shells, section 5, Toxoglossa, pi. 1, fig. 5. 1961 Turris tigrina Lamarck, Habe, Shells of Japan 2, pi. 38, fig. 18 (microsticta form). 1961 Turris notata Sowerby, Habe, Shells of Japan 2, pi. 38, fig. 16. Types — The type of notata is in the British Mu- seum (Natural History); it measures 55 x 16mm. Plate 234. Lophiotoma acuta (Perry). Holotype of Pleuro- toma notata Sowerby, 1888. Hong Kong. [22 - 915] 306 Lophiotoma A. W. B. Powell Turridae Records — RED SEA: Massaua (type of jickelii); (Cal- vert coll., MCZ). PERSIAN GULF: (Brit. Mus.). ANDA- MAN ISLANDS: Port Blair (Winckworth coll., Brit. Mus.). SEYCHELLES: N.W. Bay, Mahe (Winckworth coll., Brit. Mus.). CEYLON: (USNM). ZANZIBAR: Chwaka, 0-4 ft.; 1/2 mi. W.S.W. Ras Mungwe, 8 fins.; 1 mi. W.N.W. Ras Mbweni, 7 fms.; S. side of Pwakuu Id., 11-18 fms.; between Mwamba, Ukombi and Chumbe Ids., 4-6 fms.; Kizim- kazi, 0-3 ft.; Kiwengwa, 0-3 ft. (ANSP). MADAGAS- CAR: 9-12 ft., Pte. du Cratere, S.W. Nossi Be; 9 fms., 3 mi. N.N.E. of Nossi Fali, E. of Nossi Be (ANSP). MAURITIUS: (USNM). PORTUGUESE EAST AFRICA: Mozambique City (ANSP). NORTH BORNEO: shore at Kudat (Mrs. M. Saul, ANSP). MALAYA: Singapore (ANSP). THAILAND: 4 mi. E. of Phuket Harbour, 80 ft. (R. Tucker Abbott). DUTCH NEW GUINEA: 3 ft., black sand and silt, % mi. S. of Kaipoeri Village, Koeroedoi Id. (nearest to Lamarck’s 1816 marmorata); N.E. Ambai Id., Japen Id., 5 ft., old reef; 2 mi. N.W. Rani Id., Schouten Ids., 4-40 ft.; island off N.E. tip of Ambai, Japen Id., 10-15 ft.; 1.3 mi. S.W. of Cape Mantoewoeri, Koeroedoi Id., 4 fms.; /2— 1 mi. E. of Mios Woendi Id., 3-30 ft. (ANSP); near Hollandia (USNM). HONG KONG: (type of notata) . JAPAN: Osima Osumi (USNM); Kii (ANSP) (A. W. B. Powell coll.). RYUKYU ISLANDS: Ohama Bay; Nago; Taketomi Id. (D. Thaanum coll.). PHILIPPINES: Island of Cebu ( cotype of microsticta ) ; Port Binang, Subic Bay (USNM); Tabaco, Albay Province, Luzon; Zamboanga, Mindanao Id.; 12-24 ft., E. side Jagoliao Id., N.W. end of Bohol Id.; Calatagan. Batangas, Luzon Id. (ANSP); Davao, Mindanao Id., (A. W. B. Powell coll.). PALAU ISLANDS: S.E. comer of Eil Malk. Kayangel Lagoon, 5-15 ft.; E. of Ferry Dock. Peleliu, dredged; N. of entrance to Karamando Bay, West Babelthuap; 3 mi. N.E. of Eil Malk, E. of Yoo Passage. Malakal Harbour, dredged; E. lagoon off Mele- keiok. East Babelthuap Id.; 1 mi. S. of West Passage, Babelthuap Id.; Kossol Passage, west entrance, 20 fms. (all ANSP). MARSHALL ISLANDS: Taka Id., Taka Atoll; Bikini Id., N.E. shore; E. side Enyer Id., Bikini; 2 mi. W. of Busch, 20 fms.; 2 mi. N. of Ime, Amo Atoll; 1-2 mi. N. of Jiyabo Lagoon, Amo Atoll (USNM) (all microsticta form). NEW HEBRIDES: (Australian Mus.). SOLOMON ISLANDS: Ugi Id., (USNM). NEW CALEDONIA: 4-10 ft., barrier reef, Touho Bay (ANSP). AUSTRALIA: Queens- land: Challenger Bay, Palm Id. (ANSP); Dunk Id.; King’s Reef (A. W. B. Powell coll.); Hope Id., 5-10 fms.; Lizard Id.; Murray Id.; Palm Id.; Mast Head Ids.; Rocky Ids.; Cape Flattery; Crescent Reef ( Australian Mus. ) ; Moreton Bay (Mrs. J. Kerslake). FITI: Beea Id.; Vatulele Id. (D. Thaanum coll.); Yasawas Id. (USNM); Akuilau Id., 2-5 ft.: Wading Id.; Namotu Id.. 4-5 fms. (W. Jennings). SAMOA: Toala. N.W. Upolu Id.: 10 ft., sand, lagoon off Mulifanua, Upolu Id.; Tutuila (ANSP). Fossil Records — PMiocene, West Sumatra; Pliocene, Sonde beds, Java; Timor. Pleistocene, Celebes (van der Vlerk, 1931, p. 219). Lophiotoma albina (Lamarck, 1822) (Color pi. 180, figs. 11, 12; pi. 235) Range — Madagascar to the Carolines, Microne- sia. Remarks — Although closely allied to acuta this species is remarkably constant. The chief distin- guishing points are the triple-corded sinus rib, relatively weak spiral sculpture, strong overlay of dense axial folds, a very distinctive pattern con- sisting of an overall sprinkling of minute brown dots, and a peripheral row of regularly spaced dark-brown rectangular markings, each of which is in the form of three horizontal lines defining each of the three sinus cords. The general appear- ance of the shell is white with a revolving row of peripheral maculations. Whorls 10, plus the proto- conch, a remnant of which remains in one speci- men, suffice to show that it is of a similar multi- spiral form as that of acuta. Adult size 37 to 41.5 mm. (about VA inches) in height. Spire about three-fourths the height of the aperture plus canal, angle 23° to 25°. Whorls only slightly angled by the peripheral carina. Measurements (mm.) — height 74.0 width spire angle Balabac, Philippines 57.0 14.5 - Andaman Ids. 41.5 11.2 23° Moluccas 37.0 11.0 25° Moluccas Synonymy — 1822 Pleurotoma albina Lamarck, Anim. sans Vert. 7, p. 96. (no locality). 1843 Pleurotoma albina Lamarck, Reeve, Conch. Iconica 1, pi. 9, fig. 77. Records — MADAGASCAR: Mayotte, 50 metres, coarse sand in lagoon (ANSP). ANDAMAN ISLANDS: (Tomlin coll., Nat. Mus. Wales). MOLUCCAS: (Casey coll., USNM). Java: (Tomlin coll.). Caroline Islands: Yap (C. Kile, 1960, ANSP). PHILIPPINES: Balabac, Palawan Id. (coll. A. D’Attilio, New York); Lubang Id. (Tomlin coll.). Lophiotoma albinoides (Martin, 1883) (PI. 246, figs. 1, 2) Range — Java, Tji Longan, near Selatjan, Njalin- doeng beds, Lower Miocene. Remarks — This species appears to be quite dis- tinct judging from the illustrations which also show an enlargement of the sculptural detail. The shell is slender like the Recent albina, but the Plate 235. Lophiotoma albina (Lamarck). Molucca Is- lands. 41 mm. [22 - 916] March 31, 1964 INDO-PACIFIC MOLLUSC A, vol. 1, no. 5 Lophiotoma 307 Plate 236. Geographical distribution of Lophiotoma (Lophiotoma) acuta (Perry) in solid dots and L. (L.) albina (Lamarck) in open circles. whorls are not rendered so angulate by the spiral keels. Also, it would appear from the figures that the sinus rib is above the middle and consists of two closely spaced, beaded spirals with a heavier smooth one below. The subsutural spirals are weak. It is more likely, however, that the smooth peri- pheral carina will prove to be coincident with the sinus, the shape and nature of which is not appar- ent in the illustrations. Until the type is critically examined, the generic status of this species cannot be determined. It is referred to Lophiotoma mainly on account of Mar- tin’s comparison of the species with albina La- marck. No measurements were given. Synonymy — 1883 Pleurotoma albinoides Martin, Samml. Geol. Reichs- mus., Leiden, 1, p. 227, pi. 10, figs. 23, 23a. 1921 Pleurotoma albinoides Martin, Vredenburg, Rec. Geol. Surv. India 53(2), p. 97. Lophiotoma abbreviata subspecies abbreviata (Reeve, 1843) (PI. 237, figs. 1, 2; pi. 238, figs. 1, 2) Range — Mauritius to Fiji. Remarks — This species closely resembles acuta except for the truncated anterior canal, more broadly conic spire and heavier sinus rib. The lat- ter is composed of two strong, closely spaced, smooth spiral cords. The coloration is similar, ex- cept that the subsutural maculations are heavier. This species seems to occur only in shallow water on coral reefs and in sandy pockets. A var- iant in populations at Rodriguez Island, Mauritius and the Andaman Islands has the sinus rib less prominent, coupled with a pattern in which the speckles are more prominent and the subsutural maculations less so, but they seem to occur along with the typical form. Description — Adult shell 25 to 36 mm. ( 1 to VA inches ) in height. Spire 1.3 times the height of the aperture plus canal, angle 28° to 35°. Whorls 8 or 9, exclusive of the protoconch which is missing in Plate 237. Figs. 1, 2, Lophiotoma abbreviata (Reeve). Near Suva, Viti Levu Id., Fiji. 36 mm. Figs. 3, 4, L. abbre- viata lifuensis (Sowerby). Lifu Id., Loyalty Islands. 19 mm. [22 - 917] 308 Lophiotoma A. W. B. Powell Turridae all examples examined. Spire rather broadly conic and strongly angulated at the middle by a bicari- nate sinus rib. Spire whorls with a broadly convex subsutural fold and a secondary cord between the periphery and the lower suture. On the base there are two closely spaced primary cords or sub-keels, the uppermost emergent at the suture, plus 4 or 5 weaker primary cords on the middle area of the base, plus 5 or 6 from the neck to the tip of the anterior canal. The whole surface is crowded with fine, crisp, spiral threads. The sinus rib is composed of a pair of smooth rounded cords, with a slight concavity between. Aperture strongly lirate a little distance within the outer lip. Sinus narrow, of moderate depth. Colour creamy white, speckled with light-brown on the spirals, plus prominent dark-brown rectangular maculations which are seri- ally arranged around the subsutural fold. Measurements (mm.) — height width 36.0 + 14.5 Fiji 32.0 12.3 Philippines 29.5 11.5 Fiji 26.5 11.0 Andaman Ids. 25.0 11.0 lectotype 24.5 10.5 Fiji Synonymy — 1843 Pleurotoma abbreviate Reeve, Conch. Icon. 1, pi. 10, fig. 86. 1884 Pleurotoma abbreviate Reeve, Tryon, Manual of Conch. 6, p. 167, pi. 3, fig. 25. Plate 238. Figs. 1, 2, Lophiotoma abbreviate (Reeve, 1843). Two of four syntypes in the British Museum (Na- tural History). Reefs at low water, Masbate Island, Philip- pines. Lectotype, fig. 1, is 25 x 11 mm. Fig. 3, L. abbrevi- ate ustulata (Reeve, 1846). Holotype, no locality, Brit. Mus., 31 x 12 mm. Types — Lectotype, one of four syntypes here selected, is in the British Museum (Natural His- tory). Records — MAURITIUS: ex N. Pike (USNM); Rodriguez Id. (USNM). ANDAMAN ISLANDS: (USNM). HALMA- HERA GROUP: Morotai (USNM). PHILIPPINES: Mas- bate Id., on reefs at low water (type); Mariveles, Bataan, Luzon Id. (ANSP); Lubang (Winckworth coll., Brit. Mus.). PALAU ISLANDS: S. end of Gorokottan Id.; 1 mi. S. of West Passage, Babelthuap Id. (ANSP). NEW CALE- DONIA: Touho Bay, 0-1 ft., fringe reef, with sand pockets; La Roche Percee, Burail (ANSP). FIJI: ex Garrett (ANSP); Bega Id. ( D. Thaanum coll. ) ; reef at Suva, Viti Levu Id. (A. W. B. Powell coll.); Levuka, Vanua Levu Id. (USNM). Lophiotoma abbreviata subspecies lifuensis (Sowerby, 1907) (PI. 237, figs. 3, 4) Range — Lifu, Loyalty Islands. Remarks — This form which is known only from the type locality of Lifu is much broader-spired and more squat than the typical subspecies. The peri- pheral keel is more prominently projecting and the two basal sub-keels are much stronger. It may be only an ecological form, but so far it has not turned up from other localities. Description — Adult shell 17 to 20 mm. ( about 1 inch ) in height. Whorls 5+, exclusive of protoconch, missing in all examples examined. Spire a little more than the height of the aperture plus canal, angle 40° to 50°. Coloration and sculptural detail as in the typical subspecies, except that the peri- pheral keel, which is the sinus rib, strongly pro- jects, flange-like, giving a ledged appearance to the shoulder. Measurements (mm.) — height width 19.7 10.0 19+ 11.0 17.0 10.0 holotype Synonymy — 1907 Pleurotoma abbreviate lifuensis Sowerby, Proc. Mai. Soc. 7, p. 300, pi. 25, fig. 5. Records — LOYALTY ISLANDS: Lifu (type locality); (ANSP, Brit. Mus.; USNM). Lophiotoma abbreviata subspecies ustulata (Reeve, 1846) (PI. 238, fig. 3) Range — Probably Indian Ocean, Mauritius and Andaman Islands. Remarks — It is likely that this shell will prove to be a synonym of abbreviata, or at most a geo- graphic subspecies of it. Reeve’s type is from an [22 - 918] March 31, 1964 INDO-PACIFIC MOLLUSC A, vol. 1, no. 5 Lophiotoma 309 Plate 239. Geographical distribution of Lophiotoma (Loph- iotoma) abbreviata (Reeve) and its subspecies ustulata ( Reeve ) and lifuensis ( Sowerby ) . Plate 240. Fig. 1, Lophiotoma brevicaudata (Reeve). Gulf of Suez. 22 mm. Fig. 2, Lophiotoma ruthveniana (Melvill). Holotype. Mauritius. 41.5 mm. unknown locality but similar shells are known from the two Indian Ocean localities mentioned above; not from the Pacific range of the typical species, however. The type specimen is badly faded but the charac- teristic colour pattern of the typical species, par- ticularly the subsutural maculations, is still dis- cernable. The sculpture, however, is much weaker than is normal for abbreviata, the bicingulate peri- pheral keel being more like that of acuta; the an- terior canal, however, is truncated. Until more is known of this form it had better be admitted as a probable subspecies of abbreviata. Measurements (mm.) — height width 31 12 holotype Synonymy — 1846 Pleurotoma ustulata Reeve, Conch. Iconica 1, pi. 40, fig. 369 (no locality). 1884 Pleurotoma ustulata Reeve, Tryon, Manual of Conch. 6, p. 167, “Mauritius.” Types — The holotype is in the British Museum (Natural History). Records — INDIAN OCEAN : Mauritius; Andaman Ids. ( USNM ). Lophiotoma ruthveniana (Melvill, 1923) (PL 240, fig. 2) Range — Mauritius. Remarks — This handsome shell appears closely allied to abbreviata and is distinguished from that species mainly by the overall strongly tessellated colour pattern. In abbreviata the larger maculations are confined to the subsutural fold but in ruthven- iana the peripheral carina bears heavier macula- tions. In the absence of well-preserved apical whorls in the four examples seen, including the holotype, the species is only provisionally placed in Lophiotoma. Description ( from Melvill ) — “Shell fusiform, thick; whorls, especially the upper, somewhat com- pressed, being ten in number, inclusive of the two apical. Colour bright chestnut brown, with squar- rose, fairly regular, white tessellations on the spiral [22 - 919] 310 Lopliiotoma A. W. B. Powell Turridae carinae. These revolving keels appertain through- out—one, in particular, central, and subdivided by a shallow sulcus; the lesser tornate keels increase nu- merically in each of the lower whorls, till, on the body whorl, they total five or six, all beautifully variegated with white and chestnut alternately, as mentioned above. Mouth ovate-oblong, canal wide, abbreviate, sinus well developed, wide and deep, columellar margin fairly straight. Long. 41.5, lat. 14 mm. “A handsome species, standing somewhat alone, and conspicuous for its bright coloration and tes- sellated carinal ornamentation.” Synonymy — 1923 Turris ruthveniana Melvill, Proc. Mai. Soc. 15, p. 162, pi. 4, fig. 2. Types — The type locality is Mauritius. The holo- type is in the British Museum (Natural History), London. Lophiotoma brevicaudata (Reeve, 1843) (PI. 240, fig. 1) Range — Gulf of Suez, New Guinea, the Philip- pines and New Caledonia. Remarks — Three syntypes of Reeve’s species are in the British Museum and there is little to add to his original description: “Shell shortly fusiform, solid, yellowish, brown at the base and apex; whorls convex, encircled with a single keel round the upper portion and a double keel round the lower; last whorl encircled with single and double keels alternately down to the base, lip simple and acute, sinus large; aperture small, short, canal rather short. Hab. Island of Ticao (found on reefs), Cuming.” It is an easily recognized little shell due to its regular smooth spirals and uniform yellowish brown colour, except for the brown staining of the anterior end. The apical whorls are eroded in all three syn- types. Tryon (1884, p. 169) made this species a syno- nym of fascialis Lamarck = polytropa Helbling, but they are certainly not con-specific. Apart from the type material and imperfect spec- imens from New Guinea and New Caladonia, the best preserved example seen is one from the Gulf of Suez (ex McAndrew coll., Brit. Mus.) in the Museum of Comparative Zoology, Harvard, and a description of the specimen follows: Description ( Red Sea specimen ) — Adult shell 22 mm. (slightly under 1 inch) in height, fusiform, with a tall spire, but with a relatively short anter- ior canal. Whorls 7, exclusive of the protoconch, sufficient of which remains in one specimen to indi- cate that it is narrowly conic of several whorls. Spire 1.3 times the height of the aperture plus canal. Adult sculpture of narrow, sharp cords and threads, one cord on a moderately inflated subsutural fold, five threads over a deeply concave shoulder. There is a double-corded peripheral keel, enclosing a con- cave interspace with a single thread in the middle. There is one primary cord and two threads between the peripheral carina and the lower suture. Three cords, stronger than the rest, are on the upper part of the base, with each interspace having three threads, followed by an alternation of cords and threads over the neck, and finally closely-spaced threads over the anterior canal. Sinus deep, U- shaped, on the peripheral carina. Colour pale yel- lowish buff, with all the spirals continuously lined in light-brown and with a dark-brown staining on the anterior canal. Measurements (mm.) — height width 24.0 8.3 lectotype 22.2 7.6 Gulf of Suez Synonymy — 1843 Pleurotoma brevicaudata Reeve, Conch. Iconica 1, pi. 15, fig. 126. 1884 Pleurotoma fascialis Lamarck, Tryon, Manual of Conch. 6, p. 169 (non Lamarck, 1822). Types - British Museum (Nat. Hist.), three syn- types, lectotype, 24 x 8.3 mm., here selected. The type locality is Ticao Id., Philippines, Hugh Cum- ing, collector. Records — PHILIPPINES: Ticao Id. (type locality); Cal- apan, Mindoro Id. (MCZ). NEW CALEDONIA: (Aus- tralian Mus.). WEST NEW GUINEA: Geelvink Bay, & mi. S. of Maroepi, Ambai, Japen Id., 14-25 fms., blue mud, shell and coral (ANSP). RED SEA: Gulf of Suez (MCZ). [22 - 920] March 31, 1964 INDO-PACIFIC MOLLUSCA, vol. 1, no. 5 Lophiotoma 311 Subgenus Lophioturris new subgenus Type: Turns indica Roding, 1798 The adult shell is identical to that of typical Lophiotoma, in that it is fusiform, with a tall spire, usually with a long straight anterior canal, and with a deep and narrow sinus situated on the peripheral keel. However, the protoconch is paucispiral, blunt- tipped, smooth and rounded. I hereby designate Turris indica Roding, 1798, as the type of this new subgenus. Lophiotoma indica (Roding, 1798) (Color pi. 175, figs. 2, 3, 9, 16; pi. 242) Range — Ceylon to Australia and Fiji. Remarks — This shell is better known by the name Pleurotoma marmorata Lamarck, 1822, a name, however, which cannot be used because Lamarck had already employed it in 1816 for a dif- ferent shell which now falls under the synonymy of Lophiotoma acuta (Perry). MacNeil ( 1960, p. 101 ) referred this species to his new genus Unedogemmula (type species, unedo Kiener), but indica has a paucispiral, smooth, dome-shaped protoconch and none of the examples I have seen exhibit granulation of the peripheral keel at any stage of growth. The diagnostic charac- teristics for Unedogemmula are a conical, multi- spiral protoconch, as in Gemmula, and there are peripheral gemmules on the early whorls, some- times persisting over all the whorls. Sowerby, 1888, p. 211, described a Pleurotoma bulowi from the China Sea as follows: “P. testa elongata-fusiformi utrinque attenuata, pura alba. Anfractus 13, primi 2 rotunde convexi, politi; cae- teri acute carinati, ubique dense spiraliter lirati; spira acutissima. Anfractus ultimus leviter con- vexus, ad basin valde contractus et multo produc- tus. Apertura ovata, sinu elongato, angusto emar- ginata; canali gracili, elongato. Long. 35, maj. diam. 10.” “ A pure white, keeled, and closely ridged species of the typical form of P. babylonicay A specimen labelled bulowi, from Hong Kong, in the British Museum, not the type which could not be found, is here figured. Bleached and eroded ex- amples from Darwin, N.W. Australia are of similar appearance to the above and others from Gladstone, Queensland, are pure white also, but not eroded, which suggests that bulowi is merely an albinistic form of indica. Typical maculated shells occur within the range of these albino records. Description — Adult shell 85 to 93 mm. ( about 3 to 4 inches) in height, elongate-fusiform, with a tall spire and a long, straight anterior canal, closely simulating Fusinus. Whorls 14, plus a small blunt, smooth, paucispiral protoconch of 1/2 whorls, fol- lowed by a half-whorl of brephic axials. Sculpture consisting of a strong but narrowly-rounded, smooth, peripheral carina located at a little below median whorl height, a subsutural fold carrying a moderately strong smooth spiral cord at its lower extremity, plus a weaker cord and several threads above it. One to three smooth primary spiral cords are present between the periphery and the lower suture, and there are 18 to 20 primary spirals on the body whorl from the periphery to the end of the anterior canal. The whole of the interstitial surface is crowded with smooth, spiral threads of varying Trw \yY V FORMOSA M4«.,vt V *vj rt MARIANA • Woko ... . ■ .Wot/0 Isjf \ 1 \ | LOPHI / • • V. '4 > ■ O /j iyi ■ \ CW b '% --'f- ifi -» COOK • Plate 260. Geographical distribution of Tunis spectabilis (Reeve) in solid dots, and Tunis annulata (Reeve) in open circles. covered by the last whorl. Sinus rib narrow, low and inconspicuous on the spire’s whorls and situ- ated immediately below the weak subsutural fold. Surface further sculptured with fine threads, 2 or 3 on the subsutural fold, 2 to 4 between the peri- pheral keels of the last whorl and 12 to 15 on the base and neck. Aperture plus canal half the spire height. Pillar twisted and with a prominent fasciole and false umbilical chink. Measurements (mm.) — height width no. whorls 78.0 21.0 14+ Bohol, Philippines 67.0+ 21.0 — holotype of cryptorrhaphe Sby. 67.0 17.7 11+ Philippines 59.0 16.5 - Bohol, Philippines Synonymy — 1825 Pleurotoma cryptorrhaphe Sowerby, Tankerville Cat., Append., p. 14, no. 1503 (no locality). 1828 Murex bicarinatus Wood, Index Testaceol. Suppl., p. 15, pi. 5, fig. 7 (no locality). 1839-40 Pleurotoma woodii Kiener, Coq. Viv. 5, Pleuro- tome, p. 12, pi. 7, fig. 1 (new name for bicarinatus Wood). 1842 Pleurotoma cryptorrhaphe Sowerby, Reeve, Conch. Syst. 2, p. 188, pi. 234, fig. 16 ( 15 in error on plate ) . 1843 Pleurotoma cryptorrhaphe Sowerby, Reeve, Conch. Icon. 1, pi. 1, fig. 7. 1875 Pleurotoma cryptorrhaphe Sowerby, Weinkauff, Conchyl.-Cab., 2nd series, Pleurotomidae 4(3), p. 20, pi. 4, figs. 4-7. 1884 Pleurotoma cryptorraphe (sic) Sowerby, Tryon, Man- ual of Conch. 6, p. 168, pi. 3, figs. 30, 31. 1913 Pleurotoma (Hemipleurotoma) cryptorhaphe (sic) Sowerby, Schepman, Siboga Exped., pt. 5, 49e, p. 400. Types — The holotype is in the British Museum (Natural History) in London. Records — INDONESIA: North Ubian, 16-23 metres (Schepman, 1913). PHILIPPINES: 12-24 ft, E. side of Jagoliao Id., N.W. end of Bohol Id. ( duPont-Acad. Exped., 1958, ANSP); Zamboanga City, Mindanao Id. (MCZ); Masbate Id. (Brit. Mus.). NEW GUINEA: Milne Bay (Stanford Univ.); MARSHALL ISLANDS: Kwajalein Atoll (USNM). FIJI: Ngualito Id., off north coast, under rocks partially buried in sand ( W. Jennings, 1961 ) . Turris spectabilis (Reeve, 1843) (Color pi. 181, figs. 16, 17) Range — Western Indian Ocean to Melanesia and the Phoenix Islands. Remarks — This gaily painted shell was relegated to the synonymy of babylonia by Tryon ( 1884, p. 162) but it is distinct from that species in having a shorter anterior canal, different sculpture and a characteristic colour pattern. It resembles garnonsii in its slender whorls and tendency toward wide dash-like maculations but differs in the shorter canal and broad fight-brown zones both on the shoulder and on the base. It is evidently an uncommon spe- cies. Description — Shell up to 80 mm ( 3 inches ) in height. Spire 30° to 32°, tall but rather broad, 1.2 times the height of the aperture plus canal. Spire whorls with a single, narrowly rounded, spiral cord in place of the subsutural fold. Sinus rib narrow and rounded, weakly gemmate over the early whorls. Peripheral carina massive, followed below by a less prominent spiral keel, just emergent at the lower suture. On the base there are 6 additional primary cords which are strong, although narrowly rounded and wide-spaced. Colour variegated, basically golden-brown except for a broad, white, peripheral band. All primary spirals maculated with dark- brown dashes and dots. [22 - 986] March 31, 1964 INDO-PACIFIC MOLLUSC A, vol. 1, no. 5 Tunis 337 Measurements ( i mm.) — height 80.2 width 22.3 Direction Id., Cocos-Keeling 76.0 20.0 (ANSP) Cebu Id., Philippines ( Brit. Mus.) 68.5 20.0 Cebu Id., Philippines ( Brit. Mus.) 64.0 19.5 Mios Woendi, Dutch New Guinea 55.0 16.5 (ANSP) Cebu Id., Philippines (Brit. Mus.) 42.0 15.0 Mauritius ( Powell coll. ) Synonymy — 1843 Pleurotoma spectabilis Reeve, Conch. Iconica 1, pi. 1, fig. 6, a, b. 1875 Pleurotoma spectabilis Weinkauff, Martini and Chemn., Conch.-Cab., Pleurotomidae 4(3), p. 22, pi. 4, fig. 9. 1922 Turris spectabilis Hedley, Rec. Australian Mus. 13(6), p. 217. Types — The type locality is Ticao Id., Philip- pines, Hugh Cuming, collector. The type was not located in the British Museum (Natural History). There are three specimens in the Cuming collection fastened to a wooden tablet and the middle one resembles Reeve’s figure, but they are labelled “Island of Zebu.” Records — MAURITIUS: (A. W. B. Powell coll.). ADEN: (Australian Mus.). COCOS-KEELING: Direction Id. (V. Orr and R. Ostheimer, 1963, ANSP). ANDAMAN ISLANDS: near Port Blair (Natal Mus.). PHILIPPINES: Ticao Id. (type); Cebu Id. (Cuming coll., Brit. Mus.). DUTCH NEW GUINEA: reef 1 mi. N.E. of Mios Woendi Id., Padaido Ids. (NSF, 1956, ANSP). SOLOMON IS- LANDS: Malaita Id. (Powell coll.). AUSTRALIA: Queens- land: 20 fms., Damley Id. (Australian Mus.); Tryon Id., Capricorn Group (Mrs. J. Kerslake). MARSHALL IS- LANDS: Eniaetok Id., Rongelap (USNM). FIJI: Ngualito Id., off north coast, under rocks partially buried in sand (W. Jennings, 1961). PHOENIX ISLANDS: Canton Id. (Cal. Acad. Sci. ) . Turris ambages Barnard, 1958 Range — Off Natal and Zululand, South Africa. Remarks — No figure of the complete shell is given, just two line drawings, one of the protoconch and one of the sculptural detail of one whorl (which whorl not stated). The only clue to the location of this species is the author’s comparison of it with fagina. Nothing further can be done with this species without recourse to the type specimens and from the description it would appear that all of these are immature. Description (from the original) — “Protoconch 3 whorls, diam. 1.25 to 1.3 mm., alt. 1.3 to 1.5 mm., with axial riblets (as in Turris). Postnatal whorls with four strong spiral keels (profile of whorl 4- lobate ) , 1st rather sharp, forming the cingulum, 2nd rounded forming the lip sinus, with growth-lines producing more or less conspicuous squamiform nodules, 3rd the most prominent, rounded, forming the periphery, below this the smaller 4th keel; base with about 10 additional lirae. 17 x 7 mm. ( proto- conch and 7 whorls ) ; 25 x 8 ( protoconch and 8 whorls); a broken specimen 30 mm. long, with 5 whorls corresponding to the 5th to 8th whorls plus a portion of the 9th whorl. The 17 mm. example is whitish, with traces of irregular brown spots and streaks.” Measurements (mm.) — (See above) Synonymy — 1958 Turris ambages Barnard, Ann. South African Mus., 44, p. 148, fig. 23b. Types — The holotype is in the South African Museum. Records — SOUTH AFRICA: Off Cape Natal (Durban), 54 fms., 2 broken specimens; off Umkomaas River, 41 fms., 1 (17 mm. long); off O’Neil Peak (Zululand), 90 fms., 1 (25 mm. long). Australian Tertiary Species Turris selwyni (Pritchard, 1904) Range — Balcombian Miocene of Victoria, Aus- tralia. Remarks — Related to septemlirata but a smaller shell, 28 to 38 mm. ( 1 to 1 M inches ) in height, pro- portionately broader, with the spire less than height of aperture plus canal. Sculpture consisting of spiral keels, two on early whorls, three medially and a fourth half-emergent over the last half-whorl. The sinus rib is composed of a pair of sharp cords with a smooth, narrow, concavity between, and this is situated just above the greatest convexity, at about two-thirds whorl height. This and the following Australian Miocene spe- cies resemble Recent annulata. Measurements (mm.) — height width 38.0 17 holotype 32.7 14 Balcombe Bay, Victoria Synonymy — 1904 Pleurotoma selwyni Pritchard, Proc. Roy. Soc. Viet. 17, p. 326, pi. 19, fig. 1. 1944 Turris selwyni Pritchard, Powell, Rec. Auck. Inst. Mus. 3(1), p. 8. Records — AUSTRALIA: Muddy Creek, lower beds, Vic- toria (type); Balcombe Bay, Victoria (Balcombian, Mio- cene ) . [22 - 987] 338 Tunis A. W. B. Powell Turridae Turris septemlirata (Harris, 1897) Range — Balcombian Miocene of Victoria, Aus- tralia. Description — Adult shell 50 to 60 mm. ( 2 to 2M inches) in height, broadly fusiform, with an atten- uated spire but a somewhat truncated and twisted anterior canal. Spire 1.3 times the height of the aperture plus canal. Sculpture of prominent, rounded spiral keels, 4 on the penultimate; the rather deep and narrow sinus is situated on the second keel from the top, which is the one just above the greatest convexity of the whorls. The species bears some resemblance to the Recent an- nulate Reeve. Measurements (mm.) — height width 60.0 22.0 holotype 46.0 16.0 Muddy Creek, lower beds Synonymy — 1896 Pleurotoma perarata (Tate ms.) Cossmann, Ess. Pal. Comp. 2, p. 77. 1897 Pleurotoma septemlirata Harris, Cat. Tert. Moll. Brit. Mus. 1, p. 39, pi. 2, figs. lOa-d. 1898 Pleurotoma septemlirata Harris, Tate, Proc. Roy. Soc. N.S.W. 31, p. 392. 1944 Turris septemliratus Harris, Powell, Rec. Auck. Inst. Mus. 3(1), p. 8. Types — The holotype is in the British Museum (Natural History). Records — AUSTRALIA: Muddy Creek, Victoria, “Eo- cene” = Balcombian, Miocene ( type ) ; Muddy Creek, lower beds; Grice’s Creek and Altona Shaft, Victoria (Balcomb- ian). ? Turris ugaliensis Makiyama, 1927 Range — Japan, Kakegawa Series (Lower Plio- cene) and Tano (Middle Miocene). Remarks — This species should not have been described, for the type specimen consists solely of a broken shell of which only the body whorl remains. The Miocene material ascribed to this species, with reservation, by Shuto (1961, p. 79) is also imper- fect. The original author compared his species with unedo Kiener (which is an Unedogemmula ) to leucotropis ( which is a Lophioturris ) and gending- anensis Martin (which is considered to be a syn- onym of Lophiotoma ( Lophioturris ) indica (Rod- ing ) ) . No useful purpose can be served by trying to evaluate this species on the evidence at present available. Synonymy — 1927 Turris ugaliensis Makiyama, Mem. Coll. Sci. Kyoto Imp. Univ., Ser. B, 3(1), p. 93, pi. 4, fig. 18. 1952 Turris ugaliensis Makiyama, Hatai and Nisiyama, Sci. Rep. Tohoku Univ., 2nd Ser., Geol. Spec. 3, p. 263. 1961 Turris cf. ugariensis (sic) Makiyama, Shuto, Mem. Fac. Sci. Kyushu Univ., Ser. D, Geol., 11(2), p. 78. [22 - 9881 March 31, 1964 INDO-PACIFIC MOLLUSCA, vol. 1, no. 5 Antiplanes 339 Genus Antiplanes Dali, 1902 (PL 261) Type: Fleur otoma voyi Gabb, 1866 This moderately large, sinistral shell and its dex- tral forms Rectiplanes and Rectisulcus are mostly deep-water and characteristic of low temperatures. They range from off the Californian coast to Alaska and around the Bering Sea to Kamchatka and Ja- pan. Also post-Pleistocene of California. The genus could be regarded as a benthic. Boreal, cold-water development from Tunis , in which a great reduction in sculpture and the acquisition of a thick periostracum has taken place, this latter feature being typical of high latitude shellfish that have to contend with the greater erosive action of ice-diluted sea water. The sinus is broadly open, extending from the suture to below the periphery but the actual apex of the sinus is immediately above the greatest peri- pheral convexity. The shell ranges to 55 mm. (2/4 inches) in height, has a tall narrow spire but a truncated anterior end. Operculum leaf-shaped, with apical nucleus. Protoconch smooth, paucispiral and globular with an asymmetric inrolled top. Radula a pair of “wishbone”-shaped marginals plus a vestigial central tooth. Plate 261. Antiplanes voyi (Gabb, 1866). Eastern Pacific Recent, deep-water. 40 mm. ( perversa Gabb, 1865, non Philippi, 1847). Although this genus occurs off the coasts of warm temperate localities, Japan for instance, it is con- fined to the colder waters of the sea bottom, and thus cannot be considered inclusive in the Indo- Pacific fauna. Synonymy — 1902 Antiplanes Dali, Proc. U. S. National Mus., 24, p. 513. Type by original designation: Pleurotoma perversa Gabb, 1865 (non Philippi, 1847) = voyi Gabb, 1866. Characteristic Species — POST-PLIOCENE TO RECENT: voyi Gabb, 1866; contraria Yokoyama, 1928; kamchatica Dali, 1919; major Bartsch, 1944. Subgenus Rectiplanes Bartsch, 1944 (PI. 262) Type: Pleurotoma (Antiplanes) santarosana Dali, 1902 This subgenus differs from Antiplanes in being dextral instead of sinistral. Synonymy — 1944 Rectiplanes Bartsch, Proc. Biol. Soc. Washington, 57, p. 59. Type by original designation: Pleurotoma (Antiplanes) santarosana Dali, 1902. Characteristic Species — RECENT: litus Dali, 1919; rotula Dali, 1921; sanctiioannis E. A. Smith, 1875; santarosana Dali, 1902; thalaea Dali, 1902; kawamurai Habe, 1938. Subgenus Rectisulcus Habe, 1958 Type: Rectiplanes (Rectisulcus) motojimai Habe, 1958 This subgenus differs from Rectiplanes only in having spiral cords on the surface and in lacking longitudinal cords on the early whorls. Recent, Japan. It seems to have insufficient distinction from Rectiplanes. Synonymy — 1958 Rectisulcus Habe, Venus, Tokyo 20(2), p. 184. Type by original designation: Rectiplanes (Rectisulcus) motojimai Habe, 1958. Characteristic Species — RECENT : isaotakii Habe, 1958; motojimai Habe, 1958 ( both off Choshi, Chiba Pref., Honshu Id., Japan). Plate 262. Radula of Rectiplanes santarosana Dali. Eastern Pacific, Recent (prepared and drawn by J. P. E. Morrison, U.S. Nat. Mus.). [23 - 021] 340 Rectiplanes A. W. B. Powell T urridae [These occasional blank areas occur between genera and subgenera to permit the insertion of new material and future sections in their proper systematic sequence.] [23 - 022] March 31, 1964 INDO-PACIFIC MOLLUSCA, vol. 1, no. 5 Index 341 INDEX TO TURRINAE NAMES IN VOL. 1, NO. 5 Looseleaf subscribers should place this index at the begin- ning of the family Turridae. The subfamily Turrinae begins on p. [22-661]. In this index, the number following the name refers to the pagination found at the top of the page in vol. 1, no. 5. The column at the right is the looseleaf pagination. abbreviata Reeve, 307 [looseleaf] 22-917 acuta Perry, 303 22-913 acutangularis Deshayes, 244 22-696 acutirostra Conrad, 281 22-823 adela Cossmann & Pissarro, 238 22-672 adelaidensis Ludbrook, 301 22-881 adenica n.sp., 286 22-838 adolescens Harris, 283 22-825 adrastia Dali, 281 22-823 aelomitra Dali, 250 22-702 aelomitra Tinker, 250 22-702 aethiopica Thiele, 257 22-729 albicarinata Sowerby, 316 22-936 albida Perry, 316 22-936 albina Lamarck, 306 22-916 albinoides Martin, 306 22-916 albirupsis Harris, 283 22-825 Alio Jousseaume, 237 22-671 alternata Conrad, 244 22-696 amabilis Weinkauff, 261 22-733 ambages Barnard, 337 22-987 americanus Aldrich, 281 22-823 arnica Casey, 244 22-696 amichel Gardner, 282 22-824 amicta E. A. Smith, 334 22-984 ancilla Casey, 244 22-696 annae Hoernes & Auinger, 244 22-696 annulata Reeve, 333 22-983 Antiplanes Dali, 339 23-021 antwerpiensis Vincent, 244 22-696 aquensis Grateloup, 241 22-685 archimedis Bellardi, 281 22-823 aspera Edwards, 244 22-696 asukana Yokoyama, 294 22-866 Austrogemmula Laseron, 297 22-875 Austroturris Laseron, 237 22-671 babylonia Linnaeus, 327 22-977 babylonica Roding, 328 22-978 babylonius Gmelin, 328 22-978 badensis Hoernes & Auinger, 244 22-696 balteata Kiener, 241 22-685 barretti Guppy, 316 22-936 Bathybermudia Haas, 289 22-851 [looseleaf] Bathytoma, 237 22-671 284 22-826 beachportensis Cotton & Godfrey, 300 22-880 bemmeleni Oostingh, 273 22-765 benthina Dali, 279 22-791 bernayi Boury, 283 22-825 bezanconi Boury, 283 22-825 bhagothorensis Vredenburg, 265 22-737 238 22-672 bicarinatus Wood, 336 22-986 bicatenata Lamarck, 283 22-825 bilirata Boury, 284 22-826 bimarginata Suter, 266 22-738 binda Garrard, 270 22-762 birmanica Vredenburg, 264 22-736 bisculpta Powell, 318 22-948 bisinuata Martens, 289 22-851 bonneti Cossmann, 238 22-672 265 22-737 bosqueti Nyst, 244 22-696 brevicaudata Reeve, 310 22-920 bulowi Sowerby, 311 22-931 312 22-932 cainei Harris, 283 22-825 callifera Edwards, 244 22-696 Campylacrum Finlay & Marwick, 282 22-824 cancellata Deshayes, 244 22-696 Candida Laseron, 300 22-880 carinata Gray, 245 22-697 carinata Griffith & Pidgeon, 246 22-698 carinata Laseron, 301 22-881 carinata Link, 245 22-697 Carinoturris Bartsch, 281 22-823 carodenta Harris, 244 22-696 carola Thiele, 286 22-838 carya Harris, 283 22-825 carynae Haas, 289 22-851 casearia Hedley & Petterd, 285 22-837 castanella, Dali, 325 22-965 castanella Tinker, 324 22-964 casteri Harris, 244 22-696 cedilla Edwards, 283 22-825 cerithiformis Tinker, 324 22-964 childreni Lea, 281 22-823 cingulifera Lamarck, 322 22-962 Cinguliturris n.subgenus, 319 22-949 Clamturris Iredale, 322 22-962 clifdenensis Powell, 266 22-738 cochlea Harris, 283 22-825 cochlearis Conrad, 315 22-935 [22 - 651] 342 Index A. W. B. Powell T urridae cochleata Powell, 318 [looseleaf] 22-948 complicata Suter, 283 22-825 concinna Dunker, 248 22-700 congener E. A. Smith, 251 22-713 conifera Edwards, 241 22-685 conjuncta Casey, 244 22-696 constricta Edwards, 283 22-825 contabulata Deshayes, 283 22-825 contigua Brocchi, 244 22-696 contraria Yokoyama, 339 23-021 coraliger Harris, 244 22-696 corona Laseron, 320 22-950 coronata Goldfuss, 244 22-696 Coronia Ehrenberg, 281 22-823 Coronia Gregorio, 281 22-823 coronifer Martin, 254 22-716 coronifera Bellardi, 244 22-696 coronifera Martin, 254 22-716 corrugata Kiener, 241 22-685 cosmoi Sykes, 252 22-714 cossmanni Peyrot, 244 22-696 costifera Laseron, 301 22-881 crenulata Lamarck, 284 22-826 crenulosa Casey, 315 22-935 crispa Lamarck, 330 22-980 Cryptogemma Dali, 279 22-791 cryptorrhaphe Sowerby, 335 22-985 curvicosta Lamarck, 283 22-825 cypris Orbigny, 244 22-696 dampierana n.sp., 248 22-700 debile Finlay & Marwick, 282 22-824 denticula Basterot, 244 22-696 281 22-823 denticulata E. A. Smith, 286 22-838 denticulata Thiele, 286 22-838 depygis Conrad, 283 22-825 deshayesii Doumet, 270 22-762 desnoyersii Lea, 283 22-825 difficilis Giebel, 241 22-685 diomedea n.subsp., 253 22-715 disjuncta Laws, 266 22-738 disjuncta Peyrot, 244 22-696 distans Deshayes, 283 22-825 distanticosta Cossmann & Pissarro, 283 22-825 ducalis Thiele, 259 22-731 duchastelii Nyst, 241 22-685 duplex Suter, 266 22-738 Echinoturris Powell, 317 22-947 edita Powell, 295 22-871 elegans Wood, 314 22-934 Eodrillia Casey, 282 22-824 Eopleurotoma Cossmann, 282 [looseleaf] 22-824 Eoturris Finlay & Marwick, 283 22-825 Epalxis Cossmann, 284 22-826 Epideira Hedley, 238 22-672 Epidirella Iredale, 297 22-875 Epidirona Iredale, 299 22-879 erythraea Weinkauff, 323 22-963 Eugemmula Iredale, 244 22-696 expedita Deshayes, 283 22-825 explanata Koenen, 241 22-685 fagina Adams & Reeve, 334 22-984 fascialis Lamarck, 314 22-934 Fenestrosyrinx Finlay, 237 22-671 finlayi Powell, 317 22-947 flammulata Bouge & Dautzenberg, 322 22-962 flexicosta Boury, 283 22-825 flexiplicata Kautsky, 241 22-685 flindersi Cotton & Godfrey, 301 22-881 fluctuosa Deshayes, 283 22-825 formosissima E. A. Smith, 316 22-936 fortis Bartsch, 281 22-823 francisci Raincourt, 283 22-825 fresvillensis Cossmann & Pissaro, 283 22-825 fusca Hombron & Jacquinot, 262 22-734 fusiformis Thiele, 279 22-791 Fusiturris Thiele, 241 22-685 gabensis Hedley, 301 22-881 gajensis Vredenburg, 252 22-714 garnonsii Reeve, 329 22-979 gellibrandensis Chappie, 268 22-740 gemmata Reeve, 243 22-695 gemmavia Harris, 283 22-825 Gemmula Weinkauff, 243 22-695 gemmulina Martens, 260 22-732 gendinganensis Martin, 312 22-932 genitiva Casey, 281 22-823 gentilis Sowerby, 244 22-696 gilchristi Sowerby, 249 22-701 goosensi Boury, 244 22-696 gracillima Weinkauff, 331 22-981 graeffei Weinkauff, 263 22-735 grandis Gray, 331 22-981 grandis Griffith & Pidgeon, 331 22-981 granifera Deshayes, 283 22-825 granosa Helbling, 246 22-698 gryi Ravn, 244 22-696 guadurensis Melvill, 245 22-697 hastula Reeve, 272 22-764 hawleyi Iredale, 263 22-735 haydeni Vredenburg, 274 22-766 [22 - 652] March 31, 1964 INDO-PACIFIC MOLLUSCA, vol. 1, no. 5 Index 343 hedleyi Iredale, 301 [looseleaf] 22-881 Hemipleurotoma Cossmann, 281 22-823 Hesperiturris Gardner, 282 22-824 hindsiana Berry, 243 22-695 hoeninghausii Lea, 283 22-825 hombroni Hedley, 262 22-734 humilis Koenen, 244 22-696 husamaru Nomura, 238 22-672 hyugaensis Shuto, 293 22-865 ickei Martin, 274 22-766 imitatrix Martin, 254 22-716 ina MacNeil, 273 22-765 incredula Iredale, 325 22-965 inculta Sowerby, 283 22-825 indagatoris Finlay, 287 22-839 indica Deshayes, 271 22-763 indica Roding, 311 22-931 indoviciana (Vaughan) Harris, 281 22-823 inermis Hoernes, 241 22-685 inflexa Lamarck, 283 22-825 Infracoronia Palmer, 281 22-823 infraeocaenica Cossmann, 241 22-685 insensus Finlay, 283 22-825 insueta Boury, 283 22-825 intricata n.subsp., 332 22-982 invicta Melvill, 269 22-761 iravadica Vredenburg, 238 22-672 iris Vredenburg, 264 22-736 isaotakii Habe, 339 23-021 jacksonella Casey, 315 22-935 jaffaensis Verco, 301 22-881 jhirakensis Cossmann & Pissarro, 238 22-672 jickelii Weinkauff, 303 22-913 kaiparaensis Marshall, 266 22-738 kamchatica Dali, 339 23-021 karangensis Martin, 265 22-737 kawamurai Habe, 339 23-021 kieneri Doumet, 246 22-698 kingae n.sp., 325 22-965 kishimaensis Shuto & Ueda, 266 22-738 koeneni Glibert, 241 22-685 koolhoveni Oostingh, 274 22-766 kotorai Nomura & Zinbo, 265 22-737 Kuroshioturris Shuto, 293 22-865 kyushuensis Shuto, 296 22-872 laevis Bell, 301 22-881 laevis Pritchard, 301 22-881 lajonkairei Deshayes, 283 22-825 lancea Casey, 244 22-696 larteti Deshayes, 283 22-825 laticlavia Beyrich, 244 [looseleaf] 22-696 laubrierei Cossmann, 241 22-685 lawsi Powell, 266 22-738 legitima Iredale, 322 22-962 lemoinei Boury, 284 22-826 lepta Edwards, 283 22-825 lerchi Vaughan, 244 22-696 leucotropis Adams & Reeve, 312 22-932 lifuensis Sowerby, 308 22-918 lima Edwards, 283 22-825 lirata Pease, 324 22-964 lisboncola Harris, 283 22-825 litus Dali, 339 23-021 lobata Sowerby, 290 22-852 longaeva Edwards, 244 22-696 longwoodensis Powell, 267 22-739 Lophiotoma Casey, 303 22-913 Lophioturris n.subgenus, 311 22-931 Lucerapex Iredale, 285 22-837 ludocarola Harris, 244 22-696 lunulifera Koenen, 244 22-696 luzonica n.sp., 278 22-790 major Bartsch, 339 23-021 margaritata Marshall, 267 22-739 margaritosa Casey, 281 22-823 marmorata Lamarck, 311 22-931 martini Tesch, 257 22-729 mekranica Vredenburg, 252 22-714 mercati Bellardi, 241 22-685 Micantapex, 237 22-671 Micropleurotoma Thiele, 237 22-671 microsticta Casey, 304 22-914 millepunctata Sowerby, 323 22-963 miocoronifera new name, 254 22-716 molengraaffi Tesch, 287 22-839 molleri Laseron, 301 22-881 monerma Edwards, 244 22-696 monile Brocchi, 301 22-881 monile Kiener, 301 22-881 monilifera Cooper, 244 22-696 monilifera Lea, 250 22-702 monilifera Pease, 249 22-701 monilis Brocchi, 244 22-696 mordax Suter, 283 22-825 motojimai Habe, 339 23-021 multicincta Marshall, 283 22-825 multicostata Deshayes, 283 22-825 multigyrata Deshayes, 284 22-826 multinoda Lamarck, 283 22-825 multiseriata E. A. Smith, 299 22-879 murrayana Pritchard, 288 22-840 murrayi n.sp., 248 22-700 [22 - 653] 344 Index A. W. B. Powell Turridae [looseleaf] [looseleaf] narica Vredenburg, 238 22-672 Pleurotomus Montfort, 327 22-977 neglecta Suter, 283 22-825 plicaria Deshayes, 283 22-825 nilssoni Deshayes, 244 22-696 plumbella Harris, 283 22-825 nipponica Shuto, 293 22-865 polita Marshall, 267 22-739 nobilis Hinds, 316 22-936 politica Harris, 283 22-825 nodigera Koenen, 244 22-696 polycaste Dali, 281 22-823 nodocarinata Gabb, 282 22-824 Polystira Woodring, 315 22-935 nodulosa Laseron, 301 22-881 polytropa Helbling, 313 22-933 notata Sowerby, 304 22-914 porrecta Wood, 241 22-685 nupera Conrad, 283 22-825 pourcyensis Cossmann, 283 22-825 powelli Ludbrook, 301 22-881 obliterata Deshayes, 283 22-825 praesignis E. A. Smith, 259 22-731 odengensis Martin, 315 22-935 prestwichi Edwards, 241 22-685 odontella Edwards, 244 22-696 propinqua Deshayes, 283 22-825 odontophora Koenen, 244 22-696 pseudofascialis Martin, 315 22-935 okinavensis MacNeil, 277 22-789 Ptychosyrinx Thiele, 289 22-851 oligocolpa Cossmann, 283 22-825 pulchella Shuto, 255 22-717 optata Harris, 295 22-871 punctata Schubert & Wagner, 305 22-915 optata E. A. Smith, 287 22-839 Optoturris Powell, 295 22-871 quadricarinata Powell, 318 22-948 orangeburgensis Harris, 283 22-825 quentinensis Dali, 279 22-791 orba Marwick, 267 22-739 quoyi Desmoulins, 301 22-881 ornata Marshall, 267 22-739 ouachitensis Harris, 283 22-825 raffrayi Tapparone-Canefri, 327 22-977 owenii Reeve, 299 22-879 Rectiplanes Rartsch, 339 23-021 Oxyacrum Cossmann, 283 22-825 Rectisulcus Habe, 339 23-021 oxytropis Schumacher, 313 22-933 reevei Rellardi, 241 22-685 oxytropis Sowerby, 312 22-932 regilla Iredale, 285 22-837 316 22-936 regius Suter, 268 22-740 reticulata Marshall, 268 22-740 padangensis Thiele, 262 22-734 reticulosa Edwards, 244 22-696 pakistanica Eames, 265 22-737 rotatilis Martens, 264 22-736 panamensis Olsson, 316 22-936 rotella Edwards, 283 22-825 paracantha Tenison-Woods, 296 22-872 rotula Brocchi, 244 22-696 parkinsoni Deshayes, 244 22-696 rotula Dali, 339 23-021 paytensis Olsson, 283 22-825 rudiuscula Cossmann, 283 22-825 peaseana Dunker, 304 22-914 rugatina Harris, 283 22-825 perarata Cossmann, 338 22-988 rugosa Lea, 283 22-825 peraspera Marwick, 267 22-739 ruthveniana Melvill, 309 22-919 perksi Verco, 301 22-881 ryukyuensis MacNeil, 247 22-699 perversa Gabb, 339 23-021 philipineri Tenison-Woods, 301 22-881 sabinaria Harris, 283 22-825 philippinensis n.sp., 278 22-790 samueli Tenison-Woods, 268 22-740 picta Reeve, 316 22-936 sanctiioannis E. A. Smith, 339 23-021 picturata Weinkauff, 304 22-914 santarosana Dali, 339 23-021 Pinguigemmula MacNeil, 277 22-789 sanum Finlay & Marwick, 282 22-824 plana Giebel, 241 22-685 sayceana Chapman, 298 22-876 plebeia Sowerby, 244 22-696 sayi Lea, 283 22-825 plentopsis Harris, 244 22-696 250 22-702 Pleuroliria de Gregorio, 315 22-935 scalarata Edwards, 283 22-825 Pleurostoma Griffith & Pidgeon, 327 22-977 schoutanica May, 301 22-881 Pleurotoma Lamarck, 327 22-977 selandica Koenen, 283 22-825 Pleurotomidae, 231 22-665 selwyni Pritchard, 337 22-987 [22 - 654] March 31, 1964 INDO-PACIFIC MOLLUSC A, vol. 1, no. 5 Index 345 selysi Koninck, 241 [looseleaf] 22-685 septemlirata Harris, 338 22-988 serilla Dali, 279 22-791 sibogae Schepman, 258 22-730 sibukoensis n.sp., 258 22-730 similis Dautzenberg, 241 22-685 simillima Edwards, 244 22-696 simplex Casey, 315 22-935 simplicissima Thiele, 238 22-672 sindiensis Vredenburg, 255 22-717 Sinistrella Meyer, 281 22-823 sondeiana Martin, 274 22-766 soriensis Eames, 265 22-737 specialis Boury, 283 22-825 speciosa Reeve, 245 22-697 spectabilis Reeve, 336 22-986 spectabilis Weinkauff, 337 22-987 spiralis M. de Serres, 244 22-696 spreta Deshayes, 283 22-825 stoffelsi Nyst, 244 22-696 striata Gray, 301 22-881 subcarinata Rouault, 244 22-696 subconcava Harris, 319 22-949 subdentata Goldfuss, 244 22-696 subdeviata Gregorio, 315 22-935 sublerchi Harris, 244 22-696 submonilifera Boury, 244 22-696 subsimilis Casey, 315 22-935 Subulata Thiele, 289 22-851 suppressa Finlay, 301 22-881 supramirifica Gregorio, 315 22-935 taeniolata Edwards, 244 22-696 Taranis Lamy, 237 22-671 tasmanica May, 297 22-875 tatei Cossmann, 319 22-949 tellea Dali, 316 22-936 tenella Conrad, 244 22-696 tenuistriata Deshayes, 244 22-696 terebriformis Meyer, 281 22-823 teschi n.subsp., 291 22-853 thalaea Dali, 339 23-021 thielei Finlay, 279 22-791 thyroidifera Harris, 283 22-825 thyrsus Vredenburg, 264 22-736 tigrina Lamarck, 305 22-915 tigrinaeformis Nomura, 238 22-672 timorensis Tesch, 290 22-852 tizis Gregorio, 315 22-935 tomopleuroides Powell, 319 22-949 torquata Hedley, 301 22-881 torta Dautzenberg, 241 22-685 totomiensis Makiyama, 294 22-866 tricincta Martin, 265 [looseleaf] 22-737 trilineata Harris, 320 22-950 truncata Schepman, 292 22-854 trypanodes Melvill, 272 22-764 Trypanotoma Cossmann, 281 22-823 tuberculata Laseron, 301 22-881 Turridrupa Hedley, 237 22-671 turrifera Nyst, 244 22-696 Turris Muller, 327 22-977 Turris Roding, 327 22-977 Turritidae, 231 22-665 ugaliensis Makiyama, 338 22-988 ugariensis Makiyama, 338 22-988 undata Lamarck, 283 22-825 undatiruga Bivona, 241 22-685 undosa Lamarck, 334 22-984 unedo Kiener, 269 22-761 Unedogemmula MacNeil, 269 22-761 uniserialis Deshayes, 244 22-696 ustulata Reeve, 308 22-918 uttleyi Suter, 284 22-826 vagata E. A. Smith, 258 22-730 valdiviae Thiele, 257 22-729 vandervlerki Beets, 238 22-672 vardoni Tate, 301 22-881 varians Edwards, 244 22-696 variegata Kiener, 332 22-982 ventricosa Lamarck, 284 22-826 venusta Reeve, 328 22-978 Veruturris Powell, 318 22-948 vibex Dali, 316 22-936 virginoides Vredenburg, 275 22-767 virgo Wood, 316 22-936 Viridoturris n. genus, 320 22-950 voyi Gabb, 339 23-021 waihaoensis Finlay, 268 22-740 wateletella Harris, 244 22-696 waynensis Mansfield, 315 22-935 weisbordi Harris, 244 22-696 wetherelli Edwards, 241 22-685 wiedeyi Olsson, 283 22-825 woodii Kiener, 336 22-986 woodwardi Martin, 247 22-699 xanthophaes Watson, 297 22-875 Xenuloturris Kuroda, 322 22-962 Xenuroturris Iredale, 321 22-961 yeddoensis Jousseaume, 331 22-981 yenanensis Noetling, 238 22-672 zacatensis Gardner, 282 22-824 zonifera Bouge & Dautzenberg, 324 22-964 [22 - 655] 346 Index A. W. B. Powell Turridae Published by The Department of Mollusks Academy of Natural Sciences of Philadelphia 19th and the Parkway Philadelphia 3, Pennsylvania [22 - 656] April 30, 1965 INDO-PACIFIC MOLLUSCA, vol. 1, no. 6 Tridacnidae 347 Y 5' v - THE FAMILY TRIDACNIDAE IN THE INDO-PACIFIC by Joseph Rosewater Division of Mollusks United States National Museum Washington, D. C. The Giant Clams of the family Tridacnidae are highly specialized bivalves which today are re- stricted to the Indo-Pacific faunal region. The fam- ily is geologically young, having existed from Eo- cene to Recent times, and evidently having arisen from some cardiid-like ancestor in the Eocene. Only six species are living today. These bivalves are obligatory inhabitants of the shallower waters of coral reefs. As adults they are incapable of locomotion and rest with the umbos and hinge downward and the free edge of the shell uppermost. Depending upon the species they are either unattached or in burrows in the coral, tightly fastened with a byssus. When covered by the tide, the valves are opened and the mantle lobes pro- trude in two parallel, often highly colorful, undu- late bands. Life History Members of the family Tridacnidae are protan - drous hermaphrodites ( Stephenson 1934; Grobben, 1898; and Wada, 1942, 1952, 1954). Fertilization experiments with Hippopus hippopus conducted by Stephenson (ibid.) failed to produce normally de- veloping young. The adults spawned during the warmest months of the Australian summer. Wada examined the gonads of 6 species in the Palau Is- lands and found both male and female sex cells in larger individuals, He found most younger in- dividuals to be males with the hermaphroditic con- dition becoming apparent as maturity was reached. He also described the spawning reaction, appar- ently typical for the family, which is triggered by some factor contained in a suspension of the ani- Plate 263. Mantles of Tridacna Fig. 1, Tridacna ( Tridacna ) gigas (Linne). Close-up of in- halent siphon; note school of small, dark-green fish (at lower center) oriented to the respiratory current. Fig. 2, Tridacna (Chametrachea) squamosa Lamarck. Note gaping siphons and withdrawn mantle, probably indicating fatigue. Palau Islands; photo by George F. Kline. Fig. 3, Tridacna (Chametrachea) maxima (Roding). Two variations in color pattern. Both figs. 1 and 3 from Great Barrier Reef; photos courtesy of the National Geographic- Society ( copyrighted ) . [62 - 007] 348 Life History Joseph Rosewater Tridacnidae Plate 264. Valves agape showing the fleshy mantle ex- panded in Tridacna (Chametrachea) maxima (Roding). Dorsal view of living animal 150 mm. (6 inches) in length from Eni- wetok Atoll, Marshall Islands. (J. Rosewater photo from Turtox News, vol. 41, no. 12, 1963 ). mals’ eggs and which causes repeated contractions of the adductor muscle and closure of the valves coupled with the release of the sex cells for a period of an hour or more. I repeated these spawning re- action experiments successfully with Tridacna squamosa at Eniwetok, Marshall Islands, but at- tempts to observe the development of the appar- ently fertilized eggs failed when they degenerated (personal observations, February-March 1963). In nature, spawning in ripe individuals is probably triggered by an optimum temperature which sup- ports the normal embryological development, and many other factors may be involved. As yet the embryology, larvae and early development of the Tridacnidae are unknown, although Vaillant (1865), on the basis of morphology, suggested that an in- cubation period may be passed within the parent. Estimates on the duration of life in Tridacna range from 8 years (Pelseneer, 1894) to several hundred years (Comfort, 1957). No well docu- mented observations have been made on the growth rate or age attained. Bartsch (1945) suggested that Tridacna must grow at the same rate as Pacific corals, i.e., about 2 inches a year. My preliminary growth data from specimens placed in the lagoon off Eniwetok Island, Eniwetok Atoll, indicate that Bartsch’s estimate may be nearly correct for medium-sized specimens under “normal conditions” (personal observations, 1963). Specimen “A” of Tridacna gigas increased 6.7 percent in length ( 559 to 597 mm. ) in 135 days ( March 20 to August 2, 1963 ) . It increased an ad- ditional 2.3 percent in length during the next 6 months (610 mm. by February 8, 1964), thus hav- ing made a total increment of 51 mm. (9 percent) in 11.5 months. Specimen “B” of Tridacna gigas (597 mm.) and a Hippopus hippopus (254 mm.) unfortunately died at some time before the second measuring, so that the respective increments of 3 percent and 4 percent are inconclusive. If the growth rate of T. gigas “A” remained constant throughout life at 2 inches per year, the specimen measuring 610 mm. ( = 24 inches ) could be 12 years old. It is probable that rate of growth changes dur- ing the life of a Tridacna, being faster in young individuals than in older ones (cf. Comfort, ibid., p. 232). The experimental animals were placed in the lagoon near the Eniwetok Marine Biological Laboratory after having been moved a considerable distance from their original locations and were ob- served in the laboratory for several weeks. It is, therefore, doubtful that their condition was normal (two died); and it cannot be assumed that their growth proceeded naturally. Other unfavorable fac- tors are that the three measurements were made by different persons, and the second and third meas- urements were made under water. It is hoped that persons permanently located in the Indo-Pacific re- gions will undertake long-range studies of the life history and growth of the Tridacnidae so that re- liable data may be accumulated. Algal Symbiosis The specialized habit, habitat and morphology of the Tridacnidae may be considered adaptations to a singular method of feeding which has become [62 - 008] April 30, 1965 INDO-PACIFIC MOLLUSCA, vol. 1, Tridacnidae 349 , no. 6 highly developed in this group. Most lamellibranchs are ciliary feeders on minute particles of plant life present in the water. These are carried into the mantle cavity by a respiratory-feeding current gen- erated by ciliary tracts on the gills. Food is carried to the region of the mouth and sorted on the labial palps. Desirable particles are consumed and re- jected ones cast off. But the Tridacnidae do not entirely depend on food removed from the sur- rounding water for nourishment. As shown by Yonge (1936) the Tridacnidae are able to “farm” much food in their own tissues due to an unusual association with large numbers of unicellular, sym- biotic algae, called zooxanthellae (see Yonge, 1963, pp. 232-245). Zooxanthellae are known to be present in a vari- ety of marine organisms, the best known being the reef corals. They also live in association with proto- zoa, flatworms, other coelenterates and nudi- branchs, where their presence is said to be corre- lated with positive phototropism in these animals (Kawaguti, 1944; Hornell, 1909). The only other bivalve known to harbor zooxanthellae is Corculum cardissa (Linne) (Kawaguti, 1950). In species of Tridacna and to a lesser extent in Hippopus the zooxanthellae are “farmed” within the great expanse of thickened mantle edge ex- posed by the animals when they are covered by the tide. In the shallow water in which these bivalves live, the plants are provided with maximum light for photosynthesis. Special structures in the mantle surface of T ridacna only, called hyaline organs ( see pi. 265 ) , increase the translucency of the tissue, and around these the zooxanthellae are found in great- est numbers. The algae are always contained in the phagocytes of the clam, and when required for tood, the zooxanthellae are carried through blood spaces to the digestive gland. There digestion oc- curs, still within the phagocytes, and the products of digestion are probably stored in the digestive diverticula. The members of this family, therefore, have at their disposal a ready source of nourish- ment. This type of feeding is probably supple- mented by food obtained in the normal manner. However, in most species portions of the normal feeding apparatus appear to have undergone de- generation, indicating a great dependence upon the zooxanthellae. It would be of interest to learn how the original association of the giant clams with the algae came about, and how the clam receives its zooxanthellae during individual development. In the first case, Yonge (1936) has suggested that this might have occurred by chance infestations with zooxanthellae of the siphonal areas of ancestors of Tridacna. These may have been the same algae as those in- habiting coral animals, although a similar feeding relationship of corals with their algae does not ap- pear to exist (but see Goreau, 1964; also Yonge, 1963). Individuals able to utilize this additional food probably were more prone to survive. The capacity was enhanced in time through natural se- lection and exists today in its highly developed form. The clam-alga relationship may be termed a symbiotic mutualism (Allee, et al., 1949), an association which is beneficial to both parties, but in the end is at least partially detrimental to one of its members. The zooxanthellae are provided a [62 - 009] 350 Morphology Joseph Rosewater T ridacnidae place to live, raw materials from the catabolism of the clam for the manufacture of their food and spe- cial provision for intensification of sunlight for photosynthesis. Eventually they may be digested. However, the numbers consumed are probably never sufficient to deplete a colony in an individual clam. Obviously the clam benefits from this rela- tionship. It has a constant supply of living food, and its waste products are partially removed by the algae. Possibly, because of these factors, some of the species have attained an extremely large size. In order to use this special source of food to opti- mum advantage, the Tridacnidae have evolved unique morphological modifications (see Morphol- ogy). Commensalism In addition to the zooxanthellae contained in their tissues, the Tridacnidae serve as hosts to a number of other organisms which live on the sur- face of the valves, burrow into the shell, or inhabit the mantle cavity. As mentioned by Roscoe ( 1962 ) the valves are often encrusted with a wide variety of smaller coral reef denizens. In spite of the hard- ness of the shell it is common to find burrows of such bivalves as Lithophaga and Gastrochciena. Since Lithophaga burrows by chemically dissolving limy materials (Yonge, 1955; Turner and Boss, 1962) the hardness of the shell would not prevent its penetration. Holthuis ( 1952 ) listed the shrimps of the family Palaemonidae, subfamily Pontoniinae, which are known commensals with Tridacnidae. None of these species is the same as those commensal with mem- bers of the family Pinnidae (Holthuis, op. cit.; Rosewater, 1961 ) . The same three genera are in- volved, however: Paranchistus, Anchistus, and Conchodytes. Most of the species are enumerated later in this paper under their Tridacna hosts, but two, Anchistus maculatus (Stimpson), and A. spi- nuliferus (Miers), are reported from Tridacna “sp.” only. Morphology of the Tridacnidae The shell of a “normal” bivalve can be anatomi- cally oriented by holding its valves together in their naturally opposed position with the umbos upper- most and pointing away from the holder. In this position, the right side of the mollusk and its shell are to the holder’s right, and the left to the holder’s left. The umbos and hinge are uppermost or dorsal, and the free edges of the valves are below or ven- tral. The ends of the valves pointing away from the holder are the front or anterior end, and those op- posite are the hind or posterior end. The body within the valves is oriented similarly. Its mouth is anterior; the siphons are posterior; the foot pro- jects antero-ventrally from between the free edges of the valves. Orientation of the valves of the Tridacnidae is somewhat different from the procedure followed for a “normal” bivalve. Certain changes have taken place during the evolution of the giant clams in re- gard to the morphology of the shell and of the animal as a result of the special feeding relationship with zooxanthellae (Yonge, 1936; 1953a). These changes are easily understood when explained in terms of functional anatomy (see pis. 266 and 269). Natural selection for more efficient means of farming zooxanthellae has led to an increase in the size of the mantle. The original infestation with the algae probably occurred in the siphonal region, [62 - 010] April 30, 1965 INDO-PACIFIC MOLLUSCA, vol. 1, no. 6 Tridacnidae 351 Plate 267 Figs. 1, 3 Hippopus hippopus (Linne). Near Cebu City, Cebu Island, Philippines. 2 Tridacna (Tridacna) gigas (Linne). Young speci- men from Noekori Island, East Padaido Is- lands, New Guinea. 4-7 Tridacna (Chametrachea) squamosa Lamarck. Malakal Harbor, Koror Island, Palau Islands. 8 Tridacna ( Chametrachea ) crocea Lamarck. Near Zamboanga, Mindanao Island, Philippines. Figs. 9-12 Tridacna (Chametrachea) maxima (Roding). 9 Near Zamboanga, Mindanao Island, Philippines. 10 Moluccas Islands, Indonesia. 11 Gigmoto, Catanduanes Island, Philippines. 12 Near Darwin, Northern Territory, Australia. (all approximately 1/3 natural size) [62-011] 352 Morphology Joseph Rosewater Tridacnidae Plate 268. Nepionic valves of Tridacna (Chametrachea) maxima (Roding) from Raroia, Tuamotu Islands. Fig. 1, exterior. Fig. 2, interior. posteriorly located in the tridacnid ancestor. From that location they spread to surrounding mantle tis- sues. To provide additional area, and more bene- ficial lighting conditions for farming, individuals with genes for larger and more dorsally located siphonal regions were selected. During the evolu- tion of Tridacna , but not Hippopus, hyaline organs were developed and proved of value in increasing penetration of light through the mantle surface for growth of the algae. Eventually, the excurrent si- phon became located mid-dorsally, and the incur- rent siphon moved to a position high on the poste- rior end. During this forward and dorsal movement, associated mantle-siphon tissue spread over the dorsal, open edges of the shell, also spread laterally, and became considerably thickened. Because the mantle produces the shell, any change in the former structure is reflected in the latter. The umbos, hinge, and ligament have been pushed antero- ventrally by the anterior growth of the plant- farming tissues. Thus, in relation to the body of Tridacna, which is fixed by the byssus during the early stages of its existence, the mantle and shell have come to take a position so that the umbos are ventral instead of dorsal, and point posteriorly in- stead of anteriorly. The anterior hinge teeth are lost and the posterior ones are actually located anterior to the umbos. The open edges of the shell, usually ventral, are dorsal-most in position. The foot and byssus protrude from an area of the shell, once the antero-ventral slope, which is now posterior to the umbos, but still ventral in position. This area has probably undergone the least amount of “rotation” because of the presence here of the byssus which attaches to the substrate ventrally. Valves of Tridacnidae may be oriented anatomi- cally by holding them in an opposed position with the umbos down and the hinge away from the holder. In this position, the hinge end is anterior; the opposite end is posterior; the open edges of the valves are uppermost (dorsal), and the umbos and byssal orifice are down (ventral in position). The right valve is to the holder’s right and the left to the holder’s left (see plate 266). With the exception of the expansion of the man- tle-siphon tissues, the remainder of the soft anat- omy of Tridacnidae is grossly similar to that of other bivalves. There is evidence of slight depres- sion antero-ventrally of some organs, notably the ctenidia and palps, and there is forward displace- ment of the anus (Yonge, 1953b). During the “revolution” of the mantle and shell Plate 269. Diagrammatic sketch of Tridacna (Chame- to show organs of mantle cavity (after Stasek, 1962). trachea) maxima; right valve and half of mantle removed [62 - 012] April 30, 1965 INDO-PACIFIC MOLLUSCA, vol. 1, no. 6 Tridacnidae 353 which brought the normally dorsal umbos to a ventral position, the anterior adductor muscle was lost, and the anterior pedal retractor was displaced posteriorly and ventrally to its present location on the superior surface of the cardinal tooth (Stasek, 1962). There remain only two major, circular mus- cle scars in each valve. The posterior pedal or bys- sal retractor muscle scar is contiguous with and located anterior to the posterior adductor scar. Both are located somewhat subcentrally and vary in size depending on the species (see plate 269). Mantle retractor muscles are thickly produced and their scars are evident as a broad band extend- ing from the anteriorly located hinge around each valve subperipherally to a point near the posterior end of the byssal orifice (pallial line; see pi. 269). This long, broad muscle attachment enables the animal to retract the large amount of mantle-siphon tissue which is exposed to light during farming of the zooxanthellae. Stasek ( 1962, pp. 15-21, figs. 8, 9) by reference to the extinct genera Goniocardium, Avicularium and Bijssocardium, has shown how members of the Re- cent genera Hippopus and Tridacna may have evolved from a cardiid-like form to their present modified condition (see plate 270). The tridacnoid shell has “rotated” relative to the body proper in the sense that, allometrically, the postero-ventral margin of the mantle/shell greatly enlarges, thereby greatly changing its position relative to the body proper. Sawkinsia Cox, 1941, which was assigned to the Tridacnidae by Vokes [1953] seems to represent a side branch in the evolution of the family, not fit- ting into Stasek’s theory of a “gradual shift in the central tendency of succeeding populations towards more familiar ‘tridacnoid’ forms.” Geographic Distribution of Tridacnidae Recent Species — The geographic distribution of Tridacnidae almost appears to be correlated with maximum shell size and with ecology, but undoubt- edly many other factors are involved. Larger spe- cies, Tridacna gigas, T. derasa and Hippopus hip- popus, are limited to the western Pacific and Micronesia. These species, when mature, depend on weight alone to anchor them in position on the reef. Ecological requirements and possibly short larval life may confine them to their present range. Tridacna crocea, a deep burrower in coral, has a rather narrow range similar to the larger species. The range of T. squamosa, a medium-sized species, extends from central Polynesia (165° w. long.) westward to East Africa. This species depends for stabilization on its weight and the lodging of its valves, with their projecting scales, among the reef corals. A gelatinous byssus anchors T. squamosa, although not so firmly as in the burrowing species. Tridacna maxima is nearly pandemic in the Indo- Pacific, ranging from East Africa to southeastern Polynesia. This species protects itself and prevents dislodgement by boring shallow pockets in coral. Although details of its reproduction and develop- ment are unknown at present, it is suspected to have a longer larval life than other members of the family. Fossil Taxa — There are few fossil records of forms resembling Recent species in areas other than the Indo-Pacific. Tridacna media Pusch and T. wol- Plate 270. Fossil Tridacnidae. Right valves of various genera showing the gradual increase in the dorso-ventral angle as the relative length of the inferior or byssal edge increases. A, Goniocardium rachitis ( Deshayes ) . B, Avicu- larium aviculare (Lamarck). C, Avicularium cymbulare (Lamarck). D, Byssocardium emarginatum (Deshayes). E, Hippopus hippopus (Linne) from the Recent. Stems of brackets indicate 1 mm. (from C. R. Stasek, 1962, fig. 8). [62 - 013] 354 Economics Joseph Rosewater Tridacnidae farti Chenu, both from Europe, are very similar to T. maxima and to T. gigas respectively. These fossil species inhabited this area during the middle Tertiary apparently when arms of warmer seas ex- tended into northern Europe. The fossil Hippopus (?) gunteri Mansfield from the Tampa Limestone is the only Western Atlantic representative of this Recent genus. It probably inhabited reefs on the shores of the Tethys Sea during the Lower Mio- cene. Wholly extinct fossil genera and species which have been associated with Tridacnidae are all Eu- ropean with the exception of Avicularium trech- manni Cox, 1941 (see Stasek, 1962) and Sawkinsia matleyi Cox, 1941 (fide Vokes [1953, p. 121] ), both from the Eocene of Jamaica (see List of Recog- nized Taxa). Economic Importance Species of Tridacnidae are used as food in many parts of the Indo-Pacific. Tridacna maxima is the most popular species, and records of its consump- tion range from southeast Asia (Abbott, 1950) to the Tuamotus (H. A. Rehder, oral communication, 1962). The natives eat them raw or cooked, and there is some commerce in dried Tridacna flesh. Al- though some peoples probably consume the entire animal, the mantle is thick and leathery and is said to have an unpleasant flavor due to the presence of the symbiotic algae; therefore, as a rule, it is the large posterior adductor and pedal retractor mus- cles which are eaten. On Onotoa Atoll, Gilbert Islands, Hippopus hip- popus and species of Tridacna are either eaten raw, boiled with water or coconut milk, or dried and preserved with salt. The valves of Hippopus are also used as wash basins. Live specimens are some- times held in underwater pens to keep them fresh and allow them to reach a larger size (Ranner, 1952). On Arno Atoll, Marshall Islands, the natives infrequently eat Tridacna, but they do use giant clams to fertilize breadfruit trees. There they are considered less desirable to eat than fish (Hiatt, 1951). The hard shells of Tridacna are used by natives of the Pacific Islands for making various tools, such as mallets, hoes, scrapers, betel-nut pounders and ceremonial fetishes (personal communication, Mrs. R. T. Gallemore, W. Caroline Islands, 1962). Cen- trally perforated discs of Tridacna shell have been used as money in the Solomon Islands. Some of these discs reach 9 inches (228 mm.) in diameter and are 2 inches thick (personal communication, H. A. Rehder, 1964 ) . Hedley ( 1921 ) reported the use of Tridacna valves as catch basins for water on Warrior Island, Torres Strait. Non-nacreous pearls of Tridacnidae have little market value, except as curiosities and momentos, although one was valued at between 200 and 300 English Pounds in 1814 (Dillwyn, 1817, p. 215). The largest known Tridacna pearl came from a specimen of T. gigas collected in the Philippines. The “Pearl of Allah”, as the concretion came to be called, weighed 14 pounds and measured 9/2 by 5M inches. The clam which formed the pearl is sup- posed to have been responsible for the death of the diver who found it, and there is an interesting story connected with the acquisition of the pearl (Cobb, 1939 ) . A 14-pound pearl, undoubtedly the “Pearl of Allah”, was shown in a 1964 telecast and is now displayed in the shop of a Los Angeles, California jeweler (“Between the Tides,” 1963). So-called co- conut pearls of the East Indies have been traced to Tridacna (Reyne, 1947). A fairly recent economic use of the Tridacnidae is in the aquarists’ trade. Living specimens of one of the smaller species, T. maxima, are occasionally on sale in the Washington, D. C. area, and prob- ably in other cities. These specimens are imported from Singapore several times a year. An inquiry made at the store of a local dealer revealed that the Tridacna were maintained in aquaria constructed of heavy plastic which were filled with aerated arti- ficial seawater. According to the dealer, for the “Giant Killer Clam” to survive, it was necessary to include in the aquarium a piece of algae-covered coral “rock” for the clam to rest on and attach its byssus (see Nicol, 1963). Tridacna and Hippopus hippopus were kept in running seawater aquaria at the Eniwetok Marine Biological Laboratory for 5 or 6 weeks without coral “rocks.” Tridacna max- ima, a species which normally fastens tightly to coral by its byssus, always attached byssal threads to the floor of the aquaria. Specimens were re- moved, sometimes daily, for examination, and usu- ally were found to have secreted new attachments. Specimens seemed to survive well in aquaria, but near the end of the period of captivity showed signs of diminished vigor and mantle coloration (per- sonal observation). It is believed that this was due to the reduced light in the laboratory failing to support normal growth of zooxanthellae in the mantle tissues. The clams probably were slowly starving. It is not anticipated that Tridacna will be- come a common household pet. The price of a liv- ing specimen is currently about 20 dollars, and un- less given painstaking care it is doubtful that the clam would survive in captivity longer than a few [62 - 014] April 30, 1965 INDO-PACIFIC MOLLUSCA, vol. 1, no. 6 Tridacnidae 355 months. It may be of considerable interest to philatelists that species of Tridacna have been figured on post- age stamps of at least four countries of the Indo- Pacific region. Binker (1964) illustrated two of these: T. squamosa (French Somalia); T. maxima (Comoro Islands); and mentioned that a stylized T. gigas has been used on three stamps issued by the Cook Islands. A species ( maxima Roding) is shown on the 20 F stamp of French Polynesia. Probably the single most economically important use of the valves of the Tridacnidae is for their dis- play value. Anywhere in the world one is liable to see a pair of valves of Tridacna gigas ornamenting the front steps of a home, used on the lawn as a bird bath, or as a receptacle for a small flower bed. Large valves of Tridacna have been used in churches as baptismal fonts. Lamarck (1819, p. 105) mentioned this use, in the Church of Saint Sulpice in Paris, of a specimen given to Francis I by the Republic of Venice (36 inches in length; seen by R. T. Abbott, personal communication, 1962 ) . A substantial fishery exists for display speci- mens of Tridacna in the Palau Islands and the Philippines. These and many other species of mol- lusks are listed for sale, either singly or in bulk, in various shell dealers’ catalogues. If it were not for the large reproductive potential of most marine mollusks, we should look forward to the early ex- tinction of many of them in the areas where they are collected in such enormous quantities. Taxonomy of the Tridacnidae In the systematic portion of this paper genera and species are placed in what is believed to be an order of increasing phylogenetic complexity. This placement has been suggested by evaluating the primitiveness or complexity of a group or species through a consideration of the animals’ anatomy in relation to its environment and ecology. The geologic history of Recent genera gives little help in determining placement, since both Hip- popus and Tridacna arose in the early Miocene. Stasek (1962) has discussed the relationships of Recent Tridacnidae and their probable fossil an- cestors and proposed that Hippopus and Tridacna arose independently from some Byssocardium- like ancestor. He is undoubtedly correct in stating that the two arose separately, rather than the latter evolving from the former. Hippopus is placed first here because it resembles more closelv in shape the fossil Byssocardium, possesses a fully developed outer demibranch and lacks a definitive byssal ori- fice, although immature Hippopus is byssally at- tached, and there is a considerable gape in the ventral margin even in living adult specimens when they are relaxed and siphoning. Hyaline organs are lacking in Hippopus, which also may indicate a primitive situation. Zooxanthellae are present be- neath a fairly transparent mantle surface which may be a substitute for the lens-like structures found in Tridacna. The genus Tridacna sensu lato is the only other major taxon in the family. On the basis of its anat- omy and ecology, Tridacna gigas, the type-species of Tridacna sensu stricto, is the least specialized species of the genus. It has a complete outer demi- branch and a small byssal orifice. Adult individuals live unattached on sandy bottoms and attain a very large size ( 1370 mm. ) . Hyaline organs are very numerous in the mantle surface. Tridacna derasa differs sufficiently from other Recent species to justify the acceptance of the sub- genus Persikima for it. This species is considered intermediate between T. (T.) gigas and the remain- ing species because it attains a considerable size (514 mm. ), has a very small byssal orifice and lives, in adulthood, unattached on the reef. Peculiar mor- phological characteristics of the subgenus are the posterior displacement of the umbos and the prob- ably related anterior growth of the ctenidia which, in addition, lack complete outer demibranchs. Other living tridacnids are similar in lacking com- plete outer demibranchs and the associated food groove, although the latter structure is present on the complete inner demibranch of all species. This structure may have been lost with the increase in efficiency of algal feeding. The latter specialization parallels a tendency to bore deeper into coral. Tridacna squamosa lives attached to coral rubble by weak, mucus-covered byssal strands. Tridacna maxima excavates shallow depressions in coral boulders, or occurs nearly at the surface of the sub- strate, but is always tightly attached by a stout byssus. Tridacna crocea is usually deeply imbedded in coral and is also tightly anchored. For the above reasons, the last 3 species are placed in the sub- genus Chametrachea. Taxonomic distinctions nearly always can be made among these species on the basis of the shells or the anatomy. But such distinctions are helped by and are closely related to the ecology of the species. Mantle-color distinguishes some species at some localities (see individual species descriptions). At Eniwetok, H. hippopus, T. gigas, , T. squamosa and T. maxima were clearly separable on the basis [62 - 015] 356 Recognized Taxa Joseph Rosewater Tridacnidae of mantle-color and pattern, although variation was considerable, especially in the last two species. In the Andaman Sea and Mentawai Islands, T. maxima and T. crocea were observed alive and few, if any, distinctions were noted in the colors and patterns of their mantles. Tridacna gigas and H. hippopas were relatively stable in color and pattern where they were seen alive in these two widely separated regions (personal observations in the Malayan Archipelago and Marshall Islands, 1963). Spurious Names in Tridacnidae 1773 Concha indica Buonnani, Rerum Naturalium Historia in Mus. Kircheriano, Rome, part 2, p. 46, pi. 13, figs. 80, 81. Prashad (1932) in the Pelecypoda section of the Siboga Expedition, monograph 53c, gave extensive synonymies, but included a number of pre-Linnaean and non-binomial names as if they were valid names. Concha imbricata Rumphius, 1741; Chama aspera Regenfuss, 1748; Chama striata Petiver, 1767, are examples of names which do not belong in binomial literature. 1797 Tridachna multifolia Humphrey, Museum Calonnia- num, p. 50 (China); rejected work; name unas- signable to any species. 1798 Tridachnes arcinella Roding, Museum Boltenianum, part 2, p. 172, no. 98 [198]; refers to Gmelin [1791, Systema Naturae, ed. 13, p. 3303] species 14, Mar- tini [Chemnitz, vol. 7, pi. 52] figs. 522, 523, and to Knorr [Vergniigen der Augen und des Gemuths, etc., part 6] pi. 36, figs. 1, 2 [= Echinochama arcinella (Linne) from the tropical Western At- lantic]. 1819 Tridacna pustulosa Lamarck, Histoire Naturelle des Animaux sans Vertebres, vol. 6, part 1, p. 107 [= Productus giganteus Sowerby, 1822, a branchio- pod, fide Deshayes 1836, ibid., ed. 2, vol. 7, p. 385, and Lamy, 1932, Bull. Mus. National d’Histoire Naturelle, series 2, vol. 4, no. 3, p. 308]. The first two of the three following names of F. S. Voigt were listed under Tridacna by Sherborn (1933, part 33, p. 1047). The third name is listed here, since Voigt did not credit it to any other author, although as in the cases of the first two names it is ascribable to Lamarck. For some reason Voigt prefixed with the letters “Tr.” [= Tridacna] not only species belonging to that family, but also members of the Chamidae and Isocardiidae. 1834 Tridacna (Chama) asperella F. S. Voigt (in Cuvier) Das Thierreich, Leipzig, vol. 3, p. 508 [= Chama asperella Lamarck, 1819]. 1834 Tridacna (Chama) crenulata F. S. Voigt, ibid. [= Chama crenulata Lamarck, 1819]. 1834 Tridacna (Isocardia) moltkiana F. S. Voigt, ibid. [= Isocardia moltkiana Lamarck, 1801], 1856 Tridacna latissima Bianconi, Memorie della Accademia delle Scienze dell Istituto di Bologna, vol. 7, p. 409 (name only, not figured) [nomen nudum], 1932 Hippopus pennicornis Sherborn, Index Animalium, section 2, part 31, p. 564 [listed in error as Hip- popus; see ibid, part 19, p. 4835 = Hippopsis pen- nicornis, Insecta], 1933 Tridacna punctulosa Sherborn, ibid, part 33, p. 1047 [error for T. pustulosa Lamarck, 1819, q. v.]. List of Recognized Taxa in the Tridacnidae Below are fisted the recognized taxa of fossil and Recent Tridacnidae. Only members of the genera Tridacna and Hippopus are treated in full in the main text. A dagger f precedes fossil genera and species. Genus f Goniocardium Vasseur, 1880, Middle and Upper Eocene f rachitis (Deshayes, 1829). Type. Middle Eo- cene, France f heberti Vasseur, 1880. Eocene, France Genus f Avicularium Gray, 1853. Middle Eocene — Lower Ofigocene f aviculare (Lamarck, 1805). Type. Middle Eo- cene, France f cymbulare (Lamarck, 1819). Middle Eocene, France \trechmanni Cox, 1941. Upper Eocene, Jamaica Genus f Byssocardium Munier-Chalmas, 1882. Up- per Eocene — Lower Miocene \emarginatum (Deshayes, 1829). Type. Upper Eocene, France f andreae Tournouer, 1882. Lower Miocene, France Genus f Sawkinsia Cox, 1941. Upper Eocene \matleyi Cox, 1941. Type. Upper Eocene, Jamaica Genus Hippopus Lamarck, 1799. Lower Miocene — Recent hippopus (Linne, 1758). Type, f gunteri Mansfield, 1937. Lower Miocene, Florida Genus Tridacna Bruguiere, 1797. Lower Miocene — Recent Subgenus Tridacna s.s. ]wolfarti Chenu, 1845. Lower Miocene, Ger- many gigas (Linne, 1758). Type Subgenus Persikima Iredale, 1937. Recent derasa (Roding, 1798). Type Subgenus Chametrachea Morch, 1853. Pliocene — Recent f loczyi Kutassy, 1934. Pliocene, Indonesia f mbalavuana Ladd, 1934. Neogene, Fiji f aegyptiaca Chenu, 1845. Pliocene, Egypt squamosa Lamarck, 1819 f besairiei Collignon, 1951. Upper Cretaceous (?), Madagascar f media Pusch, 1837. “Tertiary,” Poland maxima (Roding, 1798) crocea Lamarck, 1819. Type [62 - 016] April 30, 1965 INDO-PACIFIC MOLLUSCA, vol. 1, no. 6 Tridacnidae 357 Selected Bibliography Abbott, R. Tucker, 1950. The Molluscan Fauna of the Cocos-Keeling Islands, Indian Ocean. Raffles Museum, Singapore, Bull. no. 22, pp. 68-98. Abrard, Rene. 1942. Mollusques Pleistocenes de la Cote Frangaise des Somalis. Archives du Museum National d’Histoire Naturelle, series 6, vol. 18, 105 pp., 8 pis. Allan, Joyce, 1959. Australian Shells, Revised edition, Geor- gian House, Melbourne, xx + 487 pp. Allee, W. C., A. E. Emerson, O. Park, and K. P. Schmidt. 1949. Principles of Animal Ecology. W. B. Saunders, Phila., xii + 837 pp. Banner, A. H. 1952. Preliminary Report on Marine Biology Study of Onotoa Atoll, Gilbert Islands. Atoll Research Bulletin, no. 13, part 1, pp. 1-42. Barnard, K. H. 1964. Contributions to the Knowledge of South African Marine Mollusca. Part V, Lamellibranchi- ata. Annals South African Museum, vol. 67, part 3, pp. 361-593. Bartsch, Paul, and John T. Nichols. 1945. Fishes and Shells of the Pacific World. The Macmillan Co., New York, 201 pp. Binker, Elmer J. 1964. Shell Collecting — Philatelic Style. Shells and their Neighbors, no. 23, pp. 1-4. Buchsbaum, Ralph and Lorus J. Milne. 1960. The Lower Animals; Living Invertebrates of the World. Doubleday and Co., Inc., New York, 303 pp. Cobb, W. D. 1939. The Pearl of Allah. Natural History, New York, vol. 44, no. 4, pp. 197-202, text figs. Comfort, A. 1957. The Duration of Life in Mollusks. Pro- ceedings Malacological Society of London, vol. 32, part 6, pp. 219-241. Cox, L. R. 1941. Lamellibranchs from the White Limestone of Jamaica. Proceedings Malacological Society of London, vol. 24, part 4, pp. 135-144. Dali, W. H. 1900. Transactions of the Wagner Free Institute of Science of Philadelphia, vol. 3, part 5, pp. 949-1218. Dautzenberg, Ph. 1923. Liste prehminiaire des Mollusques Marins de Madagascar, et Description De Deux Especes Nouvelles. Journal de Conchyliologie, vol. 68, pp. 21-74. 1929. Mollusques Testaces Marins de Madagascar. Mollusca II. Faune des Colonies Frangaises, vol. 3, pp. 321-636, pis. 4-7. 1932. Mollusques Testaces Marins de Madagascar; Supplement. Journal de Conchyliologie, vol. 76, p. 101. Deshayes, G. P. 1863. Catalogue des Mollusques de L’He de La Reunion ( Bourbon ) . Revue des Deux Mondes, Paris, pp. 1-44, pis. 28-41. Dillwyn, L. W. 1817. A Descriptive Catalogue of Recent Shells, vol. 1, p. 215. Dodge, Henry. 1947. The Molluscan Genera of Bruguiere. Journal of Paleontology, vol. 21, no. 5, pp. 484-492. 1952. A Historical Review of the Mollusks of Lin- naeus. Part I, The Classes Loricata and Pelecypoda. Bul- letin American Museum of Natural History, vol. 100, article 1, pp. 1-263. Franc, Andre. 1960. Classe des Bivalves. Traite de Zoologie, Tome 5, fascicle 2, pp. 1845-2133. Gillett, Keith and Frank McNeill. 1959. The Great Barrier Reef and Adjacent Isles. Coral Press Pty. Ltd., Sydney, xiii + 194 pp. Goreau, Thomas F. 1964. Mass Expulsion of Zooxanthellae from Jamaican Reef Communities after Hurricane Flora. Science, vol. 145, pp. 383-386. Grobben, Carl. 1898. Expedition S. M. Schiff “Pola” in das Rothe Meer. VIII. Zoologische Ergebnisse. Beitrage zur Morphologie und Anatomie der Tridacniden. Denkschrif- ten der Mathematisch-Naturwissenschaftleichen Classe der Kaiserlichen Akademie der Wissenschaften, Wien, vol. 65, pp. 433-444, 3 pis. Hedley, C. 1899. The Mollusca of Funafuti. Part II. Pelecy- poda and Brachiopoda. Memoirs Australian Museum, vol. 3, part 7, pp. 397-565. 1921. A Revision of the Australian Tridacna. Rec- ords Australian Museum, Sydney, vol. 13, pp. 163-172. 1923. The Largest Hippopus. Ibid., vol. 14, p. 138. Hiatt, R. W. 1951. Marine Zoology Study of Amo Atoll, Marshall Islands. Atoll Research Bulletin, no. 4, pp. i-ii + 1-13. Holthuis, L. B. 1952. The Decapoda of the Siboga Expedi- tion, Part II. The Palaemonidae. Monograph 39a, section 10, Siboga Expeditie, pp. 1-253, 110 figs. Homell, James. 1909. A Note on the Presence of Symbiotic Algae in the Integuments of Nudibranchs of the Genus Melibe. A Report to the Government of Baroda on the Marine Zoology of Okhamandal, vol. 1, pp. 145-148. Iredale, T. 1937. Middleton and Elizabeth Reefs, South Pacific Ocean. Mollusca. Australian Zoologist, vol. 8, part 4, pp. 232-261, pis. xv-xvii. Iredale, T. and Joyce Allan. 1940. A reviewer of the Rela- tionships of the Mollusca of Lord Howe Island. Ibid., vol. 9, part 4, pp. 444-451. Johnson, C. W. 1930. The Giant Clam (Tridacna gigas). The Nautilus, vol. 43, no. 4, p. 141. Kawaguti, Siro. 1944. Zooxanthellae as a Factor of Positive Phototropism in those Animals containing them. Palao Tropical Biological Station Studies, vol. 2, no. 4, pp. 681-682. 1950. Observations on the Heart Shell, Corculum cardissa (L.), and its associated Zooxanthellae. Pacific Sci- ence, vol. 4, no. 1, pp. 43-49, 8 figs., 2 tables. Keen, A Myra. 1937. Nomenclatorial Units of the Pelecypod Family Cardiidae. Bulletin Musee Royal d’Histoire Naturelle de Belgique, vol. 13, no. 7, pp. 1-22. Kellogg, James L. 1915. Ciliary Mechanisms of Lamelli- branchs with Descriptions of Anatomy. Journal of Mor- phology, vol. 26, no. 4, pp. 625-701. Kelly, K. L. and D. B. Judd. 1955. The ISCC-NBS Method of designating colors and a dictionary of color names. Na- tional Bureau of Standards Circular 553, U. S. Govern- ment Printing Office, Washington, D. C., 158 pp. Ladd, Harry S. 1934. Geology of Vitilevu, Fiji. Bernice P. Bishop Museum, Bulletin no. 119, pp. 1-263, pis. 1-44. Lamy, E. 1932. Notes sur les especes Lamarckiennes de Tridacnidae. Bulletin Museum d’Histoire Naturelle, Paris (2), vol. 4, no. 3, pp. 307-312. 1939. Revision des Tridacnidae vivants du Museum National d’Histoire Naturelle de Paris. Journal de Conchy- liologie, vol. 83, no. 4, pp. 273-302, 2 text figures. [62-021] 358 Bibliography Joseph Rosewater Tridacnidae Macnae, W. and Kalk, M. 1958. A Natural History of Inhaca Island, Mozambique. Witwatersrand University Press, Johannesburg, vi+163 pp. Mansfield, Wendall C. 1926. Fossils from Quarries near Suva, Viti Levu, Fiji Islands, and from Vavao, Tonga Islands. Carnegie Institution of Washington, Publication no. 344, pp. 85-104. Martens, E. von. 1880. Beitrage zur Meeresfauna der Insel Mauritius und der Seychellen. Berlin, pp. 181-352, pis. 19-22. McLean, R. A. 1947. A Revision of the Pelecypod Family Tridacnidae. Notulae Naturae, Philadelphia, no. 195, pp. 1-7. Melvill, J. C. and R. Standen. 1907. The Mollusca of the Persian Gulf, Gulf of Oman, and Arabian Sea, as evi- denced mainly through the Collections of Mr. F. W. Townsend, 1893-1906, with Descriptions of New Species. Proceedings Zoological Society of London (1906), pp. 783-848, 4 pis. Melvill, J. C. 1909. Report on the marine Mollusca obtained by Mr. J. Stanley Gardiner, F. R. S., among the islands of the Indian Ocean in 1905. Transactions Linnean So- ciety of London, 2nd series, Zoology, vol. 13, part 1, p. 135. Miner, Roy Waldo. 1938. On the Bottom of a Pearl Lagoon. The National Geographic Magazine, vol. 74, no. 3, pp. 365-382, color plates, 1-8. Nakazima, Masao. 1964. On the differentiation of the cre- nated-folds in the midgut-gland of Eulamellibranchia V. Crenated-folds in the two species of Tridacnidae. Venus, vol. 39 (7), pp. 67-71, pis. 4, 5. Nicol, D. 1963. Further Comments on Biotic Associations and Extinction. Systematic Zoology, vol. 12, no. 1, pp. 38-39. Nomura, S. and Zinbo, N. 1935. Fossil and Recent Mollusca from the Island of Kita-Daito-Zima. Science Reports Tohuko University, vol. 18, no. 1, pp. 41 ( i ) -51 ( ii ) , pi. v(i). Ostergaard, Jens M. 1935. Recent and Fossil Marine Mol- lusca of Tongatabu. Bernice P. Bishop Museum Bull. 131, 59 pp., 1 pi. Pelseneer, Paul. 1894. Introduction a L’Etude des Mollus- ques. Memories de la Societe Royale Malacologique de Belgique, vol. 27 (1892) pp. 31-243. Pilsbry, H. A. 1895. Catalogue of the Marine Mollusks of Japan. Frederick Steams Publ., Detroit, pp. viii + 196, 11 pis. Prashad, B. 1932. The Lamellibranchia of the Siboga Ex- pedition. Systematic Part II, Pelecypoda (Exclusive of the Pectinidae). Siboga-Expeditie 53c, E. J. Brill, Leiden, 353 pp. 19 pis. Purchon, R. D. 1955. A Note on the Biology of Tridacna crocea. Proceedings Malacological Society of London, vol. 31, pp. 95-110, 6 text figs. Rees, W. J. and A. Stuckey. 1952. The ‘Manihine’ Expedi- tion to The Gulf of Aqaba. VI. Mollusca. Bulletin British Museum (N. H.), Zoology, vol. I. (8), pp. 183-201, 3 pis. Reyne, A. 1947. On the Structure of the Shells and Pearls of Tridacna squamosa Lam. and Hippopus hippopus (L). Archives Neerlandaises de Zoologie, vol. 8, pp. 206-241, text figs., pi. 8. Roberts, John. 1963. Collecting Tridacna on Eniwetok. Ha- waiian Shell News, vol. 11, no. 7, p. 4. Roscoe, E. H. 1962. Some Records of large Tridacna speci- mens. Hawaiian Shell News, vol. 11, no. 1, p. 8. Rosewater, Joseph. 1961. The Family Pinnidae in the Indo- Pacific. Indo-Pacific Mollusca, vol. 1, no. 4, pp. 175-226. 1963. An Effective Anesthetic for Giant Clams and Other Mollusks. Turtox News, vol. 41, no. 12. pp. 300- 302. Rost, Helen and Tron Soot-Ryen. 1955. Pelecypods from the Seychelles Islands collected by Mr. W. v. Hasselberg. Acta Borealia, A. Scientia, no. 8, Tromso Museum, pp. 1-23. Shells in the News. 1963. Between the Tides. Newsletter of Gulf Coast Shell Club, Beaumont Texas, no. 8, p. 1, Nov. 1963 (mimeographed). Smith, E. A. 1898. Note on a very large specimen of Hip- popus hippopus and on large specimens of Tridacna gigas. Proc. Malacological Society of London, vol. 3, pp. Ill, 112. 1903. Marine Mollusca: In: Fauna and Geography of the Maidive and Laccadive Archipelagoes, vol. 2, part 2, pp. 589-630, pis. 35, 36. Edited by J. S. Gardiner, Cambridge, England. 1913. On a small collection of Marine Shells from Henderson Island. Annals and Magazine of Natural His- tory, ser. 8, vol. 12, pp. 409-415, 1 pi. Stasek, Charles R. 1962. The form, growth, and evolution of the Tridacnidae (Giant Clams). Archives de Zoologie Experimentale et Generale, Tome 101, fasc. 1, pp. 1-40, 14 figs. Stephenson, Anne. 1934. The breeding of reef animals, part 2, Invertebrates other than corals. Scientific Reports, Great Barrier Reef Expedition, 1928-1929, vol. 3, no. 9, pp. 247-272. Stephenson, T. A; A. Stephenson, G. Tandy, and M. Spender. 1931. The Structure and Ecology of Low Isles and other reefs. Ibid., vol. 3, no. 2, pp. 17-112, 27 pis. Turner, R. D. in: Shrock, R. R. and W. H. Twenhofel. 1953. Principles of Invertebrate Paleontology. 2nd Edition. Mc- Graw-Hill, New York, xx + 816 pp. Turner, R. D. and Kenneth J. Boss. 1962. The Genus Litho- phaga in the Western Atlantic. Johnsonia, vol. 4, no. 44, pp. 81-116. Vaillant, L. 1865. Recherches sur la famille des Tridacnides. Annales des Sciences Naturelles, Zoologie, series 5, vol. 4, pp. 64-172. pis. 8-12. van der Vlerk, I. M. 1931. Caenozoic Amphineura, Gastro- poda, Lamellibranchiata, Scaphopoda. Leidsche Geolo- gische Mededeelingen, Deel 5, pp. 206-296. Vokes, H. E. 1951-1955. Preliminary Classification of the Genera of the Pelecypoda ( an unpublished mimeo- graphed compilation). Wada, S. K. 1942. Notes on the Tridacnid Clams in Palau. Kagaku-Nanyo ( Science in the South Seas ) , vol. 5 ( 1 ) , pp. 62-69 (in Japanese). 1952. Protandric functional hermaphroditism in Tri- dacnid clams. The Oceanographical Magazine, Tokyo, vol. 4 (1), pp. 23-30. 1954. Spawning in the Tridacnid Clams. Japanese Journal of Zoology, vol. 11, pp. 273-285. [62 - 022] April 30, 1965 INDO-PACIFIC MOLLUSCA, vol. 1, no. 6 Tridacnidae 359 Yonge, C. M. 1936. Mode of Life, Feeding, Digestion and Symbiosis with Zooxanthellae in the Tridacnidae. Scien- tific Reports, Great Barrier Reef Expedition, 1928-29, vol. 1, no. 11, pp. 283-321. 1953a. Mantle Chambers and Water Circulation in the Tridacnidae. Proceedings Zoological Society of Lon- don, vol. 123, pp. 551-561, 5 figs. 1953b. The Monomyarian condition in the Lamelli- branchia. Transactions Royal Society of Edinburgh, vol. 62, no. 12, pp. 443-478, 13 figs. 1955. Adaptation to Rock Boring in Botula and Lithopliaga ( Lamellibranchia, Mytilidae) with a Discus- sion on the Evolution of this Habit. Quarterly Journal of Microscopical Science, vol. 96, part 3, pp. 383-410. 1963. The Biology of Coral Reefs. Advances in Marine Biology, vol. 1, Academic Press, London and New York, pp. 209-260. Acknowledgements A number of persons and institutions contributed toward making this study more complete. Many of them are acknowledged in the text. Thanks are due members of the Hawaiian Malacological Society for aiding in many ways. Drs. H. S. Ladd, Norman F. Sohl, and W. M. Briggs, Jr., U. S. Geological Survey, suggested ages for fossil records from re- ports in progress on Pacific mollusks, and discussed fossil species. Mr. Kenneth L. Kelly, National Bu- reau of Standards, loaned manuscript color refer- ences for the work on Tridacna mantle coloration. The following provided working space and showed every kindness during visits to study museum col- lections: Drs. Yoshio Kondo, BPBM; R. T. Abbott, R. Robertson, Virginia Orr Maes, ANSP; W. J. Clench, R. D. Turner, MCZ; Norman Tebble, BM. Drs. H. A. Rehder and J. P. E. Morrison, USNM, gave helpful suggestions and advice throughout the study. Permission has been obtained to publish this pa- per from the Secretary of the Smithsonian Institu- tion. The U. S. Atomic Energy Commission, through Drs. I. E. Wallen and R. W. Hiatt, arranged for the writer to work at the Eniwetok Marine Biological Laboratory. Dr. Hiatt and EMBL scientific assist- ants, G. A. Hiatt and J. J. Naughton, made meas- urements for the growth study. A portion of this work was supported by the National Science Foun- dation as a part of the U. S. Program in Biology, International Indian Ocean Expedition. Key to the Recent Tridacnidae 1. Opposed valves with well defined byssal orifice; byssal orifice without tightly fitting teeth Tridacna (2) Opposed valves without well defined byssal orifice; area of bys- sal orifice with tightly fitting teeth . . . Hippopus hippopus 2. Concentric sculpture lacking or limited to a few tubular spines near umbos; shell equilateral; interdigitating proc- esses elongate-triangular; radiating sculpture deeply folded Tridacna gigas Concentric sculpture present or absent; shell usually inequi- lateral; interdigitating processes bluntly triangular or rounded; radiating sculpture broad and low (3) 3. Umbos nearer anterior end or nearly central; concentric sculp- ture projecting, bladelike, or appressed and low (4) Umbos nearer posterior end; concentric sculpture absent or in low erect ridges Tridacna derasa 4. Concentric sculpture projecting, bladelike; valves nearly equi- lateral Tridacna squamosa Concentric sculpture appressed and low; valves inequi- lateral (5) 5. Posterior pedal retractor and adductor muscle scars subequal Tridacna maxima Posterior pedal retractor muscle scar less than half the size of posterior adductor muscle scar Tridacna crocea [62 - 023] 360 The Author Joseph Rosewater Tridacnidae Smithsonian Institution photo Dr. Joseph Rosewater is Associate Curator in the Division of Mollusks, U. S. National Museum, in Washington, D. C. His research has been in both marine and fresh-water mollusks. This is his second contribution to Indo-Pacific Mollusca, the first being on Pinnidae. Dr. Rosewater was born in Claremont, New Hampshire, on September 18, 1928, got his R.S. and M.S. at the University of New Hampshire, and his Ph.D. under Dr. W. J. Clench at Harvard Uni- versity. He is married and has three children. His field work in the Indo-Pacific area has taken him to southeast Asia where he participated in the Interna- tional Indian Ocean Expeditions, and to the Marine Biological Station at Eniwetok, Marshall Islands, in 1963, where he studied live Tridacna. [62 - 024] April 30, 1965 INDO-PACIFIC MOLLUSCA, vol. 1, no. 6 Hippopus 361 Family Tridacnidae Goldfuss, 1820 Genus Hippopus Lamarck, 1799 Type: Hippopus hippopus (Linne, 1758) The genus Hippopus is now limited to the Indo- Pacific region and is easily recognized by the elon- gate-triangular outline and strawberry spots on the shell of the single Recent species H. hippopus. In maturity and old age the shells usually become heavy and thickened, lose many distinguishing characters, and may reach a rather large size (397 mm. or 15.4 inches ) . Hippopus differs from Tridacna in the lack of hyaline organs in the dorsal exposed part of the mantle. The posterior adductor muscle scar is lo- cated centrally in the valves, while in Tridacna it is slightly posterior. There is a tendency for the hinge margin to become longer relative to the ventral margin in old Hippopus. Both inner and outer demi- branchs of the ctenidia are complete. Synonymy — [1787 Hippopodes Meuschen, Museum Geversianum, p. 428; a plural noun and also, a non-binomial work.] 1799 Hippopus Lamarck, Memoires de la Societe d’His- toire Naturelle de Paris, p. 86. Type by absolute tautonymy: Chama hippopus Linne, 1758. [1807 Hippopigenus Renier, Tavole per service alia Classi- ficazione e Connoscenza degle Animali, tav. 7 (re- jected name).] 1811 Pelvis, Megerle, Gesellschaft naturforschender Freunde, Berlin, vol. 4, p. 67. Type by monotypy; Pelvis hippopus [= Hippopus hippopus ( Linne, 1758)]. 1848 Cerceis Gistl, Naturgeschichte des Thierreichs fur hohere Schulen, Stuttgart, p. 172 (new name for Hippopus Lamarck, 1799 ) . Hippopus hippopus (Linne, 1758) (Pis. 267, 270, 271) Range — Malay Peninsula to eastern Melanesia. Remarks — The Horse’s Hoof, Bear Paw or Straw- berry Clam is one of the best known species in the Indo-Pacific faunal region. A combination of char- acters, which include its distinctive coloration, sculpture, size and shape, prevent the confusion of most small- and medium-sized Hippopus hippopus with any other species. In shells of adults of this species the byssal ori- fice, which gapes slightly in the young, loses its identity and its edges become tightly closed with interlocking teeth. This change may be correlated with a loss of the byssal attachment in adult speci- mens. Large animals of Hippopus (310 mm. in length) have no byssus, but smaller specimens (145 mm.) often attach byssal strands to coral rubble. The need for attachment probably diminishes as the animal attains a larger size and greater weight. Statements regarding the ease of recognition of medium-sized specimens of H. hippopus do not al- ways hold true when the species reaches old age. Depending on conditions under which the animals live, individuals apparently may reach a consider- ably larger size than was previously suspected. Smith ( 1898 ) reported what he considered to be an unusually large and thick specimen from the Philippines, 1311 inches long (336 mm.) and weigh- ing 16/2 pounds. According to Smith, this specimen varied from the “typical” H. hippopus because it had a peculiar posterior [anterior?] elongation of the valves causing the umbos to become less cen- trally located (i.e., they were nearer the posterior end ) and unusually incurved. Hedley ( 1923 ) de- scribed a large specimen of H. hippopus which he and H. A. Pilsbry collected from Pandora Reef, near Palm Islands, off Queensland. This specimen was 15/8 inches long (397 mm.) and weighed 28 lbs., 9 oz. In the same paper Hedley reported he had heard of an even larger specimen measuring over 18 inches in length (457 mm.), although I suspect the latter specimen may possibly have been confused with T. derasa, which may resemble hippopus. Some populations from the southern Philippines have a high proportion of a smooth, rather thin- shelled form which if seen separately can cause spec- ulation regarding the existence of a second living species or subspecies of Hippopus. This form inter- grades completely with H. Hippopus and does not deserve a name. Habitat — Hippopus lives on a sandy substrate in coral reef waters down to 20 feet in depth. Adults are unattached, but young specimens are found byssally attached to coral heads. Color notes — The mantle ( in a living specimen at Eniwetok Atoll) is irregularly mottled along its edges and in the center with deep yellowish green (Kelly and Judd, 1955, ISCC-NBS color name; C553 number: 132) which appears to be caused by ag- gregations of zooxanthellae. Between mottlings are interspersed areas of somewhat non-descript color ranging between moderate olive and light grayish [62 - 031] 362 Hippopus Joseph Rosewater Tridacnidae olive (C553 numbers: 107-109) with some whitish blotching. Narrow white vermiculate markings are often present. Under a dissecting microscope, these are seen to be made up of iridescent white granules. The mantle is quite translucent, and this condition may function as the hyaline organs present in Triclacna in allowing light to penetrate for growth of the zooxanthellae. Activity — Reactions to stimuli administered in the laboratory were similar to those observed in other species of Tridacnidae (see observations un- der T. gigas). When undisturbed the animals rested on the bottom of a large cement tank in running seawater with the valves open and the mantle tis- sues filling the space between (see plate 271, figs. 1, 2). The excurrent siphon was held in a flattened- Plate 271. Hippopus hippopus (Linne). Figs. 1, 2. Dorsal views of living animal from Eniwetok, Marshall Islands. Note plate-like excurrent siphon and mantle pattern. Figs. 3, 4. Exterior and interior views of left and right valves, re- spectively (97.2 mm. in length). Specimen from Zoological Museum. Copenhagen, figured by Schumacher, 1817, pi. 12. fig. 2 (Figs. 1, 2, author’s photos; figs. 3, 4, courtesy of R. T. Abbott). [62 - 032] April 39, 1965 INDO-PACIFIC MOLLUSCA, vol. 1, no. 6 Hippopus 363 disclike position, the aperture circular. The incur- rent siphon was open, but its right and left edges partially obscured the aperture with blunt tentacle- like projections. When disturbed by changes in light intensity, vibration, or prodding, the animal was capable of rapid contraction and closing of its valves. Since the mantle edges are not protruded over the edges of the valves, Hippopus is able to close more rapidly than species of Tridacna. Description — Shell reaching 397 mm. ( 15% inches, Hedley, 1923) in length; elongate- to ovate- triangular or sub-rhomboidal in shape; usually strongly inflated; with valves opposed, the byssal orifice is tightly closed in large specimens or only slightly gaping in young ones. Valves heavy, thick, usually colored with strawberry blotches arranged in irregular concentric bands, often obscured by in- crusting vermetids, annelid tubes, coral, coralline algae, other organisms and debris; color otherwise grayish white with yellowish orange tinges. Surface of valves dull or faintly shining in cleaned speci- mens. Primary radial sculpture consisting of 13 or 14 moderately convex rib-like folds over surface of valve, extending on to ventral slope where they become obsolete. Secondary radial sculpture con- sisting of low riblets on both primary folds and their interstices. Riblets varying somewhat in width and sometimes bearing rows of small- to moder- ately-sized, semi-tubular spines. When present, spines protrude in all directions giving valves a highly prickly appearance. Concentric sculpture consisting of fine, wavy lines of growth. Dorsal mar- gin undulate, with series of 8 to 12 squarish, me- dially projecting interdigitating processes repre- senting extremities of rib interstices. Hinge line as long or longer than half the length of valve. One oblong cardinal tooth in each valve; 2 elongate pos- terior laterals in right and a single blunt one in left valve. Ligament secondarily prosodetic. Umbos di- rected postero-medially. Edge of byssal orifice with a series of 6 to 12 yellowish orange, distinct to nearly obsolete plicae becoming slightly larger pos- teriorly. Ventral slope flat in young specimens, be- coming concave in old individuals. Interior of valves porcelaneous, usually white but often flushed with yellowish orange along posterior margin and in re- gion of hinge and ventral margin; the external strawberry blotches commonly showing through. Pallial line entire, only moderately wide. Muscle scars central, medium-sized; scars kidney-shaped in left valve and extending over the internal width of the 2 fold interstices; scars roundly oval in right valve and extending over only one interstice. Poste- rior pedal retractor muscle scar less than half the size of posterior adductor scar and located anterior to it. Both scars contiguous to pallial line. Area within pallial line, excluding muscle scars, dull; pallial line, muscle scars and areas to the periphery of shell shiny. Prodissoconch unknown. Byssus present in young individuals but not older ones. Measurements (mm.) — Tridacnidae are indis- criminately inequivalved, and the figure given un- der “length” and “height” is always the maximum measurement. “Width” indicates the greatest dis- tance through opposed valves, length height width 397 273 259 large; Pandora Reef, Queens- land (Hedley, 1923) 329 189 187 large: Galero Bay, Mindoro, Philippines 315 243 235 large: Rojoa Id., Eniwetok, Marshall Islands 278 172 177 average; Oyster Bay, Philip- pines 251 243 231 average; Aomoen Id., Bikini, Marshall Islands 223 145 147 average; Koror Id., Palaus 140 93 101 small; Panganan Id., Philip- pines Synonymy — 1758 Chama hippopus Linne, Systema Naturae, ed. 10, p. 691 (in M. Asiatico); refers to Buonnani, Recr. 2, figs. 81, 82; Rumphius, pi. 42, fig. C, and others. 1788 Chama asinus Barbut, Genera Vermium, p. 47, pi. 6, fig. 5 ( no locality given ) . [ 1797 Tridachna ursina Humphrey, Museum Calonnianum, p. 50 (China); refers to Chama hippopus Linne; rejected work.] 1798 Tridachnes ungula Roding, Museum Boltenianum, p. 172, no. 197 (no locality given); refers to Gmelin, Chama hippopus, species 3, and to Chem- nitz, vol. 7, pi. 50, figs. 498, 499. 1801 Hippopus maculatus Lamarck, Systeme Animaux sans Vertebres, p. 117 (no locality given); refers to Lister, pi. 349, fig. 187, pi. 350, fig. 188 and others; 1835 Tridacna maculata Quoy and Gaim- ard, Astrolabe, Zoologie, vol. 3, p. 491; Atlas, pi. 80, figs. 4-6. 1801 Hippopus brassica Bose, Histoire Naturelle des Co- quilles, Deterville ed., vol. 3, p. 91 (pi. 21, fig. 6) (dans la mer des Indes); refers to Rumphius, pi. 42, fig. C, and others. 1808 Camus hyppopus Boumon, Traite complet de la Chaux Carbonatee et de l’Arragonite, London, vol. 1, p. 326 [nomen nudum], 1853 Hippopus equinus Morch, Catalogus Conchyliorum Comes de Yoldi, part 2, p. 56 ( Ind. or. ) ; refers to Chama hippopus Linne, T. ungula Bolten [Rod- ing], and Hippopus maculatus Lamarck. Types and Nomenclature — The probable holo- type of Chama hippopus Linne is in the Linnaean Collection, London (Dodge, 1952). There seems no need to restrict Linne’s type locality, “In M. Asi- atico.” This well-known species is widely distrib- [62 - 033] 364 Hippopus Joseph Rosewater Tridacnidae uted in the western Pacific region and such a re- striction would have little meaning as Linne’s type could have come from one of many different local- ities. Locations of the type specimens of species de- scribed by Roding, Bose and Morch are unknown. The type figures of H. maculatus Lamarck, here selected, are Chemnitz, vol. 7, pi. 50, figs. 498, 499. The only possible source of confusion in the syn- onymy of this species is the name Hippopus bras- sica Bose. The name brassica was apparently taken from Linne’s translation of d’Argenville’s vernacular for the species, “la feuille de choux” viz. “Folium brassicae,” which means “the cabbage leaf.” Bose refers to his own pi. 21, fig. 6 (a copy of d’Argen- ville’s ( 1757 ) pi. 23, fig. E ) . But this is probably a poor specimen of Tridacna squamosa. His reference to “Dargenville, 1757 pi. 23, fig. 11.” is a misprint for fig. H. This figure and the remaining plate ref- erences are all definitely assignable to H. hippopus. Therefore brassica is an absolute synonym of hip- popus. The name brassica is a feminine substantive noun and must not be made to agree in gender with the masculine genus Hippopus. Records — Literature records from Madagascar and other western Indian Ocean Islands ( von Martens, 1880; Dautzen- berg, 1923, 1929; all based on Szanzin, 1843) are consid- ered erroneous, and a single dead valve recorded from Tern Island, French Frigate Shoal is probably adventitious. MA- LAYA: Singapore (USNM; MCZ; Reyne, 1947; Purchon, 1955). RYUKYU ISLANDS: Kana (MCZ); Shioya, Shana Wan; Shuri, all Okinawa (both USNM); northeast of Iheya Shima (ANSP). Philippines: Bantan Id., Batanes Group; Port San Pio Quinto, Camiguin Id., Babuyanes Group (both USNM); Subic Bay, Zambales Province (MCZ); Badang, near Gubat, Sorsogon Province, both Luzon (ANSP; MCZ); Tilig, Lubang (MCZ; USNM); Galero Bay, Mindoro (USNM); Masbate, Masbate Id. (ANSP); Nasog Point, Panay; Guimaras (both USNM); Cebu City, Cebu (A. B. Franco, ANSP; USNM); Panganan Id., between Cebu and Bohoi; Cajoagan Id. off northern Samar; Hilongas, Leyte (all USNM); Zamboanga (ANSP; USNM); Malabang, Cota- bato Province, both Mindanao (USNM); Tara Id.; Linapa- can Id.; Culion Id.; Port Colton, Busuanga, all Calamianes Group (all USNM); Puerto Princessa, Palawan (duPont- Academy Expedition, ANSP); Cagayan Id., Cagayanes Group (USNM); Cuyo Id., Cuyo Islands (ANSP); Jolo (USNM); Bongao Channel, southwest end Sanga Sanga Id.; Sibutu Id., both Sulu Archipelago (both ANSP). INDO- NESIA: Pulau Melila, south of Udjung Batu, Banjak Is- lands; Pulau Bai, Batu Group, both southwest of Sumatra (both International Indian Ocean Expedition, 1963, USNM); Tjiperwagaran, Bantam, Java (USNM); Pulau Kasiruta; Amboina; Pulau Batjan; Pulau Karakelong, Talaud Islands; Pulau Manipa, all Moluccas (all MCZ). NORTH BORNEO: Kukuban; Pulau Mandidarah; Pulau Silingaan (all Mary Saul, USNM). WESTERN AUSTRALIA: Augustus Island, (MCZ). QUEENSLAND: Torres Strait (ANSP); Portland Roads; Green Island, near Cairns (both MCZ); Brook Id.; Pandora Reef (both H. A. Pilsbry, 1933, ANSP); Hayman Id., off Bowen (USNM, MCZ); Lupton Id., Cumberland Group (MCZ). NEW GUINEA: Biak, Schouten Islands (USNM, ANSP); Seroei Bay, Japen (ANSP) both West Irian; 10-15 miles east of Aitape, North-East New Guinea (USNM); Oro Bay, Papua (ANSP). BISMARCK ARCHI- PELAGO: Koruniat Id., north coast Manus Id., Admiralty Islands; Gumlun Id., southwest of New Britain (both ANSP). SOLOMON ISLANDS: Kieta, Bougainville; Choi- seul Bay, Choiseul; Ataa District, Malaita (all ANSP); Pavuvu, Russell Islands (USNM). NEW HEBRIDES: Es- piritu Santo Island (MCZ); Efate (USNM). FIJI ISLANDS: Suva, Vitilevu (USNM). NEW CALEDONIA: lie Mouac, Poume; Plage de Poe, Bourail; Anse Vata Bay, Noumea; near Touho; Yate (all G. and M. One, ANSP). PALAU ISLANDS: Ngajangel, Kayangel Islands (USNM); 1 mile south of West Passage, Babelthuap (ANSP); Malakal Har- bor, Koror (USNM; ANSP; MCZ); Angaur (USNM). CAROLINE ISLANDS: Tomil Harbor, Yap; Fassarai Id., Ulithi; Eauripik; Ifalik; Elato; Lamotrek; Satawal; Ponape; Kapingamarangi (BPBM; all USNM). MARSHALL IS- LANDS (many atolls; see map); Eniwetok; Ujelang; Kwaja- lein; Jaluit; Bikini; Rongelap; Utirik; Ailuk; Majuro ( all USNM). GILBERT ISLANDS: Abemama (USNM); Onotoa Atoll (MCZ; USNM). TONGA: (BPBM) Fossil Records — FIJI: Late Pleistocene, Ravuka, Viti Levu (Ladd, 1934). MARIANAS ISLANDS: Miocene, Tog- pochau limestone, Saipan; Pliocene-Pleistocene, Marianas limestone, Guam (both U. S. Geological Survey). PALAU ISLANDS: Neogene (U. S. Geological Survey). Plate 272. Geographical distribution of Hippopus hippopus (Linne). [62 - 034] April 30, 1965 INDO-PACIFIC MOLLUSCA, vol. 1, no. 6 Hippopus 365 Hippopus? gunteri Mansfield, 1937 (PI. 273) Range — Fossil (Tampa Limestone: Lower Mio- cene) of Florida. Remarks — The type and several additional speci- mens from the same horizon consist of fairly well preserved molds of external sculpture in the region of the umbo and ventral slope. It is extremely diffi- cult to distinguish the sculptural pattern of these fossils from that of the Recent species, Hippopus hippopus (Linne). However, as noted by Mans- field, final generic disposition must await the dis- covery of specimens showing the dentition. It would be very helpful also to find specimens show- ing more completely the shapes of the valves. If this is actually a Hippopus, speculation is invited re- garding the former distribution of the genus which is now confined to the central and western Pacific. If confirmed it can be stated that during the middle Tertiary this member of the genus did inhabit the region which is now northwestern Florida, perhaps then a portion of the shore of the Tethys Sea. There is a vague similarity between the sculpture of this species and Avicularium trechmanni Cox, 1941 from the Upper Eocene of Jamaica. Synonymy — 1937 Hippopus ? gunteri W. C. Mansfield, Geological Bul- letin, State of Florida Department of Conservation, no. 15, p. 258, pi. 20, figs. 2, 4; Type: USNM 495963 (Station 12,300, along the channel of a disappearing stream, 1% miles northeast of Lloyd, Jefferson County, Fla). Plate 273. Hippopus? gunteri Mansfield, from 1 % miles northwest of Lloyd, Jefferson County, Florida, Lower Mio- cene (40 mm.) (from Mansfield, 1937, pi. 20, figs. 2, 4; Smithsonian photos). [62 - 039] 366 Hippopus Joseph Rosewater Tridacnidae [These occasional blank areas occur between genera and subgenera to permit the insertion of new material and future sections in their proper systematic sequence.] I [62 - 040] April 30, 1965 INDO-PACIFIC MOLLUSCA, vol. 1, no. 6 Tridacna 367 Genus Tridacna Bruguiere, 1797 The genus Tridacna is confined to the Indo- Pacific area. It contains the species of true Giant Clams which are identified by their large size and distinctive ecology, always being located on or near reef-forming corals. Three subgenera may be dis- tinguished: Tridacna s.s., Persikima Iredale and Chametrachea Morch. Subgenus Tridacna Bruguiere, 1797 Type: Tridacna gigas (Linne, 1758) Mature Tridacna gigas can be distinguished from species of Chametrachea and Persikima by its larger size, more equilateral valves, deeply folded sculp- ture, and free living habits ( not byssally attached ) . Tridacna lacks the strawberry spots, elongate-tri- angular shape and neatly interdenticulated ventral margin of Hippopus. An anatomical distinction in Tridacna sensu stricto is the possession of a com- plete outer demibranch, a character lacking in both Chametrachea and Persikima. Hippopus lacks the hyaline organs which are present in species of all subgenera of Tridacna. Synonymy — [ 1775 Tricdane d’Herbigny, Dictionnaire d’Histoire Natu- relle, vol. 3, p. 408; Tricdana, p. 411 (non- binomial).] [ 1776 Tridacna Da Costa, Elements of Conchology, p. 294, p. 7, fig. 4 (non-binomial).] [ 1792 Tridacne Bruguiere, Encyclopedic Methodique, His- toire Naturelle des Vers, vol. 1, p. 386 (vernacular name).] [1797 Tridachna G. Humphrey, Museum Calonnianum, p. 50; (rejected work).] 1797 Tridacna Bruguiere, Tableau Encyclopedique et Methodique, Vers Testacees, heading of pi. 235. Type by subsequent monotypy (Lamarck, 1799): Chama gigas Linne [= Tridacna gigas (Linne, 1758)]. 1798 Tridachnes Roding, Museum Boltenianum, part 2, p. 171. Type by subsequent designation (Iredale, 1937): Chama gigas Linne. 1807 Tridacne Link, Beschr. Nat. Samml., Univ. Rostock, part 3, p. 153. [ 1807 Tridacnigenus Renier, Tavole per servire alle Classi- ficazione e Connescenza degli Animali, Padua, tav. 7 (rejected name).] 1823 Tridacnodites Kruger, Geschichte der Urwelt, part 2, p. 464; type by monotypy: Tridacnodites gigas = Tridacna gigas (Linne)]. 1827 Gataron ‘Adanson’ Berthold (in Latreille) Natiirliche Familien des Thierreichs, p. 207 (nomen nudum); in Chamidae, fide Vokes [1955, p. 130], 1848 Tridacnoides ‘Kruger’ Bronn, Index Palaeontologicus, Stuttgart, p. 1279 “= confusio rerum” [not listed in Kruger, 1823, but credited to him by Scudder (1882), Sherbom (1931), Neave (1940), and Vokes [1951]] nomen nudum. 1875 Tridachnus Paetel, Die bisher veroffentlichten Fam- ilien und Gattungsnamen der Mollusken, p. 211 [error for Tridachnes Roding]. 1898 Tridachne Dali, Trans. Wagner Free Instit. Sci. Philadelphia, vol. 3, part 4, p. 572 (emendation for Tridacne Link, 1807). 1937 Dinodacna Iredale, Australian Zoologist, vol. 8, part 4, p. 238. Type by original designation: D. cooki- ana Iredale, 1937 [= Tridacna gigas (Linne)]. 1940 Tridachna ‘Lamarck’ Neave, Nomenclator Zoologicus, vol. 4, p. 552 [error for Tridacna Bruguiere, La- marck, 1799; perpetuated by Vokes [1951, p. 77]]. Nomenclature — The generic name Tridacna was proposed validly for the first time in 1797 on the plate heading of Tableau Encyclopedique et Meth- odique, pi. 235 (I. C. Z. N., Article 16 (a) (vii)). In spite of arguments to the contrary by Dodge ( 1947) it is quite probable that Bruguiere personally approved the plates bearing the name Tridacna and should, therefore, be listed as the author. Tridacna wolfarti Chenu, 1845 (PI. 274) Range — Upper Tertiary, Hesse, Germany. Remarks — This species is very closely allied with T. gigas. The occurrence of fossils such as T. wolf- Plate 274. Tridacna (Tridacna) wolfarti Chenu, from Hesse-Inferieure, Tertiary (reduced from Chenu, 1845, pi. 8, figs. 5, 6; Smithsonian photos). [62 - 043] 368 Tridacna Tridacnidae Joseph Rosewater Plate 275. Tridacna (Tridacna) gigas ( Linne ). Dorsal view from Linapacan Island, north of Palawan, Philippines to show elongate-triangular valve interdigitations. Specimen (length: 370 mm.; 14.6 in.) (Smithsonian photo). [62 - 044] April 30, 1965 INDO-PACIFIC MOLLUSCA, vol. 1, no. 6 Tridacna 369 arti and T. media in Tertiary deposits of Europe seemingly indicates the presence in that area dur- ing the Tertiary of a flourishing tropical fauna much like that in the Indo-Pacific. Synonymy — 1845 Tridacna wolfarti Chenu, Illustrations Conchyliologi- ques, vol. 2, Tridacna, p. 2, pi. 8, figs. 5-6 ( . . . montagnes de la Hesse-Inf^rieure ) . Tridacna gigas (Linne, 1758) (Pis. 263, 267, 275-278) Range — Philippines to Micronesia. Remarks — Tridacna gigas Linne is the largest known species of bivalve. In the phylum Mollusca it is surpassed in size and weight only by the Re- cent giant squids and some fossil nautiloids and ammonites. The largest pair of valves on record measured 4 feet, 6 inches in length ( about 137 cm. ) and weighed 507 pounds. They were brought from Tappanooly [Teluk Tapanuli?], Sumatra, and were seen at Arno’s Vale, Ireland (Dillwyn, 1817, p. 214). The heaviest reliably recorded specimen of T. gigas, one with valves weighing 579/2 pounds, is in the collection of the American Museum of Natural His- tory, New York (Miner, 1938). Only somewhat lighter are the valves weighing 550 pounds which serve as holy water fonts in the Church of St. Sul- pice in Paris and which were presented by the Re- public of Venice to Francis I in the 16th Century (Lamarck, 1819). The latter are 36 inches long (about 91.5 cm.; in litt., R. T. Abbott, 1962). In his original description of T. gigas, Linne listed the weight of the Museum Ulricae specimen as 532 pounds. There is some disagreement over the exact value of the Swedish pound in Linne’s day, and this weight was quoted as 498 pounds by Smith ( 1898 ) . Roscoe ( 1962 ) listed additional records for large T. gigas. It is possible that even larger and heavier individuals may be found in the future, but reports that the giant clam can reach half a ton or more are probably exaggerated (Ruchsbaum and Milne, 1960; Hawaiian Shell News, 1962, vol. 10, no. 10, p. 2). Tridacna gigas can be distinguished from T. crocea and T. maxima by the larger size of the former, by the more central position of the umbos, by its lack of a large byssal orifice, by the lack of crowded, undulate sculpture, and by the absence of “boring” tendencies. The mantle coloration in living T. gigas is yellowish-brown to olive-green with blue-green spots, while in crocea and maxima the mantle usually is more brilliantly colored. Certain specimens of T. gigas and T. derasa ap- pear very similar in shell characters. The two species can be distinguished by differences in ctenidial anat- omy, derasa lacking a complete outer demibranch. The shell of gigas is usually less obese; the free margins of the valves have sharply triangular inter- digitating processes, whereas these are undulate in derasa which also has much lower radial sculpture. The giant clam is larger than and lacks the scales of T. squamosa. McLean ( 1947 ) traced the de- velopment of T. gigas through a graded series rang- ing from very young juveniles to moderately large adults. All specimens were found to lack scales ex- cept near the byssal orifice where small ones are present. Tridacna squamosa has widely spaced, erect scales. Although certain specimens of gigas and squamosa agree in many characters, the two are unquestionably distinct (see subgenus Tri- dacna). Holthuis ( 1952 ) reported the following shrimps of the family Palaemonidae to be commensal with T. gigas: Anchistus mirabilis (Pesta), Conchodytes Plate 276. Tridacna (Tridacna) gigas (Linne). Dorsal view of living animal to show mantle pattern and appearance of siphonal openings. Note that byssal opening is directly beneath incurrent siphonal opening ( total length: 559 mm.) (author’s photo at Eniwetok, Marshall Islands). 162 - 045] 370 Tridacna Joseph Rosewater T ridacnidae tridacnae Peters, and Paranchistus biunguiculatus ( Borradaile ) ; the latter species was also present in T. gigas collected by the writer at Eniwetok Atoll, Marshall Islands (identified by Dr. F. A. Chace, USNM). The giant clam suffers a bad reputation from stories describing the fate of unwary persons caught between its valves. C. W. Johnson (1930) presented one view of this supposed phenomenon, implying that the giant clam is not as dangerous as the stories suggested. There are apparently very few documentations of the giant clam killing men, al- though Cobb ( 1939 ) insisted that the specimen of T. gigas which yielded the “Pearl of Allah” killed the diver who found it; and there is an elderly east coast Malaysian who attributes the loss of one of his legs to Tridacna gigas (personal communication, Johnnie Johnson, Singapore, 1963). If a diver were so unlucky as to catch an appendage between the valves of a closing T. gigas, doubtless he could Plate 277. Lectotype of Tridacna (Tridacna) gigas (Linne). Fig. 2. Internal view of left valve (length: 90 cm.). (Photos Fig. 1. External view of right valve (length: 93.5 cm.), courtesy of Zoological Institute, Uppsala). [62 - 046] April 30, 1965 INDO-PACIFIC MOLLUSCA, vol. 1, no. 6 T ridacna 37 1 drown before being released. The growing edges of valves of living T ridacna are extremely sharp and capable of inflicting deep and painful cuts if handled carelessly. Persons also have been injured by having fingers squeezed in the byssal orifice when lifting the clams (personal communication, John Roberts, 1963). Habitat — Lives on coral reefs, usually on sand, in depths of from only a few feet to several fath- oms; the depth at which this species lives may be limited by the penetration of light strong enough to permit growth of the algae contained in its tis- sues. Color Notes (see plate 263) — Main ground color of mantle ( living specimen from Eniwetok ) of medium-sized and adult T. gigas is dark yellow (Kelly and Judd, 1955, ISCC-NBS color name; C553 number 88); the outer edge, or border of mantle is usually darker, closest to dark olive brown (C553 number 96) but blotches of the former, dark yel- low, may be interspersed. Along darker outer bor- der are found in considerable numbers small rings of color, each surrounding from one to several hyaline organs; a continuous line of smaller, colored rings is found along the mantle edge at its periph- ery. Rings are numerous also around siphonal open- ings where numbers of hyaline organs may be sur- rounded by these irregular to elongate enclosures. When rings are viewed by refracted light (i.e., specimen is located between light source and ob- server) their color appears closest to strong green- ish blue ( C553 number 169 ) or strong blue ( num- ber 178). But when viewed by reflected fight (i.e., observer is between fight source and specimen ) the color of rings is brilliant bluish green (number 159). This difference in color, based on viewing Plate 278. Tridacna (Tridacna) gigas (Linne). Young speci- Dorsal view. Fig. 2. Ventral view, showing small byssal men from Green Island, near Cairns, Queensland, Australia, orifice ( 123 mm. in length ) ( Smithsonian photo ) . showing semitubular spines on summits of folds. Fig. 1. [62 - 047] 372 Tridacna Joseph Rosewater Tridacnidae position, may be responsible for some of the varied statements regarding mantle color in T. gigas. Even in the same specimen, the viewer may see rings of both blue and green at the same time when the mantle is somewhat contracted and it is struck by light from different angles. This phenomenon was not noted in T. squamosa, T. maxima or Hippopus hippopus. Toward the center of exposed mantle surface the dark yellow diminishes, becoming grayish white in spots. Scattered rather regularly over the central portion of the mantle are hyaline organs without color rings. Darker color of outer mantle edge is re- peated on edges of the siphons where, as previously mentioned, the blue or green colored enclosures sur- round clumps of hyaline organs. Small specimens of T. gigas 150-200 mm. long, were observed to lack the dark olive brown colora- tion, being more uniformly dark yellow or lighter, but with a few color rings present. Activities and Habits — Valves of Tridacna gigas remain widely gaping most of the time, except when the clam is disturbed. In the expanded con- dition the dark edges of the mantle are reflected over the edges of the valves. The incurrent siphon remains open and gaping, as does the pedal open- ing (see plate 276). It is theorized that the pedal opening can serve as a direct outlet for any large par- ticles of debris which may fall or be drawn into the incurrent opening. The excurrent siphon also is held open and gaping “hoselike” and the aperture is usu- ally pointed anteriorly, probably in order to direct the excurrent stream away from the incurrent open- ing. In T. gigas the ctenidia are comparatively large. Through the incurrent siphonal opening the inner demibranch is easily seen hanging ventrally within the mantle cavity. The outer demibranch is de- flected obliquely and laterally, so that the character of the food groove is not easily visible externally. When stimulated by a change in light intensity, such as a shadow over the excurrent siphonal area, the siphon rolls inward and closes. There follows a sudden contraction of the valves causing a great surge of water to be squirted out of the incurrent siphon. The reaction can be initiated also by touch- ing the mantle. It is probable that the reaction is a “surprise” and defensive one and may be very use- ful in frightening away fish which come to browse on the clam’s mantle ( see Stasek, 1962, pp. 30, 31 ) . The initial closing is usually retarded by the pres- ence of a considerable quantity of water within the mantle cavity. The majority of this is expelled by the initial closing, and subsequent closings bring the valve edges into closer proximity. Water Current through Mantle Cavity — An at- tempt was made to measure the rate of flow of wa- ter through the mantle cavity of T. gigas. A quan- tity of neutral red mixed with seawater to form a dense red dye was introduced to the area of the in- current siphon. The intake appeared to be very slow. The dye dropped into the opening and ap- peared to be guided laterally, down the sides of the mantle cavity to the base of the ctenida. None was pumped out of the excurrent opening, but there was a rejection reaction, i.e., a counter current was forced out of the incurrent opening several times expelling the dye. The fact that the current through the mantle cavity is slow may indicate that little “normal bivalve feeding” is taking place and that Plate 279. Geographical distribution of Tridacna (Tridacna) cles: from the literature. gigas (Linne); solid dots: specimens examined; open cir- [62 - 048] April 30, 1965 INDO-PACIFIC MOLLUSCA, vol. 1, no. 6 Tridacna 373 the major part of the animals’ food is obtained from the zooxanthellae. A large amount of water is main- tained within the mantle cavity with some continu- ous exchange, and this apparently suffices for res- piratory purposes. A specimen of T. gigas 305 mm. (12 inches) in length contained approximately 1.2 liters of water. This amount of water could be very useful to the clam in performing the defensive re- action mentioned above. Description — Shell reaching about 137 cm. ( 4 feet, 6 inches) in length; suboval to fan-shaped in outline; angle formed at umbos by hinge line and ventral margin usually exceeding 150°; moderately to strongly inflated and with small to nearly closed byssal orifice. Valves moderately thin in small speci- mens to extremely heavy and thick in large indi- viduals; color grayish white; encrusted with ver- metids, annelid tubes, coral, coralline algae and other organisms. Surface of valves dull or faintly shiny in cleaned specimens. Primary radial sculp- ture consists of from 4 to 6 strongly convex, rib-like folds which are crowded and indistinct on the ven- tral slope. Four of the folds are usually very large. Secondary radial sculpture consists of rather evenly- spaced riblets on both folds and their broad inter- stices (often grouped into bands of 3 or 4 riblets in interstices); radial riblets interrupted by fine concentric lines of growth resulting in minute retic- ulations. Where secondary and radial concentric sculptures cross, nodules may be formed. Sculpture becoming obsolete in later growth in large individ- uals. Scales limited to summits of folds near umbos; scales small, semitubular but usually worn away. Dorsal shell margins broadly fan-shaped in outline, undulate, with series of 4-5 usually rather sharply pointed, medially projecting interdigitating proc- esses representing extremities of rib interstices, often crenulate at then- edges. Hinge line somewhat longer than half the length of shell. Hinge teeth becoming obsolete in large individuals. A single, oblong cardinal tooth present in each valve; 2 elon- gate posterior lateral teeth in right valve and 1 in the left. Ligament secondarily prosodetic. Umbos directed posteromedially. Ventral margin nearly straight, but may be slightly concave in region of byssal orifice. Edge of byssal orifice with series of 4-7 nearly obsolete plicae, usually evenly spaced and becoming less distinct posteriorly. Ventral mar- gin posterior to byssal orifice weakly crenulate. Ventral slope flattened to concave in larger speci- mens, often largely covered by overgrowth of por- celaneous material from edge of byssal orifice. In- terior of valves porcelaneous and white. Interior of convexities of primary radial sculpture closed as sharp folds within pallial line, smoothly concave without. Pallial line entire, wide, roughened. Pos- terior adductor muscle scar large, subcircular, adja- cent to pallial line dorsally and located posterior to center. Posterior pedal retractor muscle scar less than half the size of posterior adductor, and con- tiguous to upper portion of latter; located just an- terior to center. Area within pallial line, excluding muscle scars, dull; area of line, muscle scars, and to periphery of shell, shiny. Prodissoconch unknown. Measurements (mm.) — length height width 1370 - - large; Tappanooly [Teluk Ta- panuli?], Sumatra 1016 — - average; Singapore, Malaysia 1003 — — average; Bikini, Marshall Is- lands 750 440 average; Pulau Siburu, north of Sipora, off southwest Sumatra 445 270 270 small; Pulau Bai, Batu Group, off southwest Sumatra 370 230 228 small; Palawan, Philippines 123 80 67 small; Green Island, near Caims, Queensland, Aus- tralia Synonymy — 1758 Chama gigas Linne, Systema Naturae, ed. 10, p. 691 (Habitat in M. Asiatico); refers to specimen in “Mu- seum Ulricae”; not T. gigas ‘Linne’ Reeve, 1862, Conchologia Iconica, vol. 14, Tridacna, pi. 1, fig. la, pi. 2, figs, lb, c which is T. maxima Roding. 1801 Tridacna gigas Lamarck, Systeme des Animaux sans Vertebres, p. 117 (no locality given) (part). 1807 Tridacne imbricata Link, Beschreibung Naturalien Sammlung Univ. Rostock. (3) p. 153 (no locality given); refers to Chama gigas ‘Linne,’ Gmelin, p. 3299, and to Chemnitz, vol. 7, pi. 49, figs. 492-493 [= T. squamosa ] 494 [= T. maxima ] (part). 1808 Camus gigantus Boumon, Traite Complet de la Chaux Carbonatee et de l’Arragonite, London, vol. 1, p. 325 [nomen nudum]. 1811 Chama gigantea Perry, Conchology, Introduction, p. 2 (no locality given). 1819 Tridacna gigas Lamarck, Histoire Naturelle des Ani- maux sans Vertebres, vol. 6, part 1, p. 105 (l’Ocean indien) (part). 1819 Tridacna mutica Lamarck, ibid., p. 106 (l’Ocean des grandes Indes) (part); Chenu, vol. 2, pi. 4, figs. 1, la [= T. gigas], pi. 6, fig. 1 [= T. maxima ]. 1903 Tridacna lamarcki Hidalgo, Memorias Real Academia de Ciencias, Madrid, vol. 21, p. 385 (Mindoro, Cebu, Mindanao and Paragua Ids.) (part); refers to Quoy and Gaimard, pi. 79, figs. 4, 5 and others [= T. gigas] ; new name for T. gigas Lamarck, not Linne. 1921 Tridacna gigas ‘Linne,’ Hedley, Records of the Aus- tralian Museum, vol. 13, no. 4, p. 168, pi. 227, fig. 1 [but not fig. 2 = T. maxima]. 1937 Dinodacna cookiana Iredale, Australian Zoologist, vol. 8, part 4, p. 238 (Gilbert Islands); refers to Sowerby, Thesaurus Conchyliorum, vol. 5, pi. 488, fig. 11 (type figure; Samoa [a doubtful record] ) . [62 - 049] 374 Tridacna Joseph Rosewater Tridacnidae Types — Linne’s reference to a large specimen of T. gigas weighing 532 pounds, preserved in the “Museum Ulricae,” is the only clue to the location of the type of this species. None of the plate refer- ences listed with the original description shows a clear-cut example of T. gigas. All of them, except one, are referrable to either T. squamosa or T. maxima. Gualtieri’s pi. 92, fig. G. may represent a young individual of either T. gigas or T. squamosa. There is present in the collection of the “Museum Ulricae,” at the University of Upsala, a large speci- men of T. gigas measuring 93.5 cm. in length. The 532 Roman pounds mentioned by Linne ( 1 Roman pound = 12 ounces ) has an English equivalent of 399 pounds, a possible weight for a giant clam measuring nearly 94 cm. Hedley (1921, p. 169) has already suggested that the type is the large speci- men said by Linne to be present in the “Museum Ulricae.” Although great care must be exercised in the recognition of type material from this collec- tion, there should be little difficulty in recognizing this specimen as the name-bearer for this species (Dodge, 1952). It is here designated the lectotype of Chama gigas Linne, 1758 (plate 277). The type locality given by Linne, “M. Asiatico,” is here re- stricted to Amboina, Moluccas, a locality from which material was available at that time. The locations of the types of T. mutica Lamarck and T. imbricata Link are unknown, and both of these are composite species. Nomenclature — The name Tridacna gigas (Linne) is universally connected with the giant clam of the Indo-Pacific. However, the name has a confused early history. As emphasized by Dodge (1952), Chama gigas Linne, 1758, is a composite species. The references given by Linne refer to the true giant clam and the species later named T. squamosa by Lamarck and T. maxima by Roding. If it were not for the reference in the original de- scription to the goliath specimen in the “Museum Ulricae,” weighing 532 pounds, the applicability of the name gigas would be in considerable doubt. However, because of that fortunate reference, the name can be restricted to this species. Such useful works as Reeve’s Conchologia Iconica show the extent of confusion regarding the true identity of T. gigas. The fine figures in this work are often cited as examples of species. Dodge (loc. cit., p. 131) referred to Reeve’s Tridacna, pi. 1, fig. la, pi. 2, figs, lb, lc, as representing T. gigas. As al- ready indicated by McLean ( 1947 ) , these figures do not represent gigas. The short hinge line, sculp- ture and delicate coloration show them to be T. maxima. Tridacna gigas was figured only once by Reeve (Conch. Icon., vol. 14, pi. 6, fig. 6, a young specimen; not T. serrifera Lamarck = T. derasa ). This species was seldom well-figured in early works. Most species were drawn natural size, which would have been impossible in the case of larger speci- mens of gigas. Smaller individuals were selected, therefore, and in many cases these were other spe- cies. Lamarck’s T. mutica apparently belongs as a syn- onym under T. gigas at least in part. The descrip- tion is not particularly helpful, but in his remarks Lamarck cites the lack of sculpture, small “lunule” aperture ( = byssal opening ) and relatively large size, 37 cm. of this species. Records — RYUKYU ISLANDS: Itoma Jima (Pilsbry, 1895, p. 181). PHILIPPINES: Looc Bay, Lubang (P. de Mesa, MCZ); Busin Harbor, Rurias Id.; Santa Cruz Id., off Zamboanga, Mindanao; Linapacan Id., Calamianes Group (all USNM). INDONESIA: Pulau Melila, south of Udjung Batu, Banjak Islands; Pulau Bai, Batu Group; Pulau Siburu, north of Sipora, all southwest of Sumatra (all International Indian Ocean Expedition, 1963, USNM). AUSTRALIA: Brook Island (H. A. Pilsbry, 1933, ANSP); Green Island, near Cairns, both Queensland (J. K. Howard, MCZ). NEW GUINEA: Poelau Rouw; Poelau Maransabadi, both Aoeri Group, Geelvink Bay; Poelau Noekori, east Padeaido Is- lands, all West Irian (all ANSP). BISMARCK ARCHI- PELAGO: Kumbun Island, southwest of New Britain (ANSP). SOLOMON ISLANDS: Kieta, Bougainville Id.; Choiseul Bay; Bambatana, both Choiseul (all ANSP). NEW HEBRIDES: Efate (USNM). PALAU ISLANDS: Koror (BPBM). MARSHALL ISLANDS: Bikini; Eniwetok (both USNM). GILBERT ISLANDS: Onotoa (Banner, 1952 [as Hippopus ] ) . Fossil records — INDONESIA: Lower and Upper Miocene (Tjilanang beds), Java (K. Martin, 1880, p. 119, pi. 19, fig. 3). MARIANAS ISLANDS: Pliocene-Pleistocene, Marianas limestone, Guam (U. S. Geological Survey). [62 - 050] April 30, 1965 INDO-PACIFIC MOLLUSCA, vol. 1, no. 6 Tridacna 375 Tridacna derasa (Roding, 1798) Subgenus Persikima Iredale, 1937 Type: Tridacna derasa (Roding, 1798) The genus Persikima was established by Iredale for the peculiar species he named P. whitleyi ( = derasa ). Sufficient characters appear to be present to allow a subgeneric distinction. These are an ex- treme posterior displacement of the umbos and an anterior extension of the ctenidia. In other sub- genera of Tridacna the umbos are either about cen- tral or displaced anteriorly; the ctenidia usually terminate at about the level of the mouth. Persi- kima appears to occupy an intermediate position between Tridacna s.s. and Chametrachea. Synonymy — 1937 Persikima Iredale, Australian Zoologist, vol. 8, part 4, p. 237. Type by original designation: P. whitleyi Iredale, 1937 [= T. derasa (Roding, 1798)]. (Pis. 280-282) Range — Philippines, East Indies, Cocos-Keeling Islands and western Micronesia. Remarks — This species is distinguished easily by its low primary and secondary radial sculpture, its often massive umbonal area and long hinge line causing the umbos to be displaced far posteriorly. Anatomically the most striking feature is the ante- rior displacement of the ctenidia which terminate far forward of the mouth. This necessitates ex- tremely elongate distal oral grooves (terminology of Kellogg, 1915; Stasek, 1962; ? = lateral oral grooves of Purchon, 1955) which lead posteriorly and ventrally to the labial palps and mouth. Because of the absence of spiny sculpture and the considerable variation in shell shape, young specimens of T. derasa can be confused with cer- tain specimens of T. gigas. The confusion may be Plate 280. Tridacna (Persikima) derasa (Roding). Mature Internal view right valve; note long hinge line and pos- specimen from outer reef, off Noumea, New Caledonia teriorly displaced umbos. Fig. 3. Dorsal view to show inter- length: 408 mm.). Fig. 1. External view left valve. Fig. 2. digitating projections (Smithsonian photos). [62 - 057] 376 Persikima Joseph Rosewater Tridacnidae dispelled by a careful comparison of shell charac- ters. Where anatomical material is available there can be no doubt of its identity, for T. gigas has complete outer demibranchs while T. derasa has these structures incomplete. Also, T. derasa never reaches the size of large specimens of T. gigas. The peculiar anterior displacement of the ctenidia in derasa is a feature distinguishing this species from all other Tridacnidae. Although there are similarities in the massive rolled-in umbos of some specimens of T. derasa and Hippopus hippopus, and also in the occurrence of orange coloration in the hinge region, there is little Plate 281. Tridacna (Persikima) derasa (Roding). Holo- type of T. obesa Sowerby, from the Philippines (length: 158 mm.; BM(NH) 1899.4.22.81). Fig. 1. Exterior of right valve. Fig. 2. Interior of left valve. Fig. 3. Ventral view; note small byssal orifice and depressed folds, (photo cour- tesy of R. T. Abbott). danger of confusing these species as they are so dif- ferent in other respects. Tridacna derasa is quite rare in collections and I have examined material from only about ten lo- calities. Collecting data with specimens from Nou- mea, New Caledonia, indicate that T. derasa lives there on the outer edge of the barrier reef. Hazards of collecting in such localities may account for the shell’s variety in collections. In the Cocos-Keeling Islands, this species was collected in quantity from the atoll lagoon. It is interesting to note that there, in a more protected habitat, specimens developed strong concentric sculpture. So far as is known from present locality data, the range of T. derasa does not extend eastward into Micronesia as far as the eastern Caroline, Marshall or Gilbert Islands. Habitat — Lives on the outer edges of barrier reefs and in coral atoll lagoons; usually unattached. Description — Shell reaching 514 mm. ( about 20/1 inches) in length; semicircular in outline, angle formed at umbos by margins of hinge line and ven- tral border usually equaling or exceeding 150°; weakly to strongly inflated and having a very small byssal orifice. Valves usually very heavy and often greatly thickened at umbos, color grayish white, and to a limited degree encrusted with coralline algae, other organisms and debris. Primary radial sculpture consisting of from 7-12 broad, often nearly obsolete rib-like folds (usually 6-7 main folds) which become crowded and indistinct on ventral slope. Secondary radial sculpture consisting of fine riblets about equally distributed over folds and their equally broad interstices. Concentric sculpture consisting of closely-spaced, undulate lines of growth which upon intersecting secondary riblets may impart locally a faint nodular appear- ance visible under low magnification. Occasionally, as in the Cocos-Keeling population, concentric sculpture can be strongly expressed, producing con- tinuous, undulate, erect ridges. Dorsal valve mar- gins undulate, with series of 6-7 terminally rounded, medially projecting, interdigitating processes repre- senting extremities of rib interstices. Hinge line usually longer than half the length of shell; umbos located considerably posterior of center. Ligament secondarily prosodetic. Ventral margin only slightly concave, byssal orifice narrow and short; plicae 6-7, elongate, often nearly obsolete. Ventral slope nar- row to broadly concave, depending on obesity of specimen. Interior of valves white; hinge area often orange-tinged. Posterior adductor muscle scar large and subcircular in outline. Posterior pedal retractor [62 - 058] April 30, 1965 INDO-PACIFIC MOLLUSCA, vol. 1, no. 6 Tridacna 377 muscle scar one fourth to one third the size of and Synonymy — anterior to dorsal border of posterior adductor. Animal: A small byssus was present in an imma- ture specimen measuring 60 mm. in length. Two larger preserved specimens showed no sign of a byssus. In other respects, the animal is similar to T. maxima, crocea and squamosa in having incom- plete outer demibranchs and guard tentacles sur- rounding the incurrent siphon. The ctenidia extend anteriorly beyond the mouth about one third of their length. Very long distal oral grooves lead downward and posteriorly from ctenidia to bases of labial palps. Measurements (mm.) — length height width 513 292 315 large; “East Indies,” U. S. Ex- ploring Expedition, USNM 408 246 231 large; reef off Noumea, New Caledonia 236 150 129 average; Cocos-Keeling Islands 213 151 144 average; Cocos-Keeling Islands 133 86 56 small; Palau Islands 60 31 15 small; Palau Islands 1798 Tridachnes derasa Roding, Museum Boltenianum, part 2, p. 172, no. 195 (no locality given; Amboina, Moluccas, here designated); refers to Gmelin, Chama gigas, sp. 2/3, and to Chemnitz, vol. 7, pi. 49, fig. 497 [type figure; a young specimen]; Hed- ley, 1921, Records of the Australian Museum, vol. 13, no. 4, p. 167, pi. 28, fig. 4. 1807 Tridacne glabra Link, Beschreibung Naturalien Samm- lung Univ. Rostock (3) p. 153 (no locality given); refers to Chama gigas, var. /3 Linne, Gmelin, and to Chemnitz, vol. 7, pi. 49, fig. 497. 1819 Tridacna serrifera Lamarck, Histoire Naturelle des Animaux sans Vertebres, vol. 6, p. 107 (l’Ocean indien?); refers to Tableau Encyclopedique Meth- odique, pi. 235, fig. 3 [this and Chenu’s pi. 8, fig. 4, are copies of Chemnitz, vol. 7, pi. 49, fig. 497]; not Reeve, vol. 14, Tridacna, pi. 6, fig. 6 ( = T. gigas). 1899 Tridacna obesa Sowerby, Proceedings of the Malaco- logical Society of London, vol. 3, p. 210, figs. pp. 210-211 (Philippines); type in British Museum (N. H.). 1937 Persikima whitleyi Iredale, Australian Zoologist, vol. 8, part 4, p. 237, pi. 15 (Middleton Reef, Coral Sea); type in Australian Museum, Sydney. Plate 282. Tridacna (Persikima) derasa (Roding). Figs. 1, 4. Young specimen from 1% miles west of Eil Malk, Palau 2. Highly sculptured form from 3 miles southwest of Home Islands (length: 133 mm.) (Smithsonian photos). Island, Cocos-Keeling Islands (length: 238 mm.). Figs. 3, [62-059] 378 Persikima Joseph Rosewater Tridacnidae Nomenclature — Roding based T. derasa on the figure of Chemnitz, which was also referred to by Link for T. glabra, and a copy of which served as the basis for Lamarck’s T. serrifera (see Synonymy). The figure is of a young specimen of this species, as evidenced by the low sculpture and undulating shell margins. The specimen which served as the basis for Sowerby’s T. obesa was a highly inflated but not particularly large individual, 16 cm. in length, whereas Iredale’s type of P. ivhitleyi is 12 inches ( about 30.5 cm. ) long. Hedley ( 1921, p. 167 ) was the first to suggest a synonymy for derasa, which included serrifera La- marck and obesa Sowerby, although the figure of Reeve which he cited for serrifera is probably not this species, but young T. gigas. I cannot agree that T. mutica Lamarck is this species. The description of the latter recalls no known species and Chenu’s plates are of young T. gigas and T. maxima. Records — (solid dots on map : specimens examined; open circles: from the literature) — PHILIPPINES: Butauanan Island, Ambos Camarines, Luzon (USNM). EAST INDIES: Pulau Mandidarah, North Borneo, Malaysia (USNM; ANSP). COCOS-KEELING ISLANDS: ( Gibson-Hill, USNM); 3 miles southwest of Home Island (International Indian Ocean Expedition, V. Orr, 1963, ANSP, USNM). AUSTRALIA: Murray Island, Torres Strait (Hedley, 1921); Gillett Cay, Swain Reefs (Aust. Mus.; USNM) both Queensland; Mid- dleton Reef, off New South Wales (Iredale, 1937). NEW GUINEA: Poelau Rouw, Aoeri Group, West Irian (ANSP). NEW CALEDONIA: barrier reef off Noumea (ANSP; USNM). PALAU ISLANDS: Malakal Harbor, Koror (ANSP); Abappaomogan, west of Eil Malk (USNM); southeast Eil Malk (ANSP). Plate 283. Geographical distribution of Tridacna (Persikima) cles: from the literature. derasa (Roding); solid dots: specimens examined; open cir- [62 - 060] April 30, 1965 INDO-PACIFIC MOLLUSCA, vol. 1, no. 6 Tridacna 379 Subgenus Chametrachea Morch, 1853 Type: Tridacna crocea Lamarck, 1819 Tridacna crocea displays in accentuated form the characteristics of this subgenus which are shown in varying degrees by all its members. They are bys- siferous, possess guard tentacles around the edge of the incurrent opening, have reduced outer demi- branchs which lack a food groove, and their valves range from being nearly equilateral with the umbos nearly central in position (T. squamosa) to strongly inequilateral ( T. maxima and crocea ) with the umbos displaced far anteriorly, never posteriorly as in Persikima. All species of Chametrachea bore into coral, this tendency being most evident in crocea. All possess a capacity for producing scaly shell sculpture, the scales becoming more reduced as the boring habit is intensified. The Recent spe- cies are arranged here in order of increasing spe- cialization. Synonymy — [1753 Chamaetrachaea Klein, Tentamen Methodi Ostra- cologicae, p. 149 (pre-Linnaean).] 1853 Chametrachea Morch, Catalogus Conchyliorum Comes de Yoldi, part 2, p. 56. Type by subsequent designation (Iredale, 1937): Tridacna crocea Lamarck. 1857 Chametrachaea H. and A. Adams, The Genera of Re- cent Mollusca, vol. 2, p. 464; refer to C. scapha Meuschen as an example, and their plate 113, fig. 2, 2a, 2b [= T. maxima Roding]. 1937 Flodacna Iredale, Australian Zoologist, vol. 8. part 4, p. 238. Type by original designation: Tridacna squamosa auct. [= T. squamosa Lamarck], 1937 Sepidacna Iredale ibid., p. 239. Type by original designation (ibid., p. 261): Tridacna troughtoni Iredale, 1927 [= T. maxima Roding] . 1937 Vulgodacna Iredale, ibid., p. 239. Type by original designation: Tridacna maxima var. fossor Hedley, 1921 [= T. maxima Roding], Tridacna loczyi Kutassy, 1934 Range — Pliocene (?) of Celebes, Indonesia. Remarks — Tridacna loczyi is probably ancestral to T. squamosa. The worn scales and shape of the fossil resemble those of large, old specimens of the latter species, as does its length of 380 mm. Synonymy — 1934 Tridacna loczyi A. von Kutassy, Verhandelingen Ge- ologisch-Mijnbouwkundig Genootschap voor Neder- land en Kolonien, Geologische Ser., deel 10, p. 313, pi. 6, fig. 5, pi. 7, fig. 1 (Kaoeroe, Celebes). Tridacna mbalavuana Ladd, 1934 (PI. 284) Range — Upper tertiary, Viti Levu, Fiji. Remarks — As Ladd suggested, the holotype of T. mbalavuana is apparently a young specimen, and it is difficult to determine its relationship with any Plate 284. Figs. 1, 2. Tridacna ( Chametrachea ) mbalavuana aegyptiaca Chenu, from Gulf of Suez, Egypt; Pleistocene (?) Ladd, from south side Walu Bay, Viti Levu, Fiji; Neogene; (reduced from Chenu, 1845, pi. 7, figs. 1, 2) (Smithsonian Holotype (53.1 mm., BPBM Geol. No. 1124) (from Ladd, photos). 1934, pi. 31, figs. 2, 3. Figs. 3, 4. Tridacna ( Chametrachea ) [62 - 067] 380 Chametrachea Joseph Rosewater Tridacnidae Recent species. The elongation of the hinge line, and posterior displacement of the umbos, suggest a relationship with Persikima. However, the widely spaced scales and characteristics of the radial sculp- ture indicate that the species is closer to T. squa- mosa in the subgenus Chametrachea. Synonymy — 1934 Tridacna mbalavuana Ladd, Bernice P. Bishop Mu- seum Bull. 119, p. 185, pi. 31, figs. 2, 3; holotype: BPBM Geol. No. 1124 (south side Walu Bay, Viti Levu, Fiji). Tridacna aegyptiaca Chenu, 1845 (PI. 284) Range — Fossil (?), Gulf of Suez. Remarks — This well-worn specimen closely re- sembles Tridacna squamosa. The single striking fea- ture, the deeply cut fold interstices, may be the result of breakage and weathering. It is doubtful that T. aegyptiaca lived earlier than the Pleisto- cene, and this conjecture is made more plausible by the statement of Chenu that the specimen is em- bedded in a matrix of recent conglomerate. Synonymy — 1845 Tridacna aegyptiaca Chenu, Illustrations Conchylio- logiques, Paris, vol. 2, Tridacna, p. 2, pi. 7, figs. 1, 2 (Gulf of Suez, Egypt). Tridacna squamosa Lamarck, 1819 (Pis. 263, 267, 285) Range — East Africa to eastern Melanesia. Remarks — The Scaly or Fluted Giant Clam is one of the strikingly beautiful inhabitants of tropi- cal Indo-Pacific coral reefs. The shell can display exquisite flutes, and certain of the animals’ patterns of coloration are extremely lovely (see plates 263 and 285). Tridacna squamosa can be distinguished from all other species in the family by its characteristic sculpture of projecting, broad, leaf-like scales (“fluted sculpture,” R. D. Turner in Shrock and Twenhofel, 1953, p. 378). Its shell is more equi- lateral then those of T. crocea and maxima, the other members of the subgenus Chametrachea, and it has a smaller byssal orifice. From T. gigas and H. hippopus it differs in lacking a complete outer demibranch, in its fluted sculpture, and in having a heavier shell for its size. Tridacna squamosa is also usually attached by a weak but copious byssus, whereas the former two species usually lack this at- tachment as adults. Tridacna (Persikima) derasa can be confused with T. squamosa, since T. derasa sometimes produces low (though continuous) pro- jecting concentric sculpture. The shell of T. derasa, Plate 285. Tridacna (Chametrachea) squamosa Lamarck. note cone-like excurrent siphon ( center ) and guard tentacles Dorsal view of living animal from Muti Island, Eniwetok of incurrent siphon (right [posterior]) (length: 235 mm.) Atoll, Marshall Islands (on coral head in 12 feet of water); (author’s photo). [62 - 068] April 30, 1965 INDO-PACIFIC MOLLUSCA, vol. 1, no. 6 Tridacna 381 however, is usually inequilateral with the umbos displaced posterior to center. Umbonal displace- ment in T. squamosa, when present, is usually ante- rior. The distinctive arrangement of the anterior ctenidial tracts of derasa can be depended upon to distinguish it further from squamosa (see Remarks and Description, T. derasa ) . Due to the plasticity of the shells of Tridacnidae, it was implied by Hedley ( 1899 ) that T. gigas and T. squamosa are the same species and (in 1921) that “two or more species distinct in youth may converge in age til they are alike Giant Clams” (also see Dautzenberg, 1929, p. 586). Although McLean (1947) maintained squamosa and gigas as separate species, he admitted the possibility “that T. squamosa may in senility lose the scales and be- come a ‘giant clam’ resembling the adult gigas.” The largest specimen of squamosa known to me (40.8 cm., from Thailand) bears little resemblance to gigas. Despite the fact that the scales of large individuals of T. squamosa can become worn down to some degree, these individuals are always dis- tinguishable from T. gigas. The species elongatissima, described by Bianconi from Mozambique, has an elongate shell and nor- mally has one or two additional radial folds. This apparent phenotype of squamosa is also present in the Red Sea and occurs occasionally elsewhere in the range of the species. Because it does not have any geographic limitations and appears to inter- grade with the typical squamosa, elongatissima is not recognized even as a subspecies. Holthuis ( 1952 ) reported the following shrimps of the family Palaemonidae to be commensal with T. squamosa: Anchistus miersi (DeMan) and Conchodytes tridacnae Peters. An additional species of Anchistus, near A. demani Kemp, was present in T. squamosa collected by the writer at Eniwetok Atoll, Marshall Islands (identified by Dr. F. A. Chace, USNM). A brachyuran crab, Xanthasia murigera White ( Pinnotheridae ) , was also found in the mantle cavity of a specimen of squamosa col- lected by the writer at Ko Sindarar Nua, Thailand (identified by Dr. R. B. Manning, USNM). Habitat — On the surface of coral reefs, usually in somewhat protected localities; in reef moats (Stephenson, et al., 1931). Color Notes — Mantles of Tridacna squamosa are variable in color and pattern. The following notes were made from a young and perhaps atypical specimen collected at Eniwetok Atoll measuring 24 cm. in length (see pi. 285). The main ground color of the mantle is dark grayish purple (Kelly and Judd, 1955, ISCC-NBS color name, C553 number 229); along the outer man- tle margin is a row of rhomboidal spots which are very light bluish green (C553 number 162); the larger irregularly-circular spots toward the center of the mantle are multicolored, their outer periphery being pale yellow (number 89), inside of which is a band of dark yellow ( number 88 ) , and entire center is near- est to light blue (number 181). Around edges of incurrent opening there is a series of the larger spots and elongate ones are also present on excur- rent opening. In older specimens the main, purple ground color seems to persist, although only one other specimen resembling that described above was seen by the writer at Eniwetok (Feb.-Mar. 1963) and none were seen later in the year (Oct.-Dee.) in waters of Malaysia, Thailand or Indonesia. The more typi- cal pattern seems to consist of smaller overall speckling, blotching and striping with yellow or tan (see plate 263). Posteriorly within the mantle cav- ity, the ctenidia of all specimens examined were brown. Appearance and Activity: Visible hyaline organs are distributed solely along the edges of the mantle. There is an irregular row of small ones on the man- tle margin, and medially another more sparse row of larger ones, some of which are on small mounds of mantle tissue. When undisturbed the mantle is reflected over the edges of the gaping valves ( plate 285 ) . The in- current opening is elongate and partially obscured by the tentacles guarding it. The excurrent open- ing is located on a cone-shaped mound of mantle tissue and is usually directed anteriorly. When the clam is disturbed a forceful jet of water can be emitted from the excurrent opening. The tissues surrounding the excurrent opening appear to be quite sensitive to an interruption of light rays. The animal portrayed in plate 263 does not appear nor- mal and may have been near death. Description — Shell reaching 408 mm. ( about 16 inches) in length; semicircular in outline; the angle formed at the umbos by the hinge line and ventral margin usually greater than 150°; strongly inflated and having medium-sized to small byssal orifice. Valves moderately heavy and thick; color usually grayish white but occasionally tinged with orange; encrusted with vermetids, annelid tubes, coral, coralline algae, other organisms and debris. Surface of valves dull. Primary radial sculpture consists of [62 - 069] 382 Cliametrachea Joseph Rosewater Tridacnidae from 4 to 12 strongly convex, rib-like folds which become crowded and obsolete on ventral slope; one fold, usually the second to fourth from anterior end, is positioned almost vertically; the others slant obliquely in an anterior or posterior direction. Usu- ally 5 or 6 of the folds are very strong. Secondary radial sculpture consists of evenly-spaced riblets on both convex folds and their relatively broad inter- stices (more noticeable on the latter), interrupted by fine concentric lines of growth resulting in either a minute reticulate sculpture or in the formation of small nodules where riblets and growth lines cross. Concentric sculpture consists of undulate lines of growth which produce widely spaced, broadly leaf- like, projecting scales on primary folds (in young specimens scales are usually semitubular). Scales usually worn or broken on ventral half of shell; but everywhere are quite delicate and easily broken. Dorsal shell margins broadly fan-shaped in outline, undulate, with series of 4-6 pointed to bluntly rounded, medially projecting, crenulated, interdig- itating processes which represent extremities of rib interstices. Hinge line usually about half the length of shell. A single, oblong cardinal tooth present in each valve; 2 elongate posterior lateral teeth pres- ent in right valve and 1 in left. Ligament second- arily prosodetic. Umbos directed postero-medially. Ventral margin slightly concave in region of byssal orifice, convex posteriorly. Edge of byssal orifice with series of 6-8 distinct to obsolete plicae, usually about evenly spaced. Posterior to byssal orifice ven- tral margin can be crenulate. Ventral slope rela- tively broad, flat, with crowded radial sculpture and often partially covered by overgrowth of por- celaneous material from edge of byssal orifice. In- terior of valves porcelaneous; white or occasionally tinged with orange. Pallial line entire, moderately wide. Posterior adductor muscle scar large, sub- circular, adjacent to pallial line dorsally and located posterior to center. Posterior pedal retractor muscle scar usually about half the size of posterior adduc- tor scar (or smaller) and contiguous to it; located just anterior to center. Area within pallial line, ex- cluding muscle scars, dull; pallial line, muscle scars and areas to the periphery of shell shiny. Prodisso- conch unknown. Measurements (mm.) — length height width 408 320 280 large; Ko Sindarar Tai, Thai- land 387 263 240 large; Eniwetok, Marshall Is- lands 276 179 186 large; Rongerik, lands Marshall Is- 208 149 122 average; Onotoa, lands Gilbert Is- 129 78 60 small; Zanzibar 89 Synonymy 60 47 small; Zanzibar [1787 Hippopodes scaphe Meuschen, Museum Geversia- num, p. 430 (part); non-binomial]. 1798 Tridacnes imbricata var. a Roding, Museum Bolteni- anum, p. 172, no. 191, refers to Chemnitz, vol. 7, figs. 492-493. 1807 Tridacne imbricata Link, Beschreibung Naturalien sammlung Univ. Rostock (3) p. 153 (no locality given); refers to Chama gigas Linne, Gmelin, p. 3299, and to Chemnitz, vol. 7, pi. 49, figs. 492 (- 493) [= T. squamosa ] 494 [= T. maxima ] (part). Non Roding, 1798. 1819 Tridacna squamosa Lamarck, Histoire Naturelle des Animaux sans Vertebres, vol. 6, part 1, p. 106 ( l’Ocean indien [Amboina, Moluccas, here re- stricted]); refers to Rumphius, pi. 43 [42], fig. A; Tableau Encyclopedique Methodique, pi. 235, fig. 4, pi. 236, fig. la, b; and others. 1845 Tridacna squamoza ‘Lamarck’ Chenu, Illustrations Conchyliologiques, vol. 2, Tridacna, caption, pi. 6, fig. 5 [misspelling], 1856 Tridacna elongatissima Bianconi, Memorie della Ac- cademia delle Scienze dell ’Instituto di Bologna, vol. 7, p. 408, pi. 25, fig. 2 (Mozambique). 1903 Tridacna lamarcki Hidalgo, Memorias Real Academia de Ciencias, Madrid, vol. 21, p. 385 (Mindoro, Cebu, Mindanao, Paragua [Palawan] (part); re- fers to Sowerby, Thes. Conch., pi. 485 (Tridacna, pi. 1) fig. 2, and others. Types — The location of the holotype of Tridacna squamosa Lamarck is unknown. It is not in the Museum National d’Histoire Naturelle, Paris, ac- cording to Lamy (1932), nor is it in Geneva (E. Binder, in litt., 1961). Rumphius’ plate 43 [42], fig. A, is here designated the lectotype of Tridacna squamosa Lamarck, 1819. The type locality, “l’Ocean indien,” mentioned by Lamarck, may be inter- preted to include the East Indies. It is here re- stricted to Amboina, Moluccas, from which Rumphius’ specimen undoubtedly came. The loca- tions of the types of T. imbricata var. a Roding and T. imbricata Link are unknown. Nomenclature — This species has been recognized generally without question under the name squa- mosa since it was described. Lamarck’s words (here translated): “large, erect, distant scales; interstices of the ribs multistriate,” leave little doubt as to its identity. Of the figures cited by Lamarck, Rum- phius, pi. 43 [42], fig. A; Gualtieri pi. 92, fig. F; and those from the Encyclopedique Methodique are un- doubtedly this species; Gualtieri, pi. 93, fig. B and the Knorr and Chemnitz figures, unfortunately, are of T. maxima. [62 - 070] April 30, 1965 INDO-PACIFIC MOLLUSCA, vol. 1, no. 6 Tridacna 383 Plate 286. Geographical distribution of Tridacna (Chame- trachea) squamosa Lamarck; solid dots: specimens examined; open circles: from the literature. Records — MOZAMBIQUE: Saco coral reef, Inhaca Is- land, Delagoa Bay (Macnae and Kalk, 1958). ZANZIBAR: Mnazi Mmoja, Zanzibar City (ANSP; USNM); Fumba; Chango Island (both ANSP). KENYA: Diani Beach, Mom- basa (USNM). RED SEA: Ras Banas, Egypt; Sharm Ubhar and Jidda, both Saudi Arabia (all USNM). MADAGASCAR: between Ambariotelo and Ambariobe, southeast Nosy Be (ANSP; USNM); Grand Recif, Tulear (R. W. Foster, ANSP). SEYCHELLES: (Rost and Soot-Ryen, 1955, p. 14). MAURITIUS: (USNM); REUNION: (Deshayes, 1863, pp. 21-22). MALDIVE ISLANDS: Kagi Island; Imma Is- land, both North Male Atoll; Wala Island, South Nilandu Atoll ( all International Indian Ocean Expedition, 1964, R. Robertson, ANSP). CHAGOS ARCHIPELAGO: (Melvill, 1909, p. 135). THAILAND: Ko Sindarar Nua (Chance Is- land); Ko Sindarar Tai (both International Indian Ocean Expedition, 1963, USNM); Ko Phuket, (MCZ; ANSP), all Andaman Sea; Ko Tao (USNM); Ko Khram (G. M. Moore, MCZ) both Gulf of Siam. MALAYSIA: Pulau Tekukor, northwest of St. John’s Island, Singapore ( International Indian Ocean Expedition, 1963, USNM). JAPAN: Hiogo [Kobe area] (Pilsbry, 1895, p. 130, ex Lischke, 1869, p. 160). PHILIPPINES: Grand Island Reef, Subic Bay; Cor- regidor, Manila Bay (both USNM); Matabungkay Cove, 14 km. south of Nasughu, Batangas Province; Tabaco, Albay Province; Barrio Lupi, Prieto Diaz, Sorsogon Province (all ANSP); Badang, near Gubat, Sorsogon Province, all Luzon (MCZ; USNM); Lubang Island (USNM); Ambil Island, both Lubang Islands (MCZ); Boac; near Cabuyo, both Marinduque (both USNM); Ibajay, Panay (MCZ); along Rio Jordan, Guimaras; Cebu, Cebu (both USNM); Olango Island, east side of Cebu (ANSP); Cagayan [off Negros Occidental]; Siaton, Negros Oriental (both USNM); Bo- rongan Village, east side of Samar Island (ANSP); Santa Cruz Island, off Zamboanga (USNM); Gulf of Davao, Mindanao Island (ANSP); Mantangal, Basilan; Lampinigan Island, off northwest Basilan; Chase Head, Endeavor Strait, Palawan; Marongas Island, off Jolo (all USNM); Bongao Channel, southwest end Sanga Sanga Island, both Sulu Ar- chipelago (ANSP). INDONESIA: South Pagi Island, Veeckens Bay, southwest of Sumatra (International Indian Ocean Expedition, 1963, USNM); Tjiperwagaran, Bantam, Java (USNM); Batjan Island; Amboina, both Moluccas (both MCZ). NORTH BORNEO: Jesselton (ANSP); Pulau Tiga (USNM); Pulau Mandidarah (ANSP); off Pulau Bohaydulong, Sempoma (USNM) (all Mary Saul, collector). AUSTRALIA: Broome, Western Australia; Murray Id., Torres Straits, Queensland (both Hedley, 1921, p. 172); Little Lagoon. Groote Eylandt, Gulf of Carpentaria (USNM). NEW GUINEA: Poelau Rouw; Poelau Maransa- badi, both Aoeri Islands; 1M miles southwest of Biak, Poelau Biak, Schouten Islands; Poelau Noesi; Poelau Konori, both Mios Woendi, Padaido Islands (all ANSP); near Hollandia (USNM) (all West Irian). ADMIRALTY ISLANDS: See- adler Harbor, Manus Island (MCZ). NEW CALEDONIA: Noumea (ANSP; USNM). FIJI: Yasawa Islands (USNM); Nananu I-Ra; Ngaloa, 3 miles west of Rovondrau Bay (both MCZ); Suva (USNM) all Viti Levu; Mbengga Island; outer reef, 3 miles southeast of Tunuloa, Vanua Levu; Dravalau, Totoya (all MCZ). MARIANAS ISLANDS: Unai Dikiki Matuis, Saipan (USNM); east Tinian (MCZ). PALAU IS- LANDS: Babelthuap (USNM; MCZ); Koror (BPBM; MCZ; ANSP; USNM); Abappaomogan, Vk miles west of Eil Malk (F. M. Bayer, 1955, USNM); Eomogan (MCZ). MAR- SHALL ISLANDS: Igurin Island, Eniwetok; Yurochi Is- land, Bikini; Burok Island, Rongelap; Letoback Island, Rongerick; Taka; Ailuk (all USNM); Majuro (BPBM); GILBERT ISLANDS: Onotoa (USNM). ELLICE IS- LANDS: Funafuti (Hedley, 1899, p. 504). SAMOA: (ANSP). TONGA: Sopu Reef (L. J. Lancaster, USNM); Niuafou (USNM). TUAMOTUS: doubtful record and pos- sibly fossil specimens. Fossil records — FRENCH SOMALIA: Pleistocene, 1.5 meters west of Obock (Abrard, 1942). DAITO ISLANDS ( south of Japan ) : Pliocene-Pleistocene, Kita Daito Jima (Nomura and Zinbo, 1935). EAST INDIES: Quaternary, Celebes (Schepman, 1907, p. 197). TONGA: Pleistocene (?), Vavau (Mansfield, 1926; USNM 352426). TUAMOTUS: Pleistocene, (?), Toau Atoll (ANSP); Pleistocene (?), Raroia Atoll (USNM). Tridacna besairiei Collignon, 1951 (PI. 287) Range — Upper Cretaceous, Antonibe, Madagas- car. Remarks — The worn and broken fragment ( pi. 287) of this species resembles most closely T. max- ima (Roding). Collignon remarked in the descrip- tion of the species that the age of this fossil is ques- tionable because no other members of the family have been found in strata older than the Tertiary. If the age is valid, then T. besairiei is the oldest fos- sil tridacnid. [62 - 071] 384 Chametrachea Joseph Rosewater Tridacnidae Synonymy — 1951 Tridacna besairiei M. Collignon, Annales Geologiques Du Service Des Mines, Bureau Geologique de Madagascar, Fascicule No. 19, Paris, p. 98, pi. 15 (3), figs. 15, 15a (Antonibe, Madagascar). Tridacna media Pusch, 1837 (PI. 287) Range — Upper Tertiary beds of Warsaw, Poland. Remarks — This fossil species appears to be most closely allied with T. maxima, although the surface sculpture of the specimen illustrated is very worn. This and T. wolfarti Chenu are the only fossil species reported from European deposits closely resembling Recent Tridacnidae. These species inhabited what must have been more expansive and tropical Euro- pean seas during the Tertiary. Synonymy — 1836 Tridacna media Pusch, Polens Palaontologie (1), p. 55, pi. 6, fig. 6a, b, c (Tertiary beds, Warsaw, Po- land ) . Tridacna maxima (Roding, 1798) (Pis. 263-265; 267-269; 288-290) Range — East Africa to eastern Polynesia except Hawaii. Remarks — This species is the most ubiquitous of the family, inhabiting reefs throughout the tropical Indo-Pacific. Both the color of the exposed mantle and the shape and sculpture of the valves are ex- tremely variable ( see plate 263 ) . Tridacna maxima can easily be distinguished from T. gigas and T. squamosa by its strongly inequilat- eral valves, the hinge line being considerably shorter than the ventral margin; sculptural differences also readily separate maxima from these species. It is most closely related to T. crocea, but differs in hav- ing more raised external valve sculpture, subequal posterior adductor and byssal retractor muscle scars, and a byssal orifice which is usually relatively shorter than that of crocea. Field observations indicate that there is consider- Plate 287. Figs. 1, 2. Tridacna (Chametrachea) besairiei 15, 15a). Figs. 3-5. Tridacna (Chametrachea) media Pusch, Collignon, from Antonibe, Madagascar, Upper Cretaceous; from Tertiary beds, Warsaw, Poland; Holotype (from Pusch, Holotype (36 mm.) (from Collignon, 1951, pi. 15 (3), figs. 1836, pi. 6, fig. 6a, b, c) (Smithsonian photos). [62 - 072] INDO-PACIFIC MOLLUSCA, vol. 1, no. 6 Tridacna 385 April 30, 1965 Plate 288. Tridacna (Chametrachea) maxima showing vari- ation in shape and sculpture of shell. Figs. 1, 2. Specimen from Eniwetok, Marshall Islands (133 mm. iength). Figs. 3, 4. From Ani Jima, Bonin Islands (injured anterodorsally; 105 mm. length). Figs. 5, 6. From Celeo Island, off North- East New Guinea (120 mm. length). Figs. 7, 8. From Nosy Be, Madagascar (127 mm. length). Muscle scars and pallial lines outlined. Figs. 1, 3, 5, 8 are left valves; figs. 2, 4, 6, 7 are right valves (Smithsonian photos). [62 - 073] 386 Chametrachea Joseph Rosewater T ridacnidae able overlap in mantle coloration and pattern be- tween maxima and crocea. Both species are exceed- ingly variable in these characters, but this may be due to convergence where these characters are sub- ject to no strong selective pressures. Mantle color in T. maxima runs the gamut of brilliant green, blue, purple and brown with great pattern variation ( see plate 263, fig. 3; and plate 264; also Gillett and Mc- Neill, 1959, p. 87; for comparison with T. crocea, see Franc, 1960, pi. 9 ) . In widely separated geographic regions color bril- liancy may vary in T. maxima. At Eniwetok, Mar- shall Islands, the mantle colors were observed to be extremely bright (personal observation, Feb.-March, 1963 ) . This seems also to be true on the Great Bar- rier Reef of Australia (Gillett, et al., op. cit.). But on the shores of the Andaman Sea in Malaysia and Thailand, and off southern Sumatra (Oct.-Dec., 1963) the colors were noted to be more subdued (similar to crocea, Franc, op. cit.). This phenome- non may be due to differences in the conditions of the animals at the time the observations were made, to a real geographic variation based on genetic dif- ference, or to environmental factors. It is evident from the study of a large series of T. maxima from many localities throughout its range that several well-known forms intergrade with this species (see Synonymy). Outstanding among the forms here considered synonyms of maxima, is T. elongata Lamarck. This, as the name suggests, tends to be elongate (see plate 289). Forms like this occa- sionally occur throughout the Indo-Pacific and show no particular geographic pattern. McLean (1947) distinguished this form from maxima. However, no consistent differences have been noted in the present study. The elongate form can be connected with more typical maxima by intermediates. The extreme variability of T. maxima, responsible at least in part for the many named forms, can be explained on the basis of the species’ ecology. The clam fives firmly attached to the coral reef. Where sufficient space is available anteriorly and posteri- orly, the valves may become more or less elongate; but where growth is blocked by a coralline obstacle, the valves may become stunted or malformed. Such malformations are common, (see plate 288). Also, apparently depending on the substrate, individuals may burrow fairly deeply, although never becoming imprisoned as does T. crocea. The depth to which a clam has burrowed can be interpreted from the parts of the valves which have been smoothed by erosion. The gross morphology of T. maxima is very similar to that of T. crocea. At maturity both are smaller animals than other Tridacnidae; the outer demi- branchs are incomplete and grossly similar in struc- ture. Hyaline organs tend to be concentrated along the edge of the mantle on papillae in maxima, while in crocea they are more diffuse on the mantle sur- face. In maxima posterior pedal retractor and adduc- tor muscles are subequal in circumference, while in crocea the posterior pedal retractor is about half the size of the adductor. The following species of shrimps ( Palaemonidae : Pontoniinae) were found commensal in the mantle cavity of T. maxima: Conchodytes tridacnae Peters (Ko Huyong, Similan Islands, Thailand); and An- chistus demani Kemp ( Ko Sindarar Nua, Thailand ) (identified by Dr. R. B. Manning, U. S. National Museum ) . Habitat — Lives in shallow water on reefs, par- tially imbedded in coral. Plate 289. Tridacna ( Chametrachea ) maxima (Roding). Lectotype of T. elongata Lamarck. Fig. 1. Exterior of left valve. Fig. 2. Interior of right valve. Fig. 3. Ventral view showing byssal orifice. (15 cm. length) (photo courtesy of E. Binder, Museum d’Histoire Naturelle, Geneva). [62-074] April 30, 1965 INDO-PACIFIC MOLLUSCA, vol. 1, no. 6 Tridacna 387 Color notes — Mantles of T. maxima ( from Eni- wetok) showed great variation in color and pattern (see plate 263, fig. 3; and plate 264). Often there is a peripheral band of color containing numerous hyaline organs; the main mantle color occurs me- dially and is sometimes spotted and streaked with other colors. Some of the main mantle colors noted were: grayish yellow (Kelly and Judd, 1955, ISCC- NBS Color Name; C553 number 90); light bluish green ( C553 no. 163 ) ; blackish blue ( C553 no. 188 ) ; grayish violet (C553 no. 215); and blackish purple (C553 no. 230). Narrow peripheral bands were noted in the following colors: light olive brown (C553 no. 94); grayish greenish yellow (C553 no. 105); and brilliant green (C553 no. 140). Spots and streaks over the surface of the mantle were noted to be: bril- liant bluish green ( C553 no. 159 ) ; and brilliant blue ( C553 no. 177 ) . Other colors were observed and it is unlikely that any two specimens of T. maxima are exactly alike in this regard. The cause of mantle coloration in Tridacna is es- pecially interesting in the cases of the variable spe- cies T. maxima and crocea. It is probably caused by the zooxanthellae. Specimens of Tridacna, probably maxima, were observed “with dead coral slabs rest- ing on their shells, beneath which the mantles lacked pigmentation, i.e., were cream-white” (personal communication R. Robertson, 1964, in Maidive Is- lands). It appears that the zooxanthellae were un- able to live in the shaded portions, thus leaving only the whitish mantle color. Mantle pattern variation apparently is based on the distribution of the algae in the tissues. Activities — Specimens of T. maxima were col- lected from the reef of Eniwetok Atoll and main- tained in the laboratory for 5-6 weeks in running water aquaria. Specimens were removed periodi- cally for photographing and examination. It was noted that the animals reattached byssal threads to the aquarium floor each time they were replaced. Tridacna maxima reacted to an interruption in light intensity by withdrawing the mantle and closing its valves. Description — Shell reaching 353 mm. ( about 13 7/8 inches) in length; elongate- to short-obtuse-tri- angular in shape, angle formed at umbos by margins of hinge and ventral border usually less than 150°, moderately to strongly inflated and having a large Plate 290. Tridacna (Chametrachea) maxima ( Roding). Figs. 3-5. Holotype of T. acuticostata Sowerby, from the Philip- 1, 2. Lectotype of T. compressa Reeve (length: 107 mm.). Figs. pines (length: 132 mm.) (photos courtesy of R. T. Abbott). [62 - 075] 388 Chametrachea Joseph Rosewater Tridacnidae byssal orifice. Valves heavy, thick, color usually gray- ish white but often tinged with yellow or pinkish orange; often incrusted with vermetids, annelid tubes, coral, coralline algae, other organisms and debris. Surface of valves dull or faintly shiny in cleaned specimens. Primary radial sculpture con- sisting of from 6 to 12 broad, moderately convex rib- like folds becoming crowded and obsolete on ventral slope. Usually 6 or 7 of these folds are very strong. Secondary radial sculpture consisting of from 10-20 evenly spaced riblets on folds and 3-7 in their narrow interstices, often interrupted on primary folds by concentric sculpture and here having nodular ap- pearance; in smooth specimens, where folds are low or worn down, secondary sculpture in interstices ap- pears more prominent and here groups of riblets may simulate raised radiating bands. Concentric sculp- ture consisting of closely spaced undulate lines of growth which produce crowded, but usually low scales on primary folds. Scales often worn off near umbos and here undulate pattern is most evident. Dorsal valve margins undulate, with a series of about 5 usually sharply triangular, medially projecting, in- terdigitating processes which represent extremities of rib interstices. Hinge line short, less than one half length of shell. A single, oblong cardinal tooth is present in each valve; 2 elongate posterior lateral teeth present in right valve and 1 in left. Ligament secondarily prosodetic. Umbos directed postero- medially. Ventral margin varying from considerably to only slightly concave in region of byssal orifice and convex posteriorly. Edge of byssal orifice with series of 4 to 9 ( usually about 6-7 ) distinct to obso- lete plicae closely spaced near umbos and becoming more distantly separated posteriorly. Ventral slope narrow, rounded or hardly to considerably flattened. Interior of valves porcelaneous, usually white but occasionally tinged with yellow or pinkish orange near dorsal margin. Pallial line entire and relatively wide. Posterior adductor muscle scar large, subcir- cular, adjacent to pallial line dorsally and located posterior to center. Posterior pedal retractor muscle scar larger than half the size of posterior adductor- scar ( usually subequal ) anteriorly contiguous to the latter, and approximately centered over byssal ori- fice. Area within pallial line, excluding muscle scars, dull; area of line, muscle scars and from there to periphery of shell shiny. Prodissoconch with its umbones directed medially and slightly anteriorly, somewhat inflated; in shape reminiscent of Cardium ; with slight anterior wing and sculptured with micro- scopic, incised, concentric lines; 1.0 mm. in length (plate 268). Measurements (mm.) — length height width 329 177 186 large; Palmyra, Line Islands 267 167 137 large; Canton Island, Phoenix Islands 202 107 103 average; Port Blair, Andaman Islands 169 78 81 average; Inner Sindu Island, Tanganyika 118 76 63 small-average; Uterik Island; Uterik Atoll, Marshall Is- land 85 60 44 small; Onotoa Atoll, Gilbert Islands Synonymy — 1 1787 Hippopodes scutra Meuschen, Museum Geversianum, p. 430; non-binomial]. [1797 Tridachna elongata Humphrey, Museum Calonni- anum, p. 50 (Madagascar); rejected work.] 1798 Tridachnes maxima Roding, Museum Boltenianum, part 2, p. 171, no. 184 (no locality given [Mauritius, here designated]); refers to Gmelin, Chama gigas, species 2 [= Argenville, pi. 23, fig. E, a foliate specimen], and to Chemnitz, vol. 7, pi. 49, fig. 495 [a smooth specimen]. 1798 Tridachnes noae Roding, ibid., p. 171, no. 186 (no locality given); refers to Gmelin, Chama gigas, species 2, and Chemnitz, vol. 7, pi. 49, fig. 494. 1798 Tridachnes imbricata Roding, ibid., p. 172, no. 190 (no locality given); refers to Gmelin, Chama gigas species 2. 1801 Tridacna gigas Lamarck, Systeme des Animaux sans Vertebres, p. 117 (no locality given) (part); re- fers to Chemnitz, vol. 7, pi. 49, fig. 495, and to Tableau Encyclopedique et Methodique, pi. 235, fig. 1; Reeve, 1862, Conchologia Iconica, vol. 14, Tridacna, pi. 1, fig. la, pi. 2, figs, lb, c (Indian and Pacific Seas). 1807 Tridacne imbricata Link, Beschreibung Naturalien Sammlung Univ. Rostock (3) p. 153 (no locality given); refers to Chama gigas Linne, Gmelin, p. 3299, and to Chemnitz, vol. 7, pi. 49, figs. 492(- 493) [= T. squamosa ] 494 [= T. maxima] (part). 1807 Tridacne maxima Link, ibid., p. 153 (no locality given); refers to Chama gigas Linne, Gmelin, and to Chemnitz, vol. 7, pi. 49, fig. 495. 1819 Tridacna gigas Lamarck, Histoire Naturelle des Ani- maux sans Vertebres, vol. 6(1), p. 105 (l’Ocean indien ) . 1819 Tridacna elongata Lamarck, ibid., p. 106 (l’Ocean indien?); Chenu, 1845, 111. Conch., vol. 2, pi. 1, fig. 2, var. /3; pi. 2, figs. 1-3. 1819 Tridacna mutica Lamarck, ibid., p. 106 (l’Ocean des grandes Indes) (part); Chenu, ibid., pi. 4, figs. 1, la [= T. gigas], pi. 6, fig. 1( [= T. maxima], 1845 Tridacna squamosa ‘Lamarck’ Chenu, ibid., pi. 3, fig. 2. 1853 Chametrachea scutrum ‘Meuschen’ Morch, Catalogus Conchyliorum Comes de Yoldi, fascicle 2, p. 56 (East Indies); refers to Gualtieri, pi. 92, fig. E; H. and A. Adams, 1857, Genera of Recent Mollusca, vol. 2, p. 465. 1857 Chametrachea scapha ‘Meuschen’ H. and A. Adams, ibid., p. 465, pi. 113, figs. 2, 2a, 2b (Pacific and Indian Oceans, and China Sea); not C. scapha ‘Meuschen’ Morch = T. crocea Lamarck. 1862 Tridacna rudis Reeve, Conchologia Iconica, vol. 14, Tridacna, pi. 5, fig. 4a, b, c (Philippine Islands). [62 - 076] April 30, 1965 INDO-PACIFIC MOLLUSCA, vol. 1, no. 6 Tiidacna 389 1862 Tridacna compressa Reeve, ibid., pi. 6, fig. 5; pi. 7, fig. 5b, (no locality given [Philippines here desig- nated] ) . 1862 Tridacna cumingii Reeve, ibid., pi. 7, fig. 7b [not fig. 7a = T. crocea ] ( Philippine Islands ) ( part ) . 1884 Tridacna lanceolata Sowerby, Thesaurus Conchyli- orum, vol. 5, p. 181, Tridacna, pi. 6, fig. 19 [18] (Philippines). 1902 Tridachnes umbricata Sherbom, Index Animalium, section 1, part 2, p. 1189 [error for T. imbricata Roding, 1798]. 1903 Tridacna lamarcki Hidalgo, Memorias Real Academia de Ciencias, Madrid, vol. 21, p. 385 (Mindoro, Cebu, Mindanao, Paragua Islands [= Palawan] ) (part); new name for T. gigas Lamarck, not Linne. 1903 Tridacna reevei Hidalgo, ibid., p. 389 (Philippines); new name for T. elongata Reeve, not Lamarck. 1912 Tridacna acuticostata Sowerby, Proceedings Malaco- logical Society of London, vol. 10, p. 30, fig. (p. 31) (Philippines). 1921 Tridacna maxima var. fossor Hedley, Records of Aus- tralian Museum, vol. 13, no. 4, p. 171, pi. 29, fig. 6; pi. 30, fig. 7; pi. 33, fig. 11 (Mast Head Island, Capricorn Group). 1927 Tridacna troughtoni Iredale, ibid., vol. 16, p. 75, pi. 5, figs. 9-10 (Vanikoro); [a young specimen]. 1939 Tridacna corallicola ‘Valenciennes’ Lamy, Journal de Conchyliologie, vol. 83, no. 4, footnote, p. 291 [nomen nudum]. Types — Rodin g’s only figure reference for T. max- ima was to Chemnitz, vol. 7, pi. 49, fig. 495, and this is here selected as the lectotype. Roding did not designate a type locality for maxima. However, in the text accompanying the type figure, Chemnitz, vol. 7, p. 124, stated that this species lives in abundance on the shores of St. Maurice (Mauritius, fide Mar- tens, 1880, p. 183). The island of Mauritius is here designated as the type locality for Tridacna maxima Roding. Three syntypes of T. elongata Lamarck are in the Museum d’Histoire Naturelle, Geneva. Of these the specimen marked “2a” (15 cm. in length) is here designated lectotype (plate 289, figs. 1-3). The length measurement matches exactly that given by Lamarck. It does not seem necessary to restrict the type locality of a synonym of the widespread T. maxima. The types of T. matica are presumed lost, and the figures of Chenu indicate that it is probably a composite species. Reeve figured three syntypes of T. compressa with the original description, and that figured in his plate 6, fig. 5, is here designated lectotype ( plate 290, figs. 1,2). The type locality here designated is the Philip- pines, a logical place from which this specimen could have come. The types of Reeve and Sowerby are in the Rritish Museum (N.H.), with the exceptions of T. rudis Reeve and T. lanceolata Sowerby which are also apparently lost. These species must be based on the figures of the respective authors. Types of Hed- ley and Iredale are in the Australian Museum, Sydney. Nomenclature — Tridacna maxima Roding ap- pears commonly in the literature under several syn- onyms, of which perhaps the best known is T. elon- gata Lamarck. The name maxima, however, has priority. Roding’s figure reference shows the ventral portion of the valves of a smooth specimen. Iron- ically, Roding called this species “the largest Claw- mussel.” Although it does not reach anything like the overall dimensions of T. gigas Linne, maxima shares with T. crocea Lamarck the capacity for de- veloping larger byssal orifices than other Tridacna. Records — SOUTH AFRICA: Chaka’s Rock, north coast Natal (Barnard, 1964). MOZAMBIQUE: Ilha da Inhaca; Ilha do Bazaruto (both MCZ; ANSP); Ilha Santa Carolina, both Bazaruto Bay (MCZ); Mozambique City (ANSP). TANGANYIKA: Mboa Magi, 9 miles south of Dar es Salaam (R. T. Abbott, USNM). ZANZIBAR: Jembiani, 5 miles south of Paje; Chumbe Island; Kizimkazi; Mnazi Mmoja; Kiwengwa (all ANSP). KENYA: Diana Beach, 20 miles south of Mombasa (USNM; MCZ; ANSP). RED SEA: Suez (USNM); Faraun Island, Gulf of Aqaba (Rees, et al., 1952, p. 199); Sharm Ubhar; Jidda Harbor, both Saudi Arabia (both USNM); Massawa, Eritrea (MCZ). PERSIAN GULF: (Mevill and Standen, 1907, p. 840). MADAGASCAR: Between Ambariotelo and Ambariobe, southeast Nosy Be (ANSP; USNM); northeast end Nosy Kalakajoro, Radama Islands; Grand Recif, Tulear (both ANSP); southeastern point lie Ste. Marie (MCZ); Tamatave (ANSP). SEYCHELLES: (ANSP). MAURITIUS: barrier reef, 1 mile southeast of Aigrette Island, Mahebourg; Flic en Flacq (both V. Orr, ANSP). MALDIVE ISLANDS: Hulule (E. A. Smith, 1903, p. 624); Kagi Island; Imma Island, both North Male Atoll (both International Indian Ocean Expedition, R. Robertson, 1964). CHAGOS ISLANDS: ( Melvill, 1909, p. 135). CEYLON: Ara Point, Nilaveli, Trincomalee (USNM); Galle (G. and M. Kline, 1956, ANSP). ANDAMAN ISLANDS: Port Blair (USNM). THAILAND: Ko Sindarar Nua (Chance Island); Ko Hu- yong, Similan Islands (both International Indian Ocean Ex- pedition, 1963, USNM); Ko Phuket, all Andaman Sea (MCZ; ANSP; USNM); Ko Khram, Gulf of Siam (G. M. Moore, MCZ). MALAYA: Singapore (ANSP). CHINA: Mirs Bay, Honk Kong (ANSP). JAPAN: Tosa, Shikoku (ANSP). RYUKYUS: Okinawa (USNM; MCZ; ANSP); northeast coast Iheya Shima (ANSP). TAIWAN: west coast Lan Yu, 45 miles southeast of Taiwan (USNM). PHILIP- PINES: Basco, Batan Island, Batanes Group; Port San Pio Quinto, Camiguin Island (both USNM); Bolinao, Pangasi- nan Province; Iba, Zambales Province; Tabaco, Albay Prov- ince; Gigmoto, Catanduanes Island (all ANSP); Barrio Rizal, near Gubat, Sorsogon Province, all Luzon (ANSP; MCZ; USNM); Tagbac Point; Tilic, both Lubang Island (both MCZ; USNM); Calapan, Mindoro (P. de Mesa, MCZ); Cebu City, Cebu; Olango Island, between Cebu and Bohol (both ANSP); Calangaman Island (USNM); Esca- lente, Negros Occidental (ANSP); Calicoan Island, off southeast Samar (MCZ); Lampinigan Island, off northwest Basilan (USNM); Cuyo Island, Cuyo Group (ANSP); Ca- gayancillo, Cagayan Islands (USNM); Bongao Channel, southwest end Sanga Sanga Island, Sulu Archipelago (ANSP). INDONESIA: Pulau Melila, south of Udjung Batu, Banjak Islands; Pulau Bai, Batu Islands; southwest tip Sanding Island, Mentawai Islands (all International In- dian Ocean Expedition, 1963, USNM); Pulau Tikus, near Bengkulu, all south Sumatra (USNM); Pulau Batjan; Pulau Amboina; Pulau Kasiruta; Pulau Burn, all Moluccas (all MCZ). NORTH BORNEO: Pulau Mandidarah (Mary Saul, [62 - 077] 390 Chametrachea Joseph Rosewater Tridacnidae 'V 1C XI ' 1 & .5^ : ' j — * — 1 £ V,M8>> : • jk J py F \0 . >■ kJ/j £L I 1 1 L-S -X J — - 4l: V . b % [jp* '■ .w ' C00(JI i ....... Plate 291. Geographical distribution of Tridacna (Chame- trachea) maxima (Roding); solid dots: specimens examined; open circles: from the literature. USNM; ANSP). COCOS-KEELING ISLANDS: reef off southeast end West Island and Pulo Maria; 3 miles south- west of New Selma (Home) Island (both International In- dian Ocean Expedition, V. Orr, 1963, ANSP; USNM). WESTERN AUSTRALIA: Augustus Island (MCZ); James Price Point, 35 miles north of Broome; % mile south of Willie Creek 17 miles north of Broome; Broome ( MCZ ) ; mouth of False Cape Creek, La Grange Bay; Quobba Point, 40 miles north of Carnarvon (all V. Orr, ANSP). NORTHERN TERRITORY : Arafura Sea, near Darwin ( A. R. Cahn, ANSP). QUEENSLAND: Murray Islands, Torres Strait; Burkitt Island, Cape York (both MCZ); Hinchinbrook Is- land; Pelorus and Orpheus Islands, both Palm Islands; Hol- boume Island (all H. A. Pilsbry, 1923, ANSP); Hook and Bait Reef, northeast of Hayman Island; Langford Reef, Whitsunday Group; Lupton Island; Hamilton Island, both Cumberland Group (all MCZ); Tryon Island; Heron Island, both Capricorn Group (both USNM). NEW GUINEA: Ambai Anchorage, Ambai Island (ANSP); Poelau Biak, Schouten Islands (ANSP; USNM); Mios Woendi (ANSP; MCZ); Poelau Noesi, both Padaido Islands (ANSP); near Hollandia (USNM) all West Irian; Celeo Island, 5 miles off Aitape, North-East New Guinea (USNM); near Gamadodo, Milne Bay, Papua (USNM). SOLOMON ISLANDS: Bam- batana, Choiseul (ANSP); Roviana Lagoon, New Georgia (MCZ); Ataa District, Malaita (J. van der Riet, ANSP); Kunggava Bay, Rennell Island (ANSP). NEW HEBRIDES: Efate (USNM). NEW CALEDONIA: 5 miles west south west of Gatope Island, Voh; Bourail; Recif Laregnere, east of Noumea; Koe, near Touho; Yate (all G. and M. Kline, ANSP). FIJI: south shore Makondronga Island, Makongai Islands (R. T. Abbott, 1940, MCZ); entrance to Suva Har- bor, Viti Levu; reef off Tohalau, Kambara Island, Lau Group (both H. S. Ladd, USNM). LORD HOWE IS- LAND: (Iredale and Allen, 1940, p. 449). BONIN IS- LANDS: Muko Jima (ANSP); Ani Jima; Chichi Jima (both Y. Kondo, 1949, USNM). MARIANAS ISLANDS: Maug Islands (USNM); Saipan (USNM; MCZ; ANSP); Guam (BPBM; USNM; MCZ; ANSP). PALAU ISLANDS: Ka- yangel Islands; Babelthuap; Malakal Harbor, Koror (all ANSP; MCZ; USNM). CAROLINE ISLANDS: Helen Reef (ANSP); Yap; Fasserai Island, Ulithi; Faraulep; Ifalik; Elato; Ponape; Kapingamarangi (all USNM). WAKE IS- LAND: (ANSP; BPBM; USNM). MARSHALL ISLANDS: Eniwetok; Ujelang; Bikar; Kwajalein; Jaluit (all USNM). GILBERT ISLANDS: Abemama; Onotoa (both USNM). LINE ISLANDS: Kingman Reef (BPBM); Palmyra Island (BPBM; USNM; MCZ; ANSP); Washington Island; Fan- ning Island; Christmas Island (all BPBM; ANSP); Jarvis Island (BPBM; MCZ). ELLICE ISLANDS: Nanumea (USNM). BAKER ISLAND: (BPBM). HOWLAND IS- LAND: (BPBM; MCZ) PHOENIX ISLANDS: Canton Is- land (BPBM; MCZ; USNM); Enderberry Island (ANSP). TOKELAU ISLANDS: Manihiki (fide R. T. Abbott, ANSP). SAMOA: Apia (USNM); Lufilufi (BPBM), both Upolu; Pago Pago, Tutuila (ANSP; USNM). TONGA: Niuafou (USNM). COOK ISLANDS: Aitutaki (USNM; ANSP; MCZ); Rarotonga (ANSP; MCZ); Mangaia (USNM). SO- CIETY ISLANDS: Bora Bora (R. Robertson, ANSP; H. A. Rehder, USNM); Raiatea (ANSP); Point Tefaao, Huahine; Moorea (both USNM; Tahiti (ANSP; USNM). AUSTRAL ISLANDS: Rurutu (Aubert de la Rue, ANSP). TUAMOTU ISLANDS: Takaroa; Pukapuka (both ANSP); Tikehau (H. A. Rehder, USNM); Toau (ANSP); Raroia (J. P. E. Mor- rison, USNM); Fakarava (ANSP; USNM); Tauere (ANSP); Mangareva (MCZ; USNM). HENDERSON ISLAND: (E. A. Smith, 1913). Fossil records — FRENCH SOMALIA: Pleistocene, 1.5 meters west of Obock (Abrard, 1942, p. 27, as crocea La- marck). EAST INDIES: Upper Miocene, Tjilanang beds, Java (K. Martin, 1880, p. 119, pi. 19, fig. 4, as T. rudis). MARIANAS ISLANDS: Pleistocene, Tanapag limestone, Saipan; Pliocene-Pleistocene, MARIANAS limestone, Guam (both U. S. Geol. Survey Coll.). PALAU ISLANDS: Up- per Tertiary, Angaur Id. (U. S. Geol. Survey coll.). TONGA: Pleistocene, Nukualofa, Tongatabu (Ostergaard, 1935; BPBM). Tridacna crocea Lamarck, 1819 (Pis. 267, 292) Range — Western coast of the Malay Peninsula eastward to Micronesia. Remarks — In its natural habitat, T. crocea ( the Crocus or Saffron-colored Giant Clam ) can be rec- ognised by its habit of being completely imprisoned in coral pockets. In order to remove the clam, the coral must be broken away and the stout byssus un- dercut. This species differs, therefore, from the ap- parently closely related T. maxima, which lives in relatively more shallow burrows in coral, but which is also tightly fastened to the substrate by a byssus. Other differences between the two are : the shapes of the adductor-retractor muscle scar complexes; char- acteristics of the interdigitating projections of the dorsal margins of the valves, the differing numbers of riblets on the radial folds, and the relative lengths [62 - 078] April 30, 1965 INDO-PACIFIC MOLLUSCA, vol. 1, no. 6 Tridacna 391 of byssal orifices (see descriptions). Valves of T. crocea are usually quite smooth and obese with de- pressed sculpture and are strongly triangularly ovate in shape; T. maxima, although sometimes nearly scaleless, usually does not have the shell sculpture so reduced and its valves tend to be more triangu- larly elongate in shape. Many of these differences appear to be correlated to some degree with the ecology of the species. Museum records indicate a much more limited range for crocea than for maxima. The latter is found throughout the tropical Indo-Pacific faunal region, but the former seems restricted to the northeastern Indian Ocean and western Pacific. It is probable that the range of crocea as indicated here is nearly correct, despite the fact that it is exceedingly diffi- cult to collect. A brachyuran crab, Xanthasia murigera White ( Pinnotheridae ) , was found as a commensal in the mantle cavity, attached to the ctenidia of T. crocea collected by the writer at Ko Sindarar Nua, Thai- land (identified by Dr. R. B. Manning, USNM). Plate 292. Tridacna ( Chametrachea ) crocea Lamarck, Figs. 1-3. Lectotype of T. crocea (96 mm. length); fig. 1. external view left valve; fig. 2. internal view right valve; fig. 3. ventral view showing byssal orifice (photo courtesy of E. Binder, Museum d’Histoire Naturelle, Geneva). Figs. 4, 5. Specimen from Sak Van, North Borneo; fig. 4. external view right valve; fig. 5. internal view left valve (99 mm. length). Fig. 6. Specimen from Mandidarah Island, North Borneo; dorsal view to show characteristics of interdigitating projections ( 105 mm. length ) ( Smithsonian photos ) . [62 - 079] 392 Chametrachea Joseph Rosewater Tridacnidae Habitat — Lives on reef flats and coral heads in water from a few inches to several fathoms in depth. Usually it is found deeply ensconced in coral pockets, the free margins of its valves nearly flush with the substrate. Description — Shell reaching 150 mm. ( about 5- 7/8 inches) in length; triangularly ovate in outline; angle formed at umbos by margins of hinge and ven- tral border usually less than 150°; moderately to strongly inflated and having an extremely large bys- sal orifice. Valves moderately heavy and thick; color usually grayish white but often tinged with yellow or pinkish orange; often encrusted with organisms and debris near free valve margins, but rubbed clean and nearly smooth ventrally. Surface of valves dull or faintly shiny in cleaned specimens. Primary radial sculpture consisting of from 6-10 broad, rather flat- tened, rib-like folds which become crowded and ob- solete on ventral slope. Of these folds, 4 or 5 are very strong. Secondary radial sculpture consisting of from 6-8 evenly spaced riblets on folds and 3-4 in their narrow interstices, often interrupted on primary folds by concentric sculpture and here having nodular ap- pearance. Concentric sculpture consists of closely spaced, undulate lines of growth which can pro- duce crowded, usually low scales on primary folds. Scales usually restricted to area directly adjacent to dorsal free margins of valves and all sculpture is worn progressively smoother as umbos are ap- proached; specimens entirely encased in coral pock- ets completely lack projecting sculpture and undu- late growth lines are pronounced. Dorsal valve margins undulate, with 4 or 5 bluntly triangular, medially projecting, interdigitating processes which represent extremities of rib interstices. Hinge line short, less than half the shell length. A single oblong cardinal tooth present in each valve; 2 elongate pos- terior lateral teeth present in right valve and one in left. Hinge teeth often somewhat obliterated by ero- sion of valves at umbos and encroachment of liga- ment. Ligament secondarily prosodetic. Umbos di- rected postero-medially. Ventral margin slightly to moderately concave. Edge of byssal orifice with series of 4-9 plicae which are closely spaced near umbos and become more distantly spaced posteri- orly. Ventral slope narrow, rounded or only slightly flattened. Interior of valves porcelaneous, usually tinged with yellow or pinkish orange especially near dorsal margin. Pallial line entire and relatively wide. Posterior adductor muscle scar large, subcircular, adjacent to pallial line dorsally and located posterior to center. Posterior pedal retractor muscle scar usu- ally smaller than half the size of the main scar. Area within pallial line, excluding muscle scars, dull; area of the pallial line, the muscle scars, and the area extending to periphery of shell is shiny. Description of animal: Externally observable de- tails of form and coloring are similar to Tridacna maxima. Measurements (mm.) — length height width 150 104 77 large; Canmahana Bay, Lu- zon, Philippines 106 93 64 average; Pulau Mandidarah, North Borneo 82 66 45 average; Helen Reef, Caroline Islands 38 27 20 small; Port Ciego, Balabac, Philippines Synonymy — [1787 Hippopodes scaphe Meuschen, Museum Geversia- num, p. 430 (part); non-binomial. 1 1798 Tridacnes maxima var. a Roding, Museum Boltenia- num, p. 171, no. 185 (no locality given); refers to Chemnitz, vol. 7, pi. 49, fig. 496. 1801 Tridacna gigas Lamarck, Systeme des Animaux sans Vertebres, p. 117 (no locality given) (part); re- fers to Rumphius, 1741 ed., pi. 43, fig. B; to Gual- tieri, pi. 92, fig. A, and others; 1819, Histoire Naturelle des Animaux sans Vertebres, vol. 6 (1), p. 105 (l’Ocean indien) (part); refers to Favanne, pi. 51, fig. B4, and others. 1819 Tridacna crocea Lamarck, Histoire Naturelle des Animaux sans Vertebres, vol. 6 (1), p. 106 (l’Ocean indien [Philippines, here restricted]); re- fers to Lister, pi. 353, fig. 190, Chemnitz, vol. 7, pi. 49, fig. 496, Tableau Encyclopedique et Meth- odique, pi. 235, fig. 2 and to Gualtieri, pi. 92, fig. A. 1845 Tridacna gigas ‘Lamarck’ Chenu, Illustrations Con- chyliogiques, vol. 2, pi. 1, figs. 1, la.; T. crocea Lamarck, ibid. pi. 4, figs. 2, 2a. 1853 Chametrachea scapha ‘Meuschen’ Morch, Catalogus Conchyliorum Comes de Yoldi, fascicle 2, p. 56 (no locality given); refers to Lister, pi. 353, fig. 190. 1862 Tridacna cumingii Reeve, Conchologia Iconica, vol. 14, Tridacna, pi. 7, fig. 7a [not fig. 7b = T. maxima ] (Philippine Islands) (part). 1862 Tridacna ferruginea Reeve, ibid., pi. 8, fig. 8a, b (no locality given [Philippines here designated]). 903 Tridacna lamarcki Hidalgo, Memorias Real Academia de Ciencias, Madrid, vol. 21, p. 385 (Mindoro, Cebu, Mindanao, Paragua [Palawan] (part); re- fers to Favanne, pi. 51, fig. B4, and others; new name for T. gigas Lamarck, not Linne. Types — Since no holotype was designated by La- marck, a probable syntype from the Lamarck Col- lection, Museum d’Histoire Naturelle, Geneva, is here designated as lectotype of Tridacna crocea (pi. 292, figs. 1-3). In the original description, Lamarck gave 102 mm. as the length of his specimen. The lectotype was found to measure 96 mm. by Dr. E. Binder, who provided the photograph. This speci- [62 - 080] April 30, 1965 INDO-PACIFIC MOLLUSCA, vol. 1, no. 6 Tridacna 393 Plate 293. Geographical distribution of Tridacna (Chame- trachea) crocea Lamarck. men is probably the same one portrayed by Chenu (1845, pi. 4, figs. 2, 2a.) The type locality “l’Ocean inclien” given by Lamarck is here restricted to the Philippines since available records indicate that the range of T. crocea barely extends into the Indian Ocean. The location of the syntypes which formed the basis for Reeve’s description of T. ferruginea is un- known, and this species must be based on his figures, both of which unquestionably are specimens of T. crocea. Of these, the larger specimen (Reeve’s pi. 8, fig. 8a ) is here designated the lectotype of T. fer- ruginea. The type locality for ferruginea is here des- ignated as the Philippines. Records — THAILAND: Ko Sindarar Nua (Chance Is- land); Pulau Tanga, Butang Islands (both International In- dian Ocean Expedition, 1963, USNM); airport beach, Ko Phuket, (R. T. Abbott, F. N. Crider, ANSP) all Andaman Sea; Ko Phangan, Gulf of Siam (USNM). MALAYA: Singapore (USNM). RYUKYU ISLANDS: Okinawa (USNM); northeast coast Iheya Shima (ANSP). PHILIP- PINES: Port Binannga, Subic Bay, Bataan Province; Na- sugbu, Batangas Province; Canmahana Bay (all USNM); Tabaco, both Albay Province; Prieto Diaz, Sorsogon Prov- ince (both ANSP) all Luzon; Polillo Islands (USNM); Tilic, Lubang Island; Ambil Island, both Lubang Islands (both MCZ; USNM); Olango Island, between Cebu and Bohol (ANSP); Cahayagan Island, Samar (USNM); Davao City; Balut Island, Sarangani Islands; Sarangani Bay; Santa Cruz Island; off Zamboanga, all Mindanao (all USNM); Port Caltom, Busuanga, Calamian Group (USNM); Cuyo Island (ANSP); Araceli, Dumaran Island (MCZ); Port Ciego, Balabac. (USNM) all Cuyo Group; Tubigan, Pangu- taran Group; Tataan, Tawi Tawi, both Sulu Archipelago (both USNM). INDONESIA: Pulau Penju, northeast of P. Simalur, South Sumatra (I.O.E., 1963, USNM) Tjiperwagaran. Ban- tam, Java (USNM); Amboina, Moluccas (MCZ). NORTH BORNEO: Pulau Mandidarah; Pulau Tiga, west of Banguey; Pulau Tigabu; Sak Van, 9 miles north of Kudat (all Mary Saul, USNM; ANSP). AUSTRALIA: Torres Strait (MCZ); Brook Islands; Pandora Reef; Challenger Bay, Palm Islands; Lindeman Island, Cumberland Group, all Queensland (all H. A. Pilsbry, ANSP). NEW GUINEA: Mios Woendi, Padaido Group, Schouten-Eilanden; Insoemanai, Wakde- Eilanden; Merauke, all West Irian (all MCZ); near Gama- dodo, Milne Bay, Papua (USNM). SOLOMON ISLANDS: Kieta, Bougainville; Faisi, Shortland Islands; Choiseul Bay, Choiseul; Auki, Malaita (all ANSP). PALAU ISLANDS: Koror (USNM; MCZ). CAROLINE ISLANDS: Helen Reef (ANSP); Tomil Harbor, Yap (USNM). Fossil Records - MARSHALL ISLANDS: Pleistocene (?) Elugelab, Eniwetok Atoll (U. S. Geological Survey). [62 - 081] 394 Chametracliea Joseph Rosewater Tridacnidae An Effective Anesthetic for Giant Clams While carrying out taxonomic studies of Tridacni- dae on Eniwetok Atoll, Marshall Islands, it became necessary to make microscopic observations without any movements on the part of the clam. The most effective anesthetic is propylene phenoxetol. A large globule of the viscous compound was dropped to the bottom of the aquarium in an amount equal to about 1 % of the water volume. The chemical dif- fuses very slowly, so that its immediate effect is not felt by the animal until too late for it to contract. After a period of about four hours, six-inch-long clams are nearly insensitive to vibration or changes in light intensity. They may be examined for several hours under a dissecting microscope, prodded, and looked into without their contracting. By replacing the animals in fresh running seawater, apparently complete recovery is affected. The compound is also effective on certain species of gastropods, including Olividae and Conidae. Propylene phenoxetol is a compact chemical, a fluid pound making about 20 liters or 25 quarts of 1% solution, or producing di- rectly a large number of globules. It is distributed by Nipa Laboratories, Ltd., Roman Wall House, 1 Crutched Friars, London E.C. 3, England, and the Goldschmidt Chemical Corp., 153 Waverly Place, New York 14, N.Y., for about $10.00 per pound. [These occasional blank areas occur between genera and subgenera to permit the insertion of new material and future sections in their proper systematic sequence.] [62 - 082] April 30, 1965 INDO-PACIFIC MOLLUSCA, vol. 1, no. 6 Tridacnidae 395 INDEX TO TRIDACNIDAE NAMES IN VOL. 1, NO. 6 Looseleaf subscribers should place this index at the begin- ning of the family Tridacnidae. The family begins on p. [62-007], In this index, the number following the name i refers to the pagination found at the top of the page in vol. column at the right is the looseleaf pagination. 1, no. 6. The [looseleaf] acuticostata Sowerby, 389 62-077 aegyptiaca Chenu, 380 62-068 andreae Tournouer, 356 62-016 arcinella Roding, 356 62-016 asinus Barbut, 363 62-033 asperella Voigt, 356 62-016 aviculare Lamarck, 356 62-016 Avicularium Gray, 353, pi. 270 62-013 356 62-016 besairiei Collignon, 383 62-071 brassica Bose, 363 62-033 Byssocardium Munier-Chalmas, 353, pi. i 170 62-013 356 62-016 Cerceis Gistl, 361 62-031 Chametrachaea H. and A. Adams, 379 62-067 Chametrachea Morch, 379 62-067 compressa Reeve, 389 62-077 cookiana Iredale, 373 62-049 corallicola Lamy, 389 62-077 crocea Lamarck, 390 62-078 cumingii Reeve, 389 62-077 392 62-080 cymbulare Lamarck, 356 62-016 derasa Roding, 375 62-057 Dinodacna Iredale, 367 62-043 “elongata Humphrey,” 388 62-076 elongatissima Bianconi, 382 62-070 emarginatum Deshayes, 356 62-016 equinus Morch, 363 62-033 ferruginea Reeve, 392 62-080 Flodacna Iredale, 379 62-067 fossor Hedley, 389 62-077 Gataron Berthold, 367 62-043 gigantea Perry, 373 62-049 gigantus Bournon, 373 62-049 gigas Lamarck, 388 62-076 gigas Linne, 369 62-045 glabra Link, 377 [looseleaf] 62-059 Goniocardium Vasseur, 353, pi. 270 62-013 356 62-016 gunteri Mansfield, 365 62-039 heberti Vasseur, 356 62-016 “Hippopigenus Renier,” 361 62-031 “Hippopodes Meuschen,” 361 62-031 Hippopus Lamarck, 361 62-031 hippopus Linne, 361 62-031 hyppopus Bournon, 363 62-033 imbricata Link, 373 62-049 382 62-070 388 62-076 imbricata Roding, 388 62-076 “indica Buonnani,” 356 62-016 Key to the Tridacnidae, 359 62-023 lamarcki Hidalgo, 373 62-049 382 62-070 389 62-077 392 62-080 lanceolata Sowerby, 389 62-077 latissima Bianconi, 356 62-016 loczyi Kutassy, 379 62-067 maculatus Lamarck, 363 62-033 matleyi Cox, 356 62-016 maxima Roding, 384 62-072 mbalavuana Ladd, 379 62-067 media Pusch, 384 62-072 “multifolia Humphrey,” 356 62-016 mutica Lamarck, 373 62-049 388 62-076 noae Roding, 388 62-076 obesa Sowerby, 377 62-059 Pelvis Megerle, 361 62-031 pennicornis Sherborn, 356 62-016 Persikima Iredale, 375 62-057 punctulosa Sherborn, 356 62-016 pustulosa Lamarck, 356 62-016 rachitis Deshayes, 356 62-016 reevei Hidalgo, 389 62-077 rudis Reeve, 388 62-076 [62 - 003] 396 Index Joseph Rosewater Tridacnidae [looseleaf] [looseleaf] Sawkinsia Cox, 356 62-016 “Tridacnigenus Renier,” 367 62-043 “scaphe Meuschen,” 382 62-070 Tridacnodites Kruger, 367 62-043 392 62-080 Tridacnoides Bronn, 367 62-043 “scutra Meuschen,” 388 62-076 Tridacnus Paetel, 367 62-043 Sepidacna Iredale, 379 62-067 troughtoni Iredale, 389 62-077 serrifera Lamarck, 377 62-059 squamosa Lamarck, 380 62-068 umbricata Sherborn, 389 62-077 squamoza Chenu, 382 62-070 ungula Roding, 363 62-033 “ursina Humphrey,” 363 62-033 trechmanni Cox, 356 62-016 Tridachna Neave, 367 62-043 Vulgodacna Iredale, 379 62-067 Tridacna Bruguiere, 367 62-043 Tridacne Link, 367 62-043 whitleyi Iredale, 377 62-059 Tridacnes Roding, 367 62-043 wolfarti Chenu, 367 62-043 INDEX TO COMMENSAL CRUSTACEA IN TRIDACNIDAE Anchistus cf demani Kemp, 381 62-069 381 62-069 386 62-074 386 62-074 Anchistus maculatus ( Stimpson ) , 350 62-010 Anchistus miersi (DeMan), 381 62-069 Palaemonidae, 350 62-010 Anchistus mirabilis (Pesta), 369 62-045 Paranchistus biunguiculatus (Borr.), 370 62-046 Anchistus spinuliferus ( Miers ) , 350 62-010 Pontoniinae, 350 62-010 Commensalism, 350 62-010 Xanthasia murigera White, 381 62-069 Conchodytes tridacnae Peters, 369 62-045 391 62-079 Published by The Department of Mollusks Academy of Natural Sciences of Philadelphia 19th and the Parkway Philadelphia 3, Pennsylvania [62 - 004] April 30, 1965 INDO-PACIFIC MOLLUSCA, vol. 1, no. 6 Strombus 397 STROMBUS (TRICORNIS) OLDI NEW SPECIES by William K. Emerson Chairman and Associate Curator Department of Living Invertebrates American Museum of Natural History Strombus oldi new species (PI. 294, figs. 1-4) Range — Known only from the Somalian coast of East Africa. Remarks — Although the fauna of the Somalian coast is not well known, it seems remarkable that this very distinctive species has not been previously dis- covered. It was first brought to light by my associate, Mr. William E. Old, Jr., who was curating a small collection of marine mollusks that was labeled “50 miles north to 60 miles south of Obbia, Italian Somaliland [Republic of Somalia], East Africa, No- vember 21, 1956.” Recently, other specimens have been found near Mogadishu [Mogadiscio]. One of these was kindly forwarded by Mrs. Orville R. Davis for examination. Of the known species of Strombus, the shell of S. oldi most closely resembles Strombus (Tricornis) sinuatus Humphrey, 1786, but it lacks the 3 or 4 tongue-like blades on the upper end of the outer lip and a thickened outer lip. In spire angle and nodulation and in the spiral ornamentation of the shell, S. oldi closely resembles Lambis ( Millepes ) digitata (Perry, 1811) from the Indian Ocean, but lacks the strong labial and parietal wall ornamenta- tion of that Lambis. This new species is character- ized by the wing-like projection on the upper end of the outer lip, by the relatively high, noduled spire, by the strong, rounded cords on the back of the outer lip, by the dark-brown blotches within the smoothish aperture, and by the rich chestnut coloring of the outer shell. Strombus solitaris Perry, 1811 (Arcana, London, vol. 1, part 2, signature Dd1, [pi. 52] is a poorly il- lustrated and inadequately described species from “Africa and the East Indies,” and, although some- what resembling S. oldi, is undoubtedly a synonym of the Caribbean S. gallus Linne, 1758, by virtue of the lower spire, the larger and fewer shoulder knobs and the slender, narrow wing-like projection on the upper end of the outer lip. Perry’s locality is un- doubtedly erroneous. I take pleasure in naming this new species in honor Plate 294. Strombus oldi Emerson ( new species). Figs. 1, paratype, 109 mm. in length, AMNH no. 97346. Both from 2, holotype, 112 mm. in length, ANSP no. 299730, Figs, 3, 4, Republic of Somalia, northwest Indian Ocean. [09 - 881] 398 Tricornis William K. Emerson Strombidae of Air. Old, who immediately recognized the signifi- cance of his discovery. Description — Adult shell 95 to 112 mm. (about 3% to 4& inches ) in length, solid, moderately heavy, with a dark-brown stain within the aperture and with 8 to 10 strong, nodular raised ribs between the anal and siphonal canals on the back of the outer lip. Color of shell is whitish with a pattern of narrow axial bands of yellowish brown on ventral portion of the body whorl and with an overlay of yellowish brown interrupted by the ribs and nodules on the dorsum. Whorls number 7 to 10; nuclear whorls are worn off. Post-nuclear whorls are ornamented by numerous, low, whitish knobs on the periphery and are colored with axial blotches between the knobs and on the concave portions of the whorls. The knobs are more prominent on the later whorls and are larg- est on the body whorl, which is also ornamented on the dorsum with numerous rows of smaller spiral ridges, two of which possess rounded beads. Be- tween the spiral ridges are fine spiral threads. Pari- etal wall is weakly glazed. Columella is stained with dark-brown and is smooth except for three small plications at the upper end of the columella, adja- cent to the anal canal. Interior of the outer lip is thickened in the region where the previous lip ter- minated. Siphonal canal is short, relatively broad and slightly twisted to the right. Stromboid notch is moderately deep and is slightly flaring to the right. Operculum, periostracum and soft parts are unknown. Measurements (mm.) — length width no. whorls 112 73 10 + holotype from Mogadiscio 109 72 8 + paratype, AMNH no. 97346 105 47 8 + paratype, AMNM no. 97346b 95 65 7 + paratype, AMNM no. 97346a Types and Records — The type locality is “Moga- discio, Republic of Somalia.” It was obtained by M. Ramazzotti, and given to the Academy of Natural Sciences of Philadelphia (no. 299730) by Mrs. Orville Davis. Paratypes, all from the vicinity of Obbia, are in the American Museum of Natural History (New York); AMNH nos. 97346, 97346a, and 97346b (im- mature ) . Published by The Department of Mollusks Academy of Natural Sciences of Philadelphia 19th and the Parkway Philadelphia 3, Pennsylvania [09 - 882] April 30, 1965 INDO-PACIFIC MOLLUSCA, vol. 1, no. 6 Strombus 399 STROMBUS (EUPROTOMUS) LISTERI T. GRAY - NEW RECORDS; ANATOMY by Takashi Okutani Tokai Regional Fisheries Research Laboratory, Tokyo, Japan Since the 1960 treatment of Strombus listen T. Gray by Abbott in Indo-Pacific Mollusca, vol. 1, no. 2, p. 115 [looseleaf p. 09-967], new records and new anatomical information have come to hand. The known range of this species is now extended to the Gulf of Oman, and, on the basis of the penis and radulae, listen is removed from the subgenus Dox- ander, and placed in the subgenus Euprotomus. I am indebted to Mr. S. Shimura, who was in charge of the Japanese Fisheries Agency research vessel, Shoyo-Maru, in 1958, for the opportunity of exam- ining preserved soft parts, and to Dr. R. Tucker Abbott and the late Mr. Richard W. Foster for rec- ords obtained from the cruises of the United States research vessel Anton Bruun. Strombus listeri T. Gray, 1852 (Pis. 295, 296) (also see p. 116 [09-968]) Range — Gulf of Oman, northwest Indian Ocean, and the Bay of Bengal. Description — Adult shell 100 to 150 mm. ( about 4 to 6 inches ) in length, rather thin but strong, with a high spire; nuclear whorls 3, smooth, convex, trans- lucent lavender; post-nuclear whorls 10 in number. First 6 whorls with numerous axial ribs all over, and fine spiral cords in between; these give the shell a cancellated appearance. The lower whorls rather angulated at the shoulder and smooth except for several spiral striae near the anterior canal area. Outer lip moderately quadrangularly expanded and with a nearly straight margin which is somewhat thickened; the inner surface smooth, polished, enamel-white with golden tint. There is a peculiar thickened part (ridge) on the central area of the Plate 295. Gross anatomy of Strombus listeri from the Bay of Bengal (shell 102 mm. in length). Fig. 1, operculum, length 22 mm. Fig. 2, external features: ct, cephalic tentacle; fa, fleshy appendage on posterior comer of tire mantle; m, mantle; op, operculum; p, penis; pr. proboscis; ps, pedal slit at anterior end of foot; sp. siphon. Fig. 3, protoconch of shell. Fig. 4, internal anatomy, an, anus; au, auricle of heart; eg, cerebral ganglion; cm, columellar muscle; g, gills; hg, hypo- branchial gland; in, intestine; k, kidney; Iv, liver; m, mantle; mo, mouth; oe, oesophagus; os, osphradium; p, penis; re, receptacle seminis; ro, renal opening; rs, radular sheath; rt, rectum; sp, siphon; st, stomach; ts, testes; vd, vas deferens; vt, ventricle of heart. Fig. 5, radulae showing rhachidian (R), marginal (M) and lateral teeth (L). Fig. 6, penis. [10-001] 400 Euprotomus Takashi Okutani S trombidae outer lip. Posterior canal very deep, so that the pos- terior corner of the outer lip protrudes very promi- nently; anterior siphonal canal well-developed; stromboid notch very deep, U-shaped. The external surface brown and whitish overlaid with chestnut undulating axial stripes and speckled bands; perio- stracum yellowish, very thin. Operculum stromboid, strongly curved, with about 17 serrations on the margin; arch length 22 mm.; width, 5 mm. Measurements (mm.) — length width 150.0 56.8 holotype of mirahilis Sowerby, Brit. Mus. 103.5 40.0 Bay of Bengal ( Shoyo-Maru coll. ) 102.0 41.0 Bay of Bengal ( “ coll.) Radula — Radular ribbon delicate, 5.5 mm. in length, 1.8 mm. in width with about 50 rows of teeth. Radula of taenioglossate type; rhachidian tooth trapezoidal in shape, with 8 to 10 obtuse cusps at the tip; lateral oblong subovate in outline, curved at the tip with 6 to 8 obtuse cusps, of which the innermost one is the largest; two marginals sickle-shaped with inconspicuous dentations near the distal extremities; formula 2-1-C-1-2. General observations on the soft parts — Head, foot, mantle, proboscis, siphon and cephalic tenta- cles uniformly fleshy in color; snout long; eyes black, pedunculated; cephalic tentacles long, tapered; an- terior end of the foot rather cylindrical with a nar- row sole; columellar muscle small. There is a short fleshy appendage on the posterior corner of the man- tle. Alimentary canal: proboscis cylindrical, interior of it longitudinally grooved; eosophagus thin-walled slender tube; salivary gland indistinct in the present specimen; radula sheath short, situated on the ven- tral side of the buccal mass; stomach apparently con- sisted of two parts and situated between anterior and posterior lobes of the liver; intestine thin-walled; rectum very thick in diameter, filled with debris-like faeces; kidney medium in size. Genital organ: penis large, keeled, with two pointed tips. Pallial organ: osphradium slender, about 15 mm. in length, situ- ated well within the mantle cavity; gill very thin and broad; hypobranchial gland irregularly ovate in outline. Records — ARABIA: off Khawr, Trucial Oman, Gulf of Oman, 40 fms., 26° 35'N. Lat.; 56° 25'E. Long. Station 259A Anton Bruun, Dec. 1, 1963. INDIA: 18 mi. S.E. of Viza- gapatnam, 79 meters. Station 90, Anton Bruun, April 28, 1963. BURMA: 17 mi. S.S.E. of Akyab, 55 meters. Station 49, Anton Bruun, April 5, 1963; 50 mi. S.W. of Irrawaddy River, Preparis North Channel, 53 meters, Gray mud. Station 43, Anton Bruun, April 1, 1963. THAILAND: 65 mi. N.N.W. of Phuket Id., Andaman Sea. Station 20, Anton Bruun, Mar. 23, 1963. Plate 296. Strombus listen T. Gray. Figs. 1 and 2, from the from off Khawr, Gulf of Oman, 40 fms.; photo by Bruce Bay of Bengal (103 mm. in length). Fig. 3, live specimen Rogers. [10-002] April 30, 1965 INDO-PACIFIC MOLLUSCA, vol. 1, no. 6 S trombus 401 MIOCENE STROMBUS (DOLOMENA) FROM INDIA by R. Tucker Abbott Pilsbry Chair of Malacology Academy of Natural Sciences of Philadelphia In his 1962 report on The Miocene Mollusca from Quilon, Kerala (India), A. K. Dey described three new fossil species of Strombus which we are includ- ing in this supplement to vol. 1, no. 2 of Indo-Pacific Mollusca. All three belong to the subgenus Dolo- mena Iredale, 1931, and are closely related to pre- viously described Tertiary species from India and Indonesia. A fourth species, “Strombus ? sp.” ( 1962, p. 66, pi. 6, figs. 4, 8) appears to be an immature specimen of a species in the subgenus Laevistrombus Kira, 1955. Strombus sedanensis subspecies daviesi Dey, 1962 (PI. 297, figs. 1, 2) Range — Known only from the Miocene Quilon beds of Kerala, S. W. India. Remarks — S. daviesi closely resembles the Mio- cene S. sedanensis K. Martin, 1899 of Java and West Pakistan, [see Indo-Pacific Mollusca, vol. 1, no. 2, looseleaf p. 09-940], and I believe should be con- sidered a subspecies of it. In daviesi the outer lip is less developed at the posterior end, hence giving the spire an appearance of being proportionately higher. Plate 297. Figs. 1, 2, holotype of Strombus daviesi Dey; Miocene of Kerala, India. 45 mm. Figs. 3, 4 holotype of This character is quite variable in some Recent as well as fossil S trombus. Vredenburg ( 1928, p. 313 ) reported S. sedanensis K. Martin from the Miocene Gaj beds, near Karachi, W est Pakistan, and his speci- mens may be daviesi Dey. This subspecies was named in honor of the British paleontologist, A. Morley Davies. Description — Adult shell 45 mm. ( 1 %, inches in length); spire moderately high; whorls about 9 in number, of which the first 3 form the protoconch; whorls shouldered; suture bordered anteriorly by a narrow cord. The ornamentation of the earlier whorls consists of close-set axial ribs and spiral threads; the ribs change to swollen nodes or blunt spines on the shoulder of the later whorls; aperture long, wide, deeply emarginate at its base; siphonal canal re- curved; inner lip and parietal wall coated with a thin, light-brown deposit or callus; outer lip thick, expanded, having a shallow, wide notch at its pos- terior end, which does not reach the suture of the preceding whorl, and a deep stromboid notch near the base; on the interior of the outer lip is a ridge which corresponds to the wrinkles on the exterior ( from A. K. Dey, 1962, p. 64 ) . Measurements (mm.) — length width no. whorls 45 29 9 holotype Strombus quilonensis Dey; Miocene of Kerala, India. 53 mm. (from Dey, 1962, pi. 6). [09 - 945] 402 Dolomena R. T. Abbott S trombidae Synonymy — 1962 Strombus daviesi Dey, Palaeontologia Indica, Memoirs Geol. Survey India, new series, vol. 36, p. 64 (pi. 6, figs. 10, 15; pi. 9, fig. 1). Types and records — The type locality is the Quilon limestone bed at Padappakara, Kerala State, southwest India; Miocene. The holotype and para- types are in the Geological Survey of India collec- tions, nos. 16437 and 16438. It is known only from the type locality. Strombus preoccupatus subspecies quilonensis Dey, 1962 (PI. 297, figs. 3, 4) Range — Known only from the Miocene Quilon beds of Kerala, S. W. India. Remarks — I can find few differences between Dey’s quilonensis and S. preoccupatus Finlay, 1927 (for illustrations see Indo-Pacific Mollusca, vol. 1, no. 2, p. 123, looseleaf page 09-979). The latter is known from the Miocene of Java and Borneo. The Indian subspecies seems to have stronger spiral threads and the spines on the lower part of the body whorl ap- pear to be less strongly developed. I believe that Dey’s Strombus cossmanni, described in the same paper and from the same locality as that of quilon- ensis, merely represents juvenile specimens of the latter. Dey believed that his cossmanni was the same as a young specimen from the Pliocene of Karikal figured in 1903 by Cossmann (Journ. de Conchyl., vol. 51, pi. 6, figs. 29, 30). However, I believe that Cossmann’s specimen was probably a young form of variabilis Swainson. I am transferring Strombus preoccupatus Finlay from the subgenus Lentigo to the subgenus Dolo- mena. Description — Adult shell about 53 mm. in length, stout, spinose, and with a rather high, pointed spire; suture bordered anteriorly by a cord, most conspic- uous on the early whorls; ornamentation of early whorls consisting of closely-spaced axial ribs with occasional varices and spiral threads. As the shell grows the axials are reduced to nodes and finally become spines on the well-marked shoulder-angle of the body-whorl. Below the shoulder-angle there are 4 main spiral ridges of which the upper 3 are no- dose, with irregularly distributed spiral threads in their interspaces; siphonal canal slightly recurved; aperture long, wide, deeply emarginate at base and extending posteriorly beyond the suture of the pre- ceding whorl; inner lip and parietal wall coated with a deposit or callus; outer lip having internally a strong crenate cord corresponding with a swollen wrinkle on the exterior (from A. K. Dey, 1962, p. 64). Measurements (mm.) — length width no. whorls 53 36 — holotype of quilonensis 34.5 17 — holotype of cossmanni Synonymy — 1962 S trombus quilonensis Dey, Palaeontologia Indica, Memoirs Geol. Survey India, new series, vol. 36, p. 64, pi. 6, figs. 1, 5. 1962 Strombus cossmanni Dey, loc. cit., p. 65, pi. 6, figs. 16, 17. Types and Records — The type locality for both quilonensis and cossmanni is the Quilon limestone bed at Padappakara, Kerala State, southwest India; Miocene. The types are in the Geological Survey of India collections: quilonensis (no. 16439); coss- manni (no. 16440). Published by The Department of Mollusks Academy of Natural Sciences of Philadelphia 19th and the Parkway Philadelphia 3, Pennsylvania [09 - 946] April 30, 1965 INDO-PACIFIC MOLLUSCA, vol. 1, no. 6 [replacing vol. 1, no. 5, pp. 317, 318; see stars] Echinoturris 403 The Xenuroturris Group Relationship appears to have existed within a group of Miocene to Recent Austro-Neozelanic genera which exhibit either alliance to or analogy with the wide-ranging Indo-Pacific Pliocene to Re- cent Xenuroturris. Of the five genera in this group only Xenuroturris has a poly gyrate protoconch; the others have a blunt cylindrical-sided, paucispiral protoconch, the tip smooth, followed by regular or brephic axials. The New Zealand Miocene Echinoturris has a relatively short canal and distinctive sculpture of a bicarinate series of sparse prickly nodules. The V- shaped sinus is on the upper carina. The southern Australian Miocene to Pliocene Veruturris has a relatively short canal compared with Turns but not truncated to the extent exhib- ited by Xenuroturris. A third group, for which a new subgenus Cin- guliturris is provided, is represented by the Austral- ian Miocene (Balcombian) species Asthenotoma tatei Cossmann, which is Xenuroturris-} ike with its truncated canal but has a paucispiral protoconch and the sinus rib as a simple spiral cord, not dupli- cate or margined in any way. Finally, the New South Wales Recent species, Xenuroturris corona Laseron, for which a new genus Viridoturris is provided, is another member of the paucispiral-apiced group, standing nearest to Xenuroturris in its truncated anterior end and de- fined sinus rib, but lacking the characteristic poly- gyrate, axially costate protoconch of that genus. If the above association of genera is a correct assumption, it would then appear that these pre- sumed offshoots have arisen independently at dif- ferent times in two areas marginal to the present tropical Indo-Pacific range of Xenuroturris typical. Genus Echinoturris Powell, 1942 (PI. 248) Type: “Tunis” finlayi Powell, 1935 The genus is so far known only by one species, the type locality for which is volcanic tuffs of Al- tonian Miocene age from Motutara, west coast, Auckland, New Zealand. The genus is characterised by small size, 8 to 10 mm., moderately long canal, blunt, round-topped, cylindrical-sided, smooth protoconch of two whorls, ending with a few closely spaced thin axials. Plate 248. Echinoturris finlayi (Powell). Holotype. Alton- ian Miocene of Motutara, Auckland, New Zealand. Key to the Xenuroturris group Protoconch polygyrate Sinus rib margined Canal truncated Xenuroturris Protoconch paucispiral Sinus rib margined or defined Canal truncated Viridoturris nov. Canal moderately long Veruturris Sinus rib not margined Canal short Sculpture cingulate Cinguliturris Canal moderately long Sculpture bicarinate-spinose Echinoturris [22 - 947a] 404 Veruturris Turridae [replacement page] A. W. B. Powell The most distinctive features are the bicarinate series of sparse prickly nodules and the rather deep V-shaped sinus situated on the upper carina, which like the lower carina is a simple rounded cord, not margined. The relationship, if any, with Xenuro- turris is certainly not close. Synonymy — 1942 Echinoturris Powell, Bull. 2, Auckland Inst, and Mus., p. 50. Type by monotypy: “Tunis” finlayi Powell, 1935 (Awamoan, Middle Miocene, New Zealand). Genus Veruturris Powell, 1944 Type: Xenuroturris (Veruturris) quadricarinatus Powell, 1944 This is a Miocene-Pliocene group from South Australia and Victoria, characterised by moderate size, 14 to 50 mm., a tall spire but only moderately long canal; paucispiral blunt protoconch of 2 to 2/2 smooth whorls followed by a half-whorl or so of brephic axials. Sinus deep, broadly V-shaped, on sinus rib situated above the middle of whorl height. This rib may be a single cord as in the type species and as in tomopleuroides, or two almost fused cords. The genus Xenuroturris differs in having a polygyrate protoconch and a very short anterior canal. The species of this genus and other turrid repre- sentatives of the Australian Tertiary are included since they are members of past faunas climatically analogous to the present tropical Indo-Pacific fauna. Synonymy — 1944 Veruturris Powell, Records Auckland Inst, and Mus. 3(1), p. 9 (subgenus). Type by original designa- tion : Xenuroturris (V.) quadricarinatus Powell, 1944. Veruturris bisculpta (Powell, 1944) Range — Abattoirs (type locality) and Wey- mouth Bores, Adelaide, Adelaidean, Lower Mid- Pliocene. Remarks — Upper half of spire whorls sculptured with about 17 vertical fold-like axials, crossed by three spiral cords which are rendered nodulose by the axials; lower half of each whorl with 2 or 3 heavy, closely spaced, plain spirals. Sinus area a deep narrow groove between two of the cords at about two-thirds whorl height. Measurements (mm.) — height width 26.7 6.5 13.9 4.5 holotype Plate 249. Veruturris subconcava (Harris). Miocene, Jan- jukian, of Victoria, Australia (from Harris, 1897, pi. 3, figs. 2a, b). Synonymy — 1944 Xenuroturris (Veruturris) bisculptus Powell, Rec. Auck. Inst. Mus. 3(1), p. 11, pi. 1, fig. 4. 1958 Xenuroturris (Veruturris) bisculptus Powell, Ludbrook, Trans. Royal Soc. South Australia 81, p. 85. Types — The holotype is in the Auckland Mu- seum (Finlay collection). Veruturris cochleata (Powell, 1944) (PI. 250) Range — Balcombian, Miocene of Muddy Creek, lower beds, Victoria (type locality); Altona shaft and Clifton Beach, Victoria. Remarks — Spire whorls with strong narrowly rounded cords, one linear-spaced pair submargin- ing the suture, then a second pair, forming the sinus rib, situated at three-quarters whorl height, followed by five or six smooth, rounded, spiral cords. Sinus narrowly V-shaped. Measurements (mm.) — height width 52 14 Synonymy — 1944 Xenuroturris (Veruturris) cochleatus Powell, Rec. Auck. Inst. Mus. 3(1), p. 10, pi. 7, fig. 11. Types — The holotype is in the Auckland Mu- seum (Finlay collection). Veruturris quadricarinata (Powell, 1944) Range — Balcombian, Miocene of Muddy Creek, lower beds, Victoria (type locality), and Altona shaft, Victoria. Remarks — Spire whorls with evenly spaced, strong, rounded cords, middle pair stronger than the other two. Sinus broadly V-shaped, situated on upper of the middle pair of cords. [22 - 948a] April 30, 1965 Lophiotoma 405 INDO-PACIFIC MOLLUSCA, vol. 1, no. 6 [replacing vol. 1, no. 5, pp. 309, 310; see stars] Plate 239. Geographical distribution of Lophiotoma (Loph- iotoma) abbreviata (Reeve) and its subspecies ustulata (Reeve) and lifuensis (Sowerby). unknown locality but similar shells are known from the two Indian Ocean localities mentioned above; not from the Pacific range of the typical species, however. The type specimen is badly faded but the charac- teristic colour pattern of the typical species, par- ticularly the subsutural maculations, is still dis- cernable. The sculpture, however, is much weaker than is normal for abbreviata, the bicingulate peri- pheral keel being more like that of acuta; the an- terior canal, however, is truncated. Until more is known of this form it had better be admitted as a probable subspecies of abbreviata. Measurements (mm.) — height width 31 12 holotype Synonymy — 1846 Pleurotoma ustulata Reeve, Conch. Iconica 1, pi. 40, fig. 369 (no locality). 1884 Pleurotoma ustulata Reeve, Tryon, Manual of Conch. 6, p. 167, “Mauritius.” Types — The holotype is in the British Museum (Natural History). Records — INDIAN OCEAN : Mauritius; Andaman Ids. (USNM). Lophiotoma ruthveniana (Melvill, 1923) (PI. 240, fig. 1) Range — Mauritius. Remarks — This handsome shell appears closely allied to abbreviata and is distinguished from that species mainly by the overall strongly tessellated colour pattern. In abbreviata the larger maculations are confined to the subsutural fold but in ruthven- iana the peripheral carina bears heavier macula- tions. In the absence of well-preserved apical whorls in the four examples seen, including the holotype, the species is only provisionally placed in Lophiotoma. Description ( from Melvill ) — “Shell fusiform, thick; whorls, especially the upper, somewhat com- pressed, being ten in number, inclusive of the two apical. Colour bright chestnut brown, with squar- rose, fairly regular, white tessellations on the spiral [22 - 919a] 406 Lophiotoma T urridae [replacement page] A. W. B. Powell carinae. These revolving keels appertain through- out—one, in particular, central, and subdivided by a shallow sulcus; the lesser tornate keels increase nu- merically in each of the lower whorls, till, on the body whorl, they total five or six, all beautifully variegated with white and chestnut alternately, as mentioned above. Mouth ovate-oblong, canal wide, abbreviate, sinus well developed, wide and deep, columellar margin fairly straight. Long. 41.5, lat. 14 mm. “A handsome species, standing somewhat alone, and conspicuous for its bright coloration and tes- sellated carinal ornamentation.” Synonymy — 1923 Tunis rutlweniana Melvill, Proc. Mai. Soc. 15, p. 162, pi. 4, fig. 2. Types — The type locality is Mauritius. The holo- type is in the British Museum (Natural History), London. Lophiotoma brevicaudata ( Reeve, 1843 ) ★ (PI. 240, fig. 2) Range — Gulf of Suez, New Guinea, the Philip- pines and New Caledonia. Remarks — Three syntypes of Reeve’s species are in the British Museum and there is little to add to his original description: “Shell shortly fusiform, solid, yellowish, brown at the base and apex; whorls convex, encircled with a single keel round the upper portion and a double keel round the lower; last whorl encircled with single and double keels alternately down to the base, lip simple and acute, sinus large; aperture small, short, canal rather short. Hab. Island of Ticao (found on reefs), Cuming.” It is an easily recognized little shell due to its regular smooth spirals and uniform yellowish brown colour, except for the brown staining of the anterior end. The apical whorls are eroded in all three syn- types. Tryon (1884, p. 169) made this species a syno- nym of fascialis Lamarck = polytropa Helbling, but they are certainly not con-specific. Apart from the type material and imperfect spec- imens from New Guinea and New Caladonia, the best preserved example seen is one from the Gulf of Suez (ex McAndrew coll., Brit. Mus. ) in the Museum of Comparative Zoology, Harvard, and a description of the specimen follows: Description ( Red Sea specimen ) — Adult shell 22 mm. (slightly under 1 inch) in height, fusiform, with a tall spire, but with a relatively short anter- ior canal. Whorls 7, exclusive of the protoconch, sufficient of which remains in one specimen to indi- cate that it is narrowly conic of several whorls. Spire 1.3 times the height of the aperture plus canal. Adult sculpture of narrow, sharp cords and threads, one cord on a moderately inflated subsutural fold, five threads over a deeply concave shoulder. There is a double-corded peripheral keel, enclosing a con- cave interspace with a single thread in the middle. There is one primary cord and two threads between the peripheral carina and the lower suture. Three cords, stronger than the rest, are on the upper part of the base, with each interspace having three threads, followed by an alternation of cords and threads over the neck, and finally closely-spaced threads over the anterior canal. Sinus deep, U- shaped, on the peripheral carina. Colour pale yel- lowish buff, with all the spirals continuously lined in light-brown and with a dark-brown staining on the anterior canal. Measurements (mm.) — height width 24.0 8.3 lectotype 22.2 7.6 Gulf of Suez Synonymy — 1843 Pleurotoma brevicaudata Reeve, Conch. Iconica 1, pi. 15, fig. 126. 1884 Pleurotoma fascialis Lamarck, Tryon, Manual of Conch. 6, p. 169 ( non Lamarck, 1822 ) . Types — British Museum (Nat. Hist.), three syn- types, lectotype, 24 x 8.3 mm., here selected. The type locality is Ticao Id., Philippines, Hugh Cum- ing, collector. Records — PHILIPPINES: Ticao Id. (type locality); Cal- apan, Mindoro Id. (MCZ). NEW CALEDONIA: (Aus- tralian Mus.). WEST NEW GUINEA: Geelvink Bay, % mi. S. of Maroepi, Ambai, Japen Id., 14-25 fms., blue mud, shell and coral (ANSP). RED SEA: Gulf of Suez (MCZ). [22 - 920a] April 30, 1965 INDO-PACIFIC MOLLUSCA, vol. 1, no. 6 Issues and Changes 407 ISSUES AND CHANGES Looseleaf subscribers should put this sheet in the front of their binder just before the Introduction [looseleaf p. 00-051]. Note that vol. 1, no. 6, con- tains additions and changes to previously published monographs. Remove staples from all issues and place the new sheets where they belong by follow- ing the looseleaf numbering at the bottom of the page. Throw away any blank, brown paper sheets. Replaced pages may be discarded or saved in the back of the binder behind the guide tab, “Replaced pages.” Notice that the index pages are numbered so that they will immediately follow their appro- priate tab. Volume 1 has not been completed. Do not bind as yet. Do not send additional payments unless you are billed. List of Guide Tabs Guide tabs are automatically sent to all those who receive the journal looseleaf. Those receiving the folded pamphlet style for future binding may, if they wish, order tabs at the rate of 10 cents (U.S.) each. The present set of ten costs $1.00. List of Issues 00-005 Guide to Contents 00-100 Phasianellidae 04-000 Strombidae 09-650 ’Cassidae 12-400 Vasidae 20-400 Turridae 22-500 Pinnidae 53-500 Tridacnidae 62-000 Replaced Pages 95-000 List of Issues Date of First page of Volume 1 Subject Author Publication no. pp. looseleaf no. 1, pp. 1- 14 Introduction Editors Mar. 31, 1959 14 00-003 15- 32 Vasidae R. T. Abbott 18 20-403 2 33-146 Strombus R. T. Abbott Nov. 23, 1960 114 09-831 3’ 147-174 Lambis R. T. Abbott Sept. 28, 1961 38 10-051 4, 175-226 Pinnidae J. Rosewater 52 53-501 5, 227-346 Turrinae A. W. R. Powell Mar. 31, 1964 120 22-661 6, 347-396 Tridacnidae J. Rosewater Apr. 30, 1965 50 62-003 397-398 Strombus oldi W. K. Emerson 2 09-881 399-400 Strombus listeri T. Okutani 2 10-001 401-402 Miocene Indian Strombus R. T. Abbott 2 09-945 * *403-406 Turrinae ( replacement ) A. W. R. Powell “ 4 22-947 407-408 Issues and Changes Editors “ 2 00-005 * Formerly spelled “Cassididae,” but there is an earlier and currently used family by that name in Insecta. * * No charge is made for these four pages because they contain editorial corrections. [00 - 005] Editors Issues Although primarily a scientific journal, INDO-PACIFIC MOLLUSCA is attrac- tively designed and richly illustrated in full color to serve the serious amateur inter- ested in marine shells. Subscription The subscription rate is U.S. per page. Numbers are issued at irregular intervals, as research is completed. It is hoped that a yearly average of a hundred pages will be maintained. A list of back numbers, their prices and an order form may be obtained from the publisher. The Journal is not avail- able on a library exchange basis, and dis- counts to magazine agencies are not made. The blue and gold-embossed, expandi- ble post binder shown here will hold 600 pages. It will protect copies waiting for binding or serve as a permanent, heavy- duty holder. Cost $4.25 each (foreign, add 50^ for postage and packing). Published by The Department of Mollusks Academy of Natural Sciences of Philadelphia 19th and the Parkway Philadelphia 3, Pennsylvania [00 - 006] May 15, 1967 INDO-PACIFIC MOLLUSCA, vol. 1, no. 7 Turrinae 409 <401 r5A faoLL- / THE FAMILY TURRIDAE IN THE INDO-PACIFIC Part la. The subfamily Turrinae concluded by A. W. B. Powell Auckland Institute and Museum Auckland, New Zealand In this section, the Turrinae of the Indo-Pacific is concluded by the inclusion of five genera of rather uncertain relationships which, upon the available evidence, seem to have more in common with the Turrinae than with any of the other sub- families. New Hawaiian Gemmula and Xenuroturris are also included. Below are listed the recognized taxa for the re- mainder of the Turrinae. The earlier list appeared in volume 1, no. 5, on pages 235-237 [pp. 22-669 to 22-671], As before, 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 dagger ( f ) . When a species occurs both Re- cent and fossil, the dagger occurs after the name. Included in the list are southern Australian and New Zealand Recent and Tertiary Turrinae. Such species are not excluded, since they have had an Indo-Pacific origin during past periods of the Ter- tiary when warm waters extended farther south than they do at present. List of Recognized Taxa in the Turrinae (concluded) Subfamily TURRINAE Heteroturris Powell, new genus sola Powell, new species. Type Turridrupa Hedley, 1922 acutigemmata (E. A. Smith, 1877). Type albofasciata (E. A. Smith, 1877) armillata (Reeve, 1845) astricta (Reeve, 1843) subsp. consobrina Powell, new subspecies bijubata (Reeve, 1843) cerithina (Anton, 1839) cincta (Lamarck, 1822) deceptrix Hedley, 1922 diffusa Powell, new species jubata (Hinds, 1843) f maoria Powell, 1942 prestoni Powell, new name weaveri Powell, new species Austroturris Laseron, 1954 steira (Hedley, 1922). Type Taranis Jeffreys, 1870 [morchi Malm, 1863], Type alio (Jousseaume) Lamy, 1934 benthicola (Dell, 1956) bicarinata (Suter, 1915 ) f gratiosa ( Suter, 1908 ) imporcata (Dell, 1962) mayi (Verco, 1909) f nexilis (Hutton, 1885) subsp. recens (Fleming, 1948) percarinata Powell, new species spirulata (Dell, 1962) ticaonica Powell, new species turritispira (E. A. Smith, 1882) vestalis (Hedley, 1903) Micropleurotoma Thiele, 1929 ?\ashiyaensis Shuto & Ueda, 1963 spirotropoides (Thiele, 1925). Type Additional Species Gemmula Weinkauff, 1875 [p. 22-703] pseudomonilifera Powell, new species interpolata Powell, new species congener new subsp. unilineata Powell tessellata Powell, new species [p. 22 -734a] microscelida (Dali, 1895) Xenuroturris Iredale, 1929 [p. 22-967] gemmuloides Powell, new species [23 - 101] 410 Turrinae A. W. B. Powell Turridae [These occasional blank areas occur genera and subgenera to permit the of new material and future sections proper systematic sequence.] between insertion in their [23 - 102] May 15, 1967 INDO-PACIFIC MOLLUSCA, vo’. 1, no. 7 Heteroturris 411 Genus Heteroturris Powell, new genus Type: H. sola Powell, 1967 A new genus is required for a species repre- sented by only one specimen, which was dredged in 162 fathoms off the Philippines by the Albatross Expedition. This shell is problematic in its rela- tionships but appears to be most nearly allied to Lophiotoma. The general appearance of an elongate-fusiform shell with a tall spire, narrow body-whorl tapered to a long straight anterior canal, and with sculpture of a few strong smooth keels, suggests Polystira, but both the protoconch and the sinus negate inclu- sion in that genus. The single thread bisecting the sinus area of the shoulder slope and the polygyrate, axially-costate protoconch recall Turridrupa, but again both the sinus and the long, tapered anterior end do not match that genus. The regular, crisp axial threads which cross the sinus area resemble those of Microdrillia but otherwise there is no simi- larity to that genus. The deep parallel-sided pos- terior sinus is not like that in the subfamily Turri- culinae. However, it does appear to be a member of the Turrinae with a modified Lophiotoma- type sinus which, instead of being narrow and peripheral, occupies the greater part of the shoulder slope be- tween a heavy subsutural margining cord and the peripheral carina. The polygyrate, tall, conical, axially-costate protoconch and the elongate-fusi- form shell show close affinities to Lophiotoma. Heteroturris sola Powell, new species (PI. 298) Range — Philippines, south east of Talicayo, Cebu, in 162 fathoms. Description — Shell rather small, 18 mm. ( % inch ) in height, narrowly fusiform with tall spire and long straight unnotched anterior canal. Whorls 8, plus a tall, narrowly conic protoconch of approxi- mately 4-4M whorls, strongly sculptured with closely spaced, somewhat flexed, axial ribs; initial whorls, estimated at 1/2, missing. The protoconch passes abruptly into the post-nuclear sculpture, which is strongly bicarinate. The lower or peripheral carina is undulatingly nodulose for the first 2-2/2 post- nuclear whorls, after which the carinae are smooth and rounded. The uppermost of the two initial carinae resolves into a strong but narrowly crested subsutural margining cord, the lower one into the heavier smooth medial peripheral carina. There is a third smooth cord on the later whorls, between the peripheral carina and the lower suture. On the last two whorls a fourth cord is half-emergent at the lower suture. About 13 smooth cords are on the base and neck from below the peripheral carina, and there is an occasional weak intermediate thread. Between the spirals there is a dense surface sculpture of distinct, crisp, axial threads, strongest on the shoulder slope where they mark the succes- sive positions of the apex of the sinus. These are crossed medially by a single smooth thread. Spire and aperture plus canal of about equal height. Aperture long and narrow, slowly tapered below. Posterior sinus broad and deep, with parallel sides, its apex square-cut, the whole occupying the entire shoulder slope from between the subsutural mar- gining cord to the peripheral carina. Colour uni- formly ivory-white. Measurements (mm.) — height width 18.0 5.5 holotype Types — The unique holotype is in the United States National Museum, Washington ( Albatross Sta. 5412; USNM 232702). Plate 298. Heteroturris sola Powell, new genus and new species. 162 fins., off Cebu Id., Philippines. 18 mm. Holo- type, USNM no. 232702. [23 - 107] 412 Austroturris A. W. B. Powell Turridae Genus Austroturris Laseron, 1954 Type: Filodrillia steira Hedley, 1922 “A genus much more closely related to Turri- drupa than to Filodrillia, from which the sinus alone distinctly separates it. In Filodrillia the sinus is subtubular like that of Etrema, in Austroturris it is wide and open immediately above the peripheral keel and with a thin inner margin. This sinus is similar to that of Turridrupa, as is the sculpture consisting primarily of a strong peripheral keel, with lesser spiral keels below and fine axial threads between the keels. The protoconch, however, is dif- ferent, of only two smooth whorls, and without a third whorl with axial riblets. It is also distin- guished from Turridrupa by a well marked fasciole area crossed by fine curved axial threads.” (Laseron) Synonymy — 1954 Austroturris Laseron, The New South Wales Turridae. Zool. Handb. Royal Zoological Society of N.S.W., p. 6: Type by original designation: Filodrillia steira Hedley, 1922. Austroturris steira (Hedley, 1922) (PI. 299) Range — New South Wales, 30-50 fathoms. Measurements (mm.) — height width 5.8 2.6 Synonymy — 1922 Filodrillia steira Hedley, Rec. Aust. Mus., 13(6), p. 224, pi. 42, fig. 11. 1954 Austroturris steira (Hedley), Laseron, The N.S.W. Turridae, Handbook Roy. Zool. Soc. N.S.W., p. 6, pi. 1, figs. 3, 4. Records - NEW SOUTH WALES: off Cape Three Points, 50 fathoms (type locality); off Sydney, 30-50 fathoms ( Laseron ) . Types — The holotype is in the Australian Mu- seum, Sydney. Plate 299. Austroturris steira (Hedley). 5.8 mm. 30-50 fms., New South Wales, Australia (from Hedley, 1922, pi. 42, fig. 11). May 15, 1967 INDO-PACIFIC MOLLUSCA, vol. 1, no. 7 Turridrupa 413 Genus Turridrupa Hedley, 1922 Type: Pleurotoma acutigemmata E. A. Smith, 1877 This is a problematic genus since several sub- family alternatives suggest themselves. In general appearance, these shells resemble small stout clavi- nids, and in accord with that subfamily, the sinus is situated, typically, at about the middle of the shoulder slope. Here, however, comparison ends, for the clavinid sinus is spout-like, constricted above, either by a heavy subsutural fold, a strong parietal tubercle, or both. In Turridrupa the sinus is simple U-shaped, not restricted above by either a subsutural fold or by a parietal tubercle, but has its apex, normally, at the termination of one or two spiral cords or threads that bisect the shoulder area. The sinus is, therefore, comparable with that of Turns, which has its apex situated on a rib above the peripheral carina. It is of interest to note that Thiele ( 1929 ) placed Turridrupa as a section of Gemmula, an undoubted member of the Turrinae. The radula, however, presents another problem, for it has a formula of 1 + 0 + 1 + 0 + 1 ( jubata ); the marginal is massive and neither “wishbone- shaped” nor bifid; the lateral is absent; and the cen- tral large and broad-based. This enlargement of the central tooth to compen- sate for the missing lateral is a common feature of the Cochlespirinae, i.e., Aforia, and in some species ascribed to Leucosyrinx ( not the type species ) , but in all these, the marginals are bifid-based, wish- bone-shaped, or separated into two components. All other characters exclude Turridrupa from the Cochlespirinae. The protoconch, typically, has about three whorls, the first two being smooth, the last one distinctly axially costate. The operculum is leaf-shaped, with a terminal nucleus. The Recent geographical range of Turridrupa covers most of the Indo-Pacific. A New Zealand fossil species, maoria Powell, 1942, takes the genus back to the upper Miocene. Of the Recent species listed by Hedley ( 1922 ) as belonging to Turridrupa, five belong elsewhere. To the clavinid genus Microdrillia, I add com- mentica (Hedley, 1915), fastoda (Hedley, 1907) and pertinax (Hedley, 1922). To the turriculinid genus Vexitomina I add rougeyroni Souverbie, which is a synonym of regia (Reeve, 1842). The species sibogae (Schepman, 1913) is probably a clavinid. It superficially resembles Turridrupa ceri- thina (Anton, 1839), but has two smooth whorls in the protoconch. Several Japanese species referred to Turridrupa in Kuroda and Habe (1952, Check List and Bibliography of the Recent Marine Mol- lusca of Japan) would also be better accommo- dated in the subfamily Clavinae. Synonymy — 1922 Turridrupa Hedley, A revision of the Australian Turri- dae, Records of the Australian Museum, vol. 13, no. 6, p. 226. Type: by original designation: Pleuro- toma acutigemmata E. A. Smith, 1877. Plate 300. Turridrupa jubata ( Hinds ) . Radula and opercu- lum from specimen from Bohol Ids., Philippines. [23 - 117] 414 Turridrupa A. W. B. Powell Turridae Key to species and subspecies of Turridrupa A. Sinus apex at end of mid-shoulder cord Sinus cord finely gemmate Colour whitish with subsutural brown band armillata (Reeve) Colour yellowish brown; anterior end, white jubata (Hinds) Sinus cords of elongated knots Small narrow shell (17-21 mm.) Knots large; few, 13-14 per whorl acutigemmata (E. A. Smith) Large broad shell ( 25-33 mm. ) Knots weak, 17 per whorl prestoni Powell All cords smooth Shell broad; unicoloured yellowish brown cincta (Lamarck) Shell tall, narrow; unicoloured buff . . deceptrix (Hedley) All cords smooth to waved Cords buff on dark brown ground .... bijubata ( Reeve ) Peripheral cord tuberculate Reddish brown; pale subperipheral zone albofasciata (E. A. Smith) B. Sinus apex at end of several mid-shoulder threads All spirals gemmate Colour pattern checquered cerithina ( Anton ) C. Sinus apex at end of peripheral cord Peripheral cord only maculated Alternate white and brown dashes .... astricta (Reeve) Unicoloured dashes, defined by blotches . . diffusa Powell All spire cords maculated Alternate light and dark dots . . astricta consobrina Powell Spire brown-streaked; base pale weaveri Powell Plate 301. Fig. 1, Turridrupa acutigemmata (E. A. Smith). Fig. 4, T. prestoni Powell, new name. 33 mm. Port Blair, Holotype, 21 mm., no locality. Fig. 2, T. deceptrix Hedley. Andaman Islands. Fig. 5, T. cincta (Lamarck). 16.5 mm. 14 mm. Damley Id., Queensland, Australia. Fig. 3, T. ju- Mauritius. bata (Hinds). 22.5 mm. Off Arakabesan Id., Palau Islands. [23-118] May 15, 1967 INDO-PACIFIC MOLLUSCA, vol. 1, no. 7 Turridrupa 415 Turridrupa acutigemmata (E. A. Smith, 1877) (PI. 301, fig. 1) Range — Persian Gulf to East Indies, Japan and north Queensland. Remarks — This species is easily recognized by its slender shape, tall spire and comparatively few, strong, laterally elliptical gemmules on the sinus rib; each gemmule is buttressed top and bottom by a swelling. In jubata, the nearest related species, the gemmules are more numerous, less conspicu- ous, and somewhat oblique, which imparts the ap- pearance of a strand of unravelled rope. Description — Shell small, 17-21 mm. ( about % inch ) in height, narrowly fusiform, with a tall spire, almost twice the height of the aperture and canal. Spire whorls sculptured with three strong keels and a fourth emergent, or half emergent, over the last two or three whorls. Base with an additional seven keels, from below the lower sutural one to the ante- rior fasciole, which bears several indistinct spirals. Subsutural keel smooth, moderately strong, sinus keel strong, with laterally elliptical gemmules, about 13 or 14 per whorl, which are buttressed top and bottom by swellings, followed by a massive smooth peripheral keel; all keels other than the sinus one are smooth. Interstices of keels with two or three fine spiral threads. Sinus deep, U-shaped, its apex at the termination of the gemmate keel. Colour uniformly yellowish brown to reddish brown. Measurements (mm.) — height width 21.0 7.5 holotype 17.0 6.0 New Britain Synonymy — 1877 Pleurotoma acutigemmata E. A. Smith, Ann. Mag. Nat. Hist., Ser. 4, 19, p. 489. 1884 Pleurotoma jubata (non Hinds) (pars), Tryon, Man. of Conch., 6, p. 171. 1896 Pleurotoma acutigemmata Smith, Melvill and Sykes, Proc. Malac. Soc. 2, p. 165. 1904 Pleurotoma acutigemmata var. minor E. A. Smith, Ann. Mag. Nat. Hist., Ser. 7, 13, p. 457 (non Pi. minor C. B. Adams, 1845; Evans and Schumard, 1857, nor Jeffreys, 1867). 1913 Pleurotoma (Hemipleurotoma) acutigemmata Smith, Schepman, Siboga Exped., Pt. 5, 49e, p. 400. 1917 Turns ( Tomopleura ) acutigemmata (Smith), Melvill, Proc. Malac. Soc., 12, p. 146, pi. 8, fig. 1. 1922 Turridrupa acutigemmata (Smith), Hedley, Rec. Aust. Mus., 13(6), p. 226, pi. 42, figs. 12, 13. 1940 Tunis (Turridrupa) acutigemmata var. minor (Smith), Winckworth, Proc. Malac. Soc., 24(1), p. 24. 1960 Turridrupa jubata (non Hinds) (pars), Oyama and Takemura, The Molluscan Shells, 4, fig. 9 ( = acuti- gemmata, non fig. 10, which is jubata). Records — Locality unknown ( type ) ; PERSIAN GULF : Gulf of Oman, Muscat, 20-40 fathoms (Melvill, 1917). CEYLON; off south coast (type of var. minor). INDO- NESIA: Bay of Bima, 55 metres; near south coast of Timor, 34 metres (Schepman, 1913). WEST NEW GUINEA: straits north of Misool Id. (Schepman, 1913). NEW BRITAIN ( AWBP coll.). JAPAN: Izu Peninsula (Oyama and Takemura, 1960). QUEENSLAND: Damley Island, 20 fathoms; Hope Island, 5-10 fathoms (Aust. Mus.); Low Isles, near Port Douglas; Lindeman Island (AWBP coll.). FIJI: main reef, Navula Passage, 2 feet (W. Jennings, Aug., 1963). Types — The holotype of acutigemmata is in the British Museum (Natural History). Plate 302. Geographical distribution of Turridrupa acuti- gemmata (E. A. Smith). [23 - 123] 416 Turridrupa A. W. B. Powell Turridae Turridrupa albofasciata (E. A. Smith, 1877) (PI. 303, figs. 3, 4) Range — Hawaiian Islands, Japan and New Cale- donia. Remarks — This common Hawaiian turrid occurs in many collections under a manuscript name of Dali’s, which I do not quote, since to the best of my knowledge the name has not as yet appeared in print. Elsewhere in his manuscript Dali referred to albofasciata Smith as an indeterminate species, since it had never been figured. However, the well- preserved holotype, in the British Museum, leaves no doubt that albofasciata is the name to be used for this Hawaiian shell. The species stands close to bijubata, from which it differs in the very strong undulations of the peripheral keel, a different form of subsutural mar- gin, fewer and stronger interstitial spiral threads, and the presence of a pale band between the pe- ripheral and lower keels. The species bijubata occurs in the Hawaiian group also but not commonly. Also, although some forms of bijubata, notably those from both Mau- ritius and New Caledonia, develop strong undula- tions, the form of the subsutural margin and the coarser, decussated interstitial sculpture remain constantly exclusive to albofasciata, as also does, apparently, the pale submedian band. It is likely that Hervier’s gatchensis from the Loyalty Islands is a synonym of albofasciata rather than a nodulose form of bijubata. Hervier’s figure, which is small and indistinct, shows a deep violet- grey shell with a subperipheral reddish brown spiral band, and both the peripheral and subpe- ripheral spirals appear to be undulating and sub- nodose. Also the Japanese shells ascribed to gatch- ensis by Oyama and Takemura ( 1960, pt. 4, figs. 6-8) appear to be Smith’s species. Description — Shell small, 14-22 mm. ( about % inch) in height, claviform, very solid. Spire-whorls sculptured firstly with a rather wide subsutural band, composed of four spirals, three of them threads, but the fourth is a moderately strong cord; then a strong median cord which is conspicuously undulated and thickened where it crosses rather distant axial folds, which spread as buttresses above the peripheral cord but not below it, and finally, another strong but smooth cord between the periphery and the lower suture. Base with six strong cords, distantly spaced between the periph- ery and the anterior fasciole, which is sculptured with 5 or 6 irregular linear-spaced cords. Interstices of the keels and cords with 3 or 4 distinct spiral lirations, densely decussated by finer crisp axial threads. Spire almost twice height of aperture plus canal. Sinus deep, U-shaped, and situated at the termination of the peripheral keel. Colour dark chocolate to dark purplish brown, with a pale buff or light-brown band between the peripheral and lower keels. In partially bleached specimens the pale band is more conspicuous. Operculum leaf- shaped with a terminal nucleus. Measurements (mm.) — height width 22.0 8.0 holotype 21.0 8.0 Honolulu, dredge deposits 18.2 7.3 Honolulu, dredge deposits 14.0 6.0 Honolulu, off Waikiki beach Synonymy - 1877 Pleurotoma albofasciata E. A. Smith, Ann. Mag. Nat. Hist., Ser. 4, 19, p. 491. 1895 Surcula gatchensis Hervier, Journ. de Conch., 44, p. 61, pi. 1, fig. 7. 1914 Surcula bijubata gatchensis Hervier, Bouge and Daut- zenberg, Joum. de Conch., 61, p. 145. 1960 Turridrupa gatchensis (Hervier), Oyama and Take- mura, The Molluscan Shells, pt. 4, figs. 6-8. Records — HAWAIIAN ISLANDS ( type locality ) ; Hawaii, Keokea, Hilo, in sand under seaweed (Thaanum coll., Bishop Mus. ); Oahu, off Waikiki, 25-50 fathoms; Honolulu Harbour entrance, 5-8 fathoms; Paumalu (USNM); Pupu- kea Beach and Waialua (USNM); Maui, off Launiupoko Camp, 4-12 fathoms (Thaanum coll., USNM); Maalaea Bay, 50 feet, in coralline algae (A. Tiedeman, 1964); Mc- Gregor’s Landing, Madaea Bay, 25-30 feet (A. Tiedeman, 1964); French Frigate Shoal (ANSP). JAPAN: Amami- Oshima Islands (Oyama and Takemura). LOYALTY IS- LANDS: Lifu (Hervier, 1895). NEW CALEDONIA: (Bouge and Dautzenberg, 1914). Types — The holotype of albofasciata is in the British Museum (Natural History); that of gatch- ensis in the collection of the Journal de Conchy li- ologie, Mus. d’Hist. Nat., Paris. Turridrupa armillata (Reeve, 1845) (PI. 305, fig. 2) Range — Philippine Islands. Remarks — This species belongs to the acuti- gemmata-jubata-prestoni series, in which the sinus keel, between the subsutural fold and the periph- ery is gemmate. In jubata and prestoni the gem- mules are rather distant, laterally elongated and knot-like, but in armillata they are numerous, closely-spaced beads. The holotype in the British Museum is the only specimen known to me, and this has a distinctive colour pattern: whitish, with a subsutural band of [23 - 124] May 15, 1967 INDO-PACIFIC MOLLUSCA, vol. 1, no. 7 Turridrupa 417 Plate 303. Figs. 1, 2, Turridrupa bijubata (Reeve). Fig. 1, Mauritius. 24 mm. Fig. 2, Island of Burias, Philippines. Syn- type, 21 mm. Figs. 3, 4, T. albofasciata (E. A. Smith). Fig. 3, Hawaiian Islands, holotype, 22 mm. Fig. 4, French Frigate Shoals, 22 mm. Fig. 5, T. weaveri Powell, new species, holotype, 21.7 mm. 45 ft. off Rabbit Island, Oahu, Hawaiian Islands. light-brown. It seems most nearly related to jubato, but unfortunately the type of that species, which should be in the British Museum, could not be located. Description — Shell small, 18.5 mm. (1 inch) in height, somewhat broadly fusiform, with spire about 1/3 times height of aperture plus canal. Spire whorls sculptured with three strong subequal keels; first smooth and submargining the suture, second, the sinus keel, densely gemmate, followed by the smooth peripheral keel; base with the addition of seven strong smooth keels to the anterior fasciole, which has 4-5 closely spaced cords. Colour as de- scribed above. Measurements (mm.) — height width 18.5 7.25 Synonymy — 1845 Pleurotoma armillata Reeve, Conch. Iconica, 1, pi. 21, fig. 176. Types — The holotype is in the British Museum ( Natural History ) . The type locality is “Philippines.” Turridrupa astricta subspecies astricta (Reeve, 1843) (PI. 305, fig. 4) Range — Tuamotu Archipelago. Remarks — This species differs from the other members of the genus in having a maculated pe- ripheral sinus rib. This rib is a single smooth keel. quite unlike the double keel of Lophiotoma. In the form of the sculpture and in the style of U-shaped sinus, the species is not very dissimilar from cincta. Unfortunately only a few worn shells are available and all have the apical whorls missing. Description — Shell small, 14 mm. (& inch) in height, drilliaform, somewhat cylindrical, very solid, sculptured with strong smooth spiral keels, the peripheral sinus keel maculated. Spire almost twice height of aperture plus canal. Spire-whorls with three equidistant keels, the median one stronger than the other two, which are subequal. Base with an additional five keels, and 4 or 5 more closely spaced spiral cords on the anterior fasciole. Inter- spaces with 3 or 4 fine, crisp, spiral threads. Sinus U-shaped, its apex at the termination of the periph- eral keel. End of the subsutural keel somewhat thickened, but no parietal tubercle or callus-pad. Anterior canal very short, with an oblique shal- lowly-notched end. Colour bright, yellowish buff, except for the peripheral keel which is white and regularly maculated with dark reddish brown dashes. A Japanese shell ascribed to astricta, by Oyama and Takemura, 1960 (see under modesta, following), has the peripheral maculations more numerous and in the form of squarish dots. The colour pattern and truncated anterior end recalls some members of Lophiotoma of the ab- breviata series, but as already remarked, the sinus in that genus is a deep narrow slit and its associ- ated keel is double-edged with a concavity in the middle. [23 - 125] 418 Turridrupa A. W. B. Powell Turridae Measurements (mm.) — height width 14.0 5.5 Tuamotu Archipelago (ex Garrett) 13.5 5.0 Lectotype; one of two co-types Synonymy — 1834 Pleurotoma interrupta Sowerby, Proc. Zool. Soc., Lon- don, for 1833, p. 138 (non Lamarck, 1816). 1843 Pleurotoma astricta Reeve. Conch. Iconica, 1, pi. 12, fig. 98 ( nom. nov. for Pi. interrupta Sowerby, 1834; non Lamarck, 1816). Records — “Island of Annaa, Chain Island” = Island of Anaa, Tuamotu Archipelago, under coral on reefs ( type locality); “Paumotu Isles” = Tuamotu Archipelago (Garrett coll., B. P. Bishop Mus. ). Types — Two co-types, one selected as lectotype, in the British Museum (Natural History). Turridrupa astricta subspecies consobrina Powell, new subspecies (PI. 305, fig. 3) Range — Hawaiian Islands and Amami-Oshima Islands, Japan. Remarks — This seems to be a North Pacific sub- species of astricta, differing slightly but constantly from the typical South Pacific species in having all three cords of the spire-whorls of equally strong development, and smaller, more numerous macula- tions, which are not confined to the sinus rib. In astricta typical, the maculations are more widely spaced, elongated dashes, confined to the sinus cord, which is of heavier build than the two other cords of the spire- whorls. Description — Shell rather small, 19 mm. (% inch) in height. Spire twice height of aperture plus canal. Whorls 14 including a narrowly conic protoconch of 4/2 whorls, which are closely sculptured with concavely arcuate axials crossed by minute indis- tinct spiral lirae. Spire-whorls with three equally strong but narrowly ridged spiral cords. Five simi- lar cords on the body-whorl from the sinus cord to the neck and a further three on the anterior fasci- ole. From 2-5 very fine spiral threads in each inter- space, except for a wider interspace on the neck, which bears about nine threads. Entering parietal callus-pad heavy. Sinus deep, U-shaped, at the ter- mination of the middle cord of the spire-whorls. Colour buff to pale golden-brown, maculated with dark reddish brown spots, about 15 per whorl on the sinus cord but irregularly disposed also on the other cords of both the spire and the base. Measurements (mm.) — height width 19.0 7.0 holotype 14.5 5.0 paratype Synonymy — 1960 Turridrupa astricta (Reeve), Oyama and Takemura, The Molluscan Shells, 4, figs. 11, 12 (non Reeve, 1843). Records — HAWAIIAN ISLANDS: Maalaea Bay, Maui Id., in and under calcareous algae, 40 to 50 feet (j. Kern and A. Tiedeman, ex. C. Weaver) (holotype). JAPAN: Amami-Oshima Islands (Oyama and Takemura, 1960). Types — The holotype and paratypes presented to the B. P. Bishop Museum, Honolulu, by Mr. J. Kern. Turridrupa bijubata (Reeve, 1843) (PI. 303, figs. 1, 2) Range — Mauritius, Japan to Hawaiian Islands, Philippines, Queensland, New Caledonia and Fiji. Remarks — This strongly keeled, solid, small shell has the keels buff-coloured on a dark chocolate ground. Three keels encircle the spire-whorls; these are not gemmulate but often the sinus keel is some- what waved and thickened on the crests of low axial folds. Description — Shell small, 18-21 mm. (about % inch) in height, drilliaform, very solid, sculptured with strong, sharply raised spiral keels, three on spire-whorls, one subsutural and slightly weaker than the other two; median one, the sinus rib, undulating and slightly thickened at the crests of very weak broad axial folds; base with an addi- tional four keels and five more closely-spaced, weaker spirals on the anterior end. Interstices of the keels with 4 or 5 very fine, crisp, spiral threads. Spire almost twice height of aperture plus canal. Sinus U-shaped, its apex on the second or middle keel of the spire-whorls. Anterior canal very short, flexed, with an oblique shallowly notched termina- tion. Colour dark purplish brown or chocolate, with the keels buff to light-brown. Measurements (mm.) — height width 21.0 8.25 one of three syntypes 18.5 8.0 Low Isles, Queensland 18.0 7.0 San Cristoval, Solomon Islands Synonymy — 1843 Pleurotoma bijubata Reeve, Conch. Iconica, 1, pi. 10, fig. 87. 1914 Surcula bijubata (Reeve), Bouge and Dautzenberg, Journ. de Conch., 61, p. 144. [23 - 126] May 15, 1967 INDO-PACIFIC MOLLUSCA, vol. 1, Turridrupa 419 Plate 304. Geographical distribution of Turridrupa jubata Smith), T. prestoni Powell and T. weaveri Powell. (Hinds), T. bijubata (Reeve), T. albofasciata (E. A. 1914 Surcula bijubata nodulosa Bouge and Dautzenberg, Journ. de Conch., 61, p. 145. 1960 Turridrupa bijubata (Reeve), Oyama and Takemura, The Molluscan Shells, pt. 4, figs. 3-5. Records — MAURITIUS (AWBP. coll.). ANDAMAN IS- LANDS: Long Island, Port Blair, dredged (Winckworth coll., Brit. Mus.). PHILIPPINES: Island of Burias, under stones at low water (type locality). JAPAN: (Oyama and Takemura) Oshima-Osumi (USNM). HAWAIIAN IS- LANDS: Oahu, Mokolea Rock, Kailu Bay, 55-65 feet, in sand pockets under coral (C. Weaver, Dec., 1963); Maui, 50 feet, in coralline algae (A. Tiedeman and j. Kern, 1964). SOLOMON ISLANDS: San Cristoval (Aust. Mus.). QUEENSLAND: Low Isles; North West Island (Mrs. J. Kerslake ) . NEW CALEDONIA: Poindimie (Mrs. J. Kers- l.ike ) . LOYALTY ISLANDS: Lifu (Aust. Mus.). FIJI: (Garrett coll., ANSP); Korolevu (Mrs. J. Kerslake); Navula Passage, main reef, 2 feet (W. Jennings, Aug., 1963). SAMOA: Vailele Beach (AWBP. coll.). Types — There are three syntypes in the British Museum (Natural History). The type locality is the Island of Bureas [Burias], Philippines. Plate 305. Fig. 1, Turridrupa cerithina (Anton), syntype of Pleurotoma digitale Reeve, 24 mm. Island of Burias, Philippines. Fig. 2, T. armillata (Reeve), holotype, 18.5 mm., Philippine Islands. Fig. 3, T. astricta consobrina Powell new subspecies, holotype, 19 mm. 40-50 ft., Maui Id., Hawaiian Islands. Fig. 4, T. astricta astricta (Reeve). 14 mm., Tuamotu Archipelago. Fig. 5, T. diffusa Powell new species, holotype, 12 mm., Samoa. 420 Turridrupa A. W. B. Powell Turridae Plate 306. Geographical distribution of Turridrupa cerithina (Anton), T. astricta (Reeve), its subspecies consobrina Powell and T. diffusa Powell. Turridrupa cerithina (Anton, 1839) ( PI. 305, fig. 1 ) Range — Mauritius to Japan, Philippines, Queens- land, New Caledonia and Fiji. Remarks — In this species all the spiral keels and cords are gemmate at the points of intersection with the numerous narrow flexuous axials. Colour dark chocolate with the gemmules picked out in white. The apex of the normal U-shaped sinus is at the termination of one or two spiral threads which bisect the shoulder sulcus. Description — Shell of moderate size, 18-24.5 mm. ( %-l inch) in height, solid, drilliaform, with a tall spire but with a truncated anterior end. Sculpture of spiral keels and cords which are densely gemmu- late at the points of intersection with numerous narrow flexuous axial ribs. The suture is submar- gined by a closely spaced pair of spirals, the upper one weak but the lower one massive. This is fol- lowed by the rather narrow, deeply concave shoulder sulcus which bears one or two fine spiral threads. Peripheral spiral on the site of a slight angulation, below which the spirals continue over the base to the anterior fasciole. There are from 1-3 spirals between the periph- eral keel and the lower suture or 7-10 between the periphery and the anterior fasciole; 5 or 6 weaker, linear-spaced, smooth cords on the anterior fasci- ole. Spire about 1A times height of aperture plus canal. Anterior canal very short and weakly notched, with an oblique termination. In senile specimens the subsutural fold thickens at its termination to form a callosity, slightly restricting the U-shaped sinus which is at the termination of the one or two threads bisecting the shoulder sulcus. Colour dark chocolate with the gemmules picked out in white; pillar light reddish brown; interior of aperture light purplish grey. Measurements (mm.) — height width 24.5 9.0 Japan 24.0 9.0 one of three syntypes 20.0 7.5 Mauritius Synonymy — 1839 Pleurotoma cerithina Anton, Verzeichniss der chylien, p. 73, sp. 2504. 1843 Pleurotoma digitale Reeve, Conch. Icon., 1, pi. 17, fig. 138. 1884 Drillia (Crassispira) digitalis (Reeve), Tryon, Man. of Conch., 6, p. 191. pi. 13, fig. 75. 1914 Drillia (Crassispira) digitalis (Reeve), Bouge and Dautzenberg, Journ. de Conch., 61, p. 143. 1922 Turridrupa cerithina (Anton), Hedley, Rec. Aust. Mus. 13(6), p. 226. Records — PHILIPPINES: Island of Burias, under stones at low water (type locality). JAPAN: Oshima-Osumi (USNM). NEW GUINEA: Finschafen (T. Garrard). QUEENSLAND: Low Isles (AWBP coll.); Buchan’s Point (Mrs. J. Kerslake ) . NEW CALEDONIA: (Bouge and Dautzenberg, 1914). FIJI (USNM). ANDAMAN IS- LANDS: Port Blair (Winckworth coll., Brit. Mus.). MAU- RITIUS: ( ANSP and AWBP coll.). Types — Three syntypes of digitale are in the British Museum (Natural History). [23 - 128] INDO-PACIFIC MOLLUSCA, vol. 1, no. 7 Turridrupa 421 May 15, 1967 Plate 307. Geographical distribution of Turridrupa cincta (Lamarck) and T. deceptrix Hedley. Turridrupa cincta (Lamarck, 1822) (PI. 301, fig. 5) Range — Mauritius, Queensland, Japan and Fiji. Remarks — This shell stands nearest to bijubata in size and general appearance but is easily distin- guished from that species by the uniform yellowish brown colour, lack of axial undulations, and differ- ent arrangement of the keels on the spire-whorls, the subsutural one being far heavier than the other two, of which the median sinus one is quite weak. Description — Shell small, 14.5-16.5 mm. ( about /I inch ) in height, drilliaform, very solid, sculptured with strong spiral keels, with 3 or 4 fine, crisp, spiral threads in each interspace. Spire-whorls with a massive subsutural keel, followed by a very weak median one, the sinus rib, and then a strong one below it; four more strong keels on the base, upper- most emergent at the suture, and five closely- spaced cords on the anterior end. Spire more than 1/2 times height of aperture plus canal. Apical whorls acuminate. Sinus small, narrowly U-shaped, its apex on the weak median spiral. Anterior canal very short, with an oblique, shallowly-notched ter- mination. Colour uniform light yellowish brown. Measurements (mm.) — height width 16.5 7.0 Mauritius 16.0 6.5 Mauritius 14.5 6.5 Fiji Synonymy — 1822 Pleurotoma cincta Lamarck, Hist. nat. anim. sans Vert., ed. 1, 7, p. 92. 1939-40 Pleurotoma cincta (Lamarck), Kiener, Icon. Coq. Viv., Pleurotome, p. 60, pi. 19, fig. 5. 1960 Turridrupa cincta (Lamarck, Oyama and Take- mura, The Molluscan Shells, pt. 4, figs. 1, 2. Records — “les mers de Pile de France” = Mauritius (type locality) (AWBP coll.); Barkley Island (Mrs. J. Kerslake). JAPAN: Amami-Oshima Islands (Oyama and Takemura). QUEENSLAND: North West Island (Mrs. J. Kerslake); Portland Roads, Cape York Peninsula (T. Garrard). FIJI: (A. Garrett, ANSP). “HEBVEY ISLANDS” = COOK IS- LANDS: (B. P. Bishop Mus.). NEW CALEDONIA: Poindimie (Mrs. J. Kerslake). Turridrupa modesta ( Sowerby, 1834 ) Localities — “Real Llejos [Nicaragua] et ad Insu- lam Annaa” [Tuamotu Archipelago]. Remarks — Upon the available evidence, the status of this species cannot be determined, for the type, which should be in the British Museum, can- not be found. Reeve in 1843 ( Conch. Icon. 1, pi. 12, fig. 99) and Tryon (1884, Man. of Conch. 6, p. 241) considered it a synonym of cincta Lamarck, 1822, and said the first mentioned locality, “Real Llejos,” was probably erroneous. Shells in the Garrett col- lection at the B. P. Bishop Museum, labelled “PL modesta Sowerby, Hervey Islands” are undoubtedly cincta, and since Garrett probably relied upon the British Museum for his identifications, Sowerby ’s species may well be a synonym of cincta as Reeve claimed. Also, Sowerby made no mention of peripheral maculations in his original description of modesta, but he did mention “cingulo mediano nigro, albo articulata” for his interrupta (i.e., astricta ), also from the Island of Anaa, and described in the same paper. [23 - 131] 422 Turridrupa A. W. B. Powell Turridae On the other hand, Oyama and Takemura (1960, The Molluscan Shells, pt. 4, figs 13, 14), figured a maculated Japanese shell as modesta, and this is certainly not cincta. A shell very similar to Oyama and Takemura’s “ modesta ” from Samoa (AWBP coll. ) likewise has the peripheral dashes uncoloured, but defined by surrounding diffused blotching in brown. See T. diffusa Powell, new species. Synonymy — 1834 Pleurotoma modesta Sowerby, Proc. Zool. Soc., Lon- don, for 1833, p. 136. 1843 Pleurotoma cincta Lamarck, Reeve, Conch. Icon., 1, pi. 12, fig. 99. Turridrupa deceptrix Hedley, 1922 (PI. 301, fig. 2) Range — North Queensland and New Guinea to the Andaman Islands. Remarks — This species resembles jubata in its crisp, narrow, spiral keels but has more the shape of acutigemmata. It differs from both, however, in the lack of gemmules. Description ( original )— “Shell elongate-conic, very solid, contracted at the base, constricted and channelled at the suture, last whorl about half the total length; eleven whorls, including the proto- conch. Colour pale ochraceous-buff, aperture lighter. Sculpture: — Last whorl with thirteen, penultimate with four, and earlier whorls with three prominent spiral keels, the furrows between which carry faint radial striae, and sometimes a small interstitial thread. Aperture: — There is a thin callus sheet on the inner lip, and a solid callus plug at the angle of the aperture; outer lip simple; sinus a semicircular notch with reflected margin; canal short, open, and slightly recurved; deep within the throat are five revolving raised threads.” Measurements (mm.) — height width 19.0 6.5 Japen Island, West New Guinea 14.0 6.0 holotype Synonymy — 1922 Turridrupa deceptrix Hedley, Rec. Aust. Mus., 13(6), p. 227, pi. 42, fig. 14. Records — QUEENSLAND: Damley Island, 30 fathoms (type locality). WEST NEW GUINEA: 1 mi. S.E. of Cape Dgarwawoffi, Japen Island, 10-16 fathoms, coarse gravel and silt ( NSF, ANSP). ANDAMAN ISLANDS: Port Blair (Winckworth coll., Brit. Mus.). INDONESIA: Sunda Strait, 30 metres (Th. Mortensen, 1922, Zool. Mus., Copenhagen). T types — The holotype is in the Australian Mu- seum, Sydney. Turridrupa diffusa Powell, new species (PI. 305, fig. 5) Range — Samoa and Japan. Remarks — This species belongs to the astricta group. It is readily distinguished by its narrow form and the reversed colour pattern, in that the periph- eral nodes are colourless, being defined by a sur- rounding diffused brown blotching. The Japanese record, based upon Oyama and Takemura’s figures, shows a shell with smaller and more numerous nodes. When more material is available this may prove to be distinct from typical diffusa. Description — Shell small, 12 mm. (M inch) in height, drilliaform, narrowly subcylindrical, solid, sculptured with strong spiral cords, smooth except for the sinus one, which bears laterally elongated slightly raised losenge-shaped nodes. These nodes are accentuated by being picked out by a surround- ing diffusion of pale reddish brown blotches on an otherwise creamy-white ground. Spire less than twice height of aperture plus canal. Postnuclear whorls about 10 ( apex eroded ) . Spire- whorls sculp- tured with a heavy, narrowly ridged, subsutural cord, followed by the weakly nodulose sinus rib, and two plain, narrowly ridged cords between it and the lower suture. The subsutural cord is strong- est, but those following it diminish only slightly in strength; seven cords on the body-whorl from below the sinus rib to the end of the anterior canal; one or two intermediate spiral threads over the region of the neck. There are about nine nodes on the sinus cord of the last whorl. Parietal callus-pad heavy. Sinus deep, U-shaped, at the termination of the nodulose rib. Measurements (mm.) — height width 12.0 4.5 holotype Synonymy — 1960 Turridrupa modesta (Sowerby), Oyama and Take- mura, The Molluscan Shells, pt. 4, figs. 13, 14 (non Sowerby, 1834). Records — SAMOA (ex A. J. Garrett) (holotype). JAPAN: Amami-Oshima Islands (Oyama and Takemura, 1960). Types — The holotype is in the Powell collection, Auckland Museum. [23 - 132J May 15, 1967 INDO-PACIFIC MOLLUSCA, vol. 1, no. 7 T urridrupa 423 Turridrupa jubata (Hinds, 1843) (PI. 301, fig. 3) Range — Japan to Melanesia. Remarks — It is difficult to be positive about the identity of jubata since the holotype can no longer be found, but there seems to be little doubt that the species is closely allied to acutigemmata, from which it differs in being more broadly fusiform and in having more numerous and smaller gemmules on the sinus rib. These gemmules are not only laterally compressed but are slightly oblique as well, which gives something of the appearance of a strand un- ravelled from a rope. Description — Shell small, 26 mm. ( 1 inch ) in height, rather broadly fusiform with tall spire, about lfi times height of aperture plus canal, which is relatively long for the genus. Sculpture of nar- row, sharply raised spiral keels, three on the spire- whorls; first smooth and subsutural; second, the sinus rib is delicately beaded ( about 25 per whorl ) ; third keel smooth. On the base are seven additional and equally strong smooth keels, plus five weak but crisp spiral cords on the fasciole. Surface smooth except for excessively fine axial growth lines. Sinus deep, narrowly U-shaped, its apex at the termina- tion of the beaded keel. Colour pale yellowish brown, sometimes with the anterior end white. Measurements (mm.) — height v vidth 26.0 - fide Hinds 22.5 8.5 Palau Is., Arakabesan Id., dredged 22.0 8.0 Palau Is., Malakal Harbour, 25-30 fathoms Synonymy — 1843 Pleurotoma jubata Hinds, Proc. Zool. Soc. for 1843, London, p. 37. 1843 Pleurotoma jubata Hinds, Reeve, Conch. Iconica, 1, pi. 7, fig. 52. 1844 Pleurotoma jubata Hinds, Voy. H.M.S. Sulphur, 2, p. 15, pi. 5, fig. 3. 1913 Pleurotoma ( Hemipleurotoma ) jubata Hinds, Schep- man, Siboga Exped., Pt. 5, 49e, p. 400. 1960 Turridrupa jubata (Hinds), Oyama and Takemura, The Molluscan Shells, pt. 4, fig. 10 (not fig. 9, which is acutigemmata) . Records — STRAITS OF MALACCA: found in mud at the depth of eighteen fathoms (type locality). JAPAN: Izu Peninsula (Oyama and Takemura, 1960); Shionomizaki, Kii ( ANSP). PALAU ISLANDS: dredged off Arakabesan Is- land (ANSP); off Malakal Harbour, 25-30 fms. (ANSP). WEST NEW GUINEA: mi. S. of Maroepi, Ambai Islands, Japen Island, 14-25 fms.; 2 mi. N. of Matas, Aoeri Islands, Geelvink Bay, 18-20 fms. (ANSP). CELEBES: between islands of Wowoni and Buton, 75-94 metres (Schepman 1913). NEW CALEDONIA: Poindimie (Mrs. J. Kerslake). Turridrupa prestoni Powell, new name (PI. 301, fig. 4) Range — Andaman Islands. Remarks — The name Pleurotoma rimata Preston, 1908, is preoccupied by Pleurotoma (Drillia) rimata E. A. Smith, 1888, so a new name is hereby pro- vided. This species stands between jubata and acutigemmata. It has the general shape of jubata but attains a much larger size. The interstices of the keels are spirally lirate, not smooth, and the gemmules are laterally elongated as in acutigem- mata, and there are more of them, 17, compared with 13 or 14 in the latter species. Description — Shell large for the genus, 25-33 mm. ( l-lfi inches) in height, broadly fusiform with tall spire, almost 1M times height of aperture plus canal, which is relatively long for the genus. Sculp- ture of stout but narrowly crested keels, of varying strength, with from 1-4 fine crisp spiral threads in most interspaces. Spire-whorls with three keels and a fourth sometimes emergent over the last two whorls; firstly a moderately strong smooth sub- sutural keel, followed by the gemmate sinus keel, also moderately strong, and then a massive smooth peripheral keel just above the lower suture. On the base, from level with the top of the aperture to the anterior fasciole, are a further eight spiral keels, followed by five cords on the fasciole. Sinus deep, U-shaped, its apex at the termination of the gem- mate median keel. Colour yellowish brown, paler over neck and anterior end. Measurements (mm.) — height width 33.0 12.5 Port Blair, Andaman Islands 33.0 12.0 Port Blair, Andaman Islands 25.5 10.0 holotype Synonymy — 1908 Pleurotoma rimata Preston, Rec. Indian Mus., 11, p. 190, pi. 17, fig. 62 (non Pleurotoma (Drillia) rimata E. A. Smith, 1888). Records — ANDAMAN ISLANDS (type locality): Port Blair (Winckworth coll., Brit. Mus.). Turridrupa weaveri Powell, new species (PI. 303, fig. 5) Range — Hawaiian Islands. Remarks — This species is not very closely allied to any other described member of the genus. It does, however, show some resemblance to cerithina (Anton, 1839) in that both have a tessellated pat- tern in dark-brown upon a pale ground, but in [23 - 133] 424 Turridrupa A. W. B. Powell Turridae cerithina the pattern is over the entire shell, whereas in weaveri it is absent from both the early spire- whorls and the base. Other points of difference between the two species are that the keels are gem- mate in cerithina but smooth in weaveri ; also there are 1-3 spiral threads in the shoulder sulcus of cerithina but none in weaveri. The species is named for Mr. Clifton Weaver of Hawaii, who generously made available all the Ha- waiian turrids in his collection. Description — Shell comparatively large, 18-23 mm. (about 1 inch) in height. Spire tall, almost twice height of aperture plus canal. Protoconch eroded away in the holotype, but in a not fully adult specimen from Mokumanu Island, Oahu, it is conical, with a low dome-shaped smooth initial whorl, followed by 1% whorls sculptured with strong, closely-spaced, vertical axials which termi- nate abruptly as the tricarinate post-nuclear sculp- ture commences. Spire- whorls sculptured with three cords plus a fourth, emergent over the last whorl. Subsutural cord massive, followed by the sinus cord of less than half that strength, and one or two cords below, which are slightly heavier than the sinus cord. Three or four strong widely spaced cords on the base, with 3 or 4 threads in each interspace, followed by 5 or 6 irregular cords on the anterior fasciole. Colour creamy white, heavily maculated with dark reddish brown in an irregular tessellated pattern. The maculations are absent from the early spire-whorls and cease abruptly at the lower suture, which leaves the base unicoloured. Measurements (mm.) — height width 18.0 7.0 Mokumanu Id., Oahu 21.7 8.75 holotype 23.0 9.0 off Kewalo, Oahu Records — HAWAIIAN ISLANDS: 45 feet, off Rabbit Island, Oahu (type) (C. Weaver, Nov., 1961); off Pokoi Bay, 40 feet, sand and coral; off Kewalo, Oahu, 20 fathoms (Mrs. M. King, March 28, 1962); off Waikiki, Oahu, 100- 190 fathoms (P. Burgess, April-June, 1953); Mokumanu Island, Oahu, 65-75 feet, attached to base of small coral chunk (A. Tiedeman and C. Weaver, March 26, 1965). Types — The holotype is presented to the B. P. Bishop Museum, Honolulu by Mr. Clifton Weaver. FOSSIL SPECIES Turridrupa maoria Powell, 1942 Range — New Zealand, upper Miocene. Remarks — The record of this genus in the New Zealand upper Miocene is based upon a single im- perfect specimen. However, the presence of a spiral rib traversing the sinus area, the tuberculate pe- ripheral spiral keel, and evidence of axial costae on what remains of the nuclear whorls, make the generic reference almost certain. Measurements (mm.) — height width 5.3 2.25 Synonymy — 1942 Turridrupa maoria Powell, Bull. No. 2, Auck. Inst. Mus., p. 117, pi. 11, fig. 10. Records — NEW ZEALAND: N. Z. Geol. Surv. loc. 1340, tuffaceous arenaceous mudstone, 2000'-3000' above the Wai- kokopu sandstone, 0.35 mi. up north-flowing nameless stream from road bridge, block 12, Ngatapa S.D. (base of Mapiri series, upper Miocene). Types — The unique holotype is in the New Zea- land Geological Survey, Wellington. I [23 - 134] May 15, 1967 INDO-PACIFIC MOLLUSCA, vol. 1, no. 7 Taranis 425 Genus Taranis Jeffreys, 1870 Type: Trophon morchi Malm, 1863 A very widely distributed genus of miniature Bathytoma- like shells, but with a much shallower sinus, the apex of which is on the peripheral keel. Angles of approach of sinus unequal, being steep and straight above but protractively arcuate below. Protoconch paucispiral, of barely two whorls, papillate, slightly globose, apparently smooth, but under high magnification there is a dense sculpture of minute stippled lirae. Pillar abruptly twisted at the commencement of a short shallowly notched anterior canal. Post-nuclear sculpture clathrate; dis- tant keels or strong spiral cords, crossed by crisp lamellate axials. There is no operculum, according to Jeffreys, in his original description of the genus. The type species of Taranis (pi. 308, fig. 1) has a wide distribution in moderately deep water from Norway to the Mediterranean and the Atlantic coast of the United States to the Gulf of Mexico. The Australasian genus Fenestrosyrinx is identi- cal with Taranis both in general appearance and also in the form and sculpture of the protoconch. Another apparent synonym is the genus Alio (from a manuscript of Jousseaume), and a substitute name for it, Feliciella, both published in the same paper by Lamy ( 1934, l.c. ) and based upon a small Red Sea shell. In addition to the species dealt with here, other members of the genus are: alexanclrina Sturany, 1896, from off Alexandria, Egypt; cirrata Brugnone, 1862, Pleistocene of Sicily and elsewhere in Europe; strongi Arnold, 1903, and inculta Moody, 1916, both from the Pleistocene of California. The Recent range of Taranis is now known to cover the North Atlantic, the Mediterranean and the Red Sea, and extends also to Hongkong, Japan, Queensland, eastern Australia, South Australia and New Zealand. Fossil occurrences are known from the Pleistocene of England, Sicily, New Zealand and California, and the Pliocene of Italy. Probably many other species, both Recent and fossil, will later be recognized as members of this wide rang- ing genus. Although Taranis resembles a Bathytotna or Mi- cantapex, in miniature, there is probably no close relationship. The genus is referred to the subfamily Turrinae solely because of the peripheral position of the sinus, but this feature is not an infallible guide to the Turrinae, as evidenced by Bathytoma and Micantapex, both of which have a different style of radula. Synonymy — 1870 Taranis Jeffreys, Ann. Mag. Nat. Hist., Ser. 4, 5, p. 447. Type by monotypy: Trophon morchi Malm, 1863. 1926 Fenestrosyrinx Finlay, Trans. N. Z. Inst., 56, p. 254. Type by original designation: Turris nexilis bicari- natus Suter, 1915. 1934 Alio (Jousseaume ms.) Lamy, Journ. de Conch., 78, pp. 67-71. Type by original designation: Alio alio (Jousseaume ms.) Lamy, 1934. 1934 Feliciella Lamy, Journ. de Conch., 78, p. 67 ( unneces- sary name for Alio alio). Type by original designa- nation: Feliciella jousseaumei Lamy, 1934. Plate 308. Taranis alio (Jousseaume), 4 mm., Djibouti, Red Sea (from Lamy, Journ. de Conch., 78, p. 67, text fig.). [23 - 145] 426 Taranis A. W. B. Powell Turridae Key to species of Taranis A. Periphery angulate, but not flanged Axials obsolete, fine growth-lines at most Periphery sharply angulate Keels strong, two on spire, two more on base imporcata (Dell) Periphery weakly angulate Keels suppressed, cords moderate, subequal spirulata (Dell) Axial and spiral sculpture subequal, dense, fenestrate Shell wide Peripheral angle medial maiji (Verco) Shell narrow Peripheral angle at about two thirds whorl height Sculpture crisply fenestrate benthicola (Dell) Sculpture indistinct, fenestrate turritispira (E. A. Smith) B. Periphery angulate but only weakly flanged Shell wide Subsidiary spiral cords weaker than peripheral one ticaonica Powell Shell narrow Subsidiary spiral cords weaker than peripheral one bicarinata (Suter) Subsidiary spiral cords almost as strong as peripheral g ratiosa (Suter) C. Periphery a projecting heavy flange Peripheral flange crenulated by axial lamellae Shoulder lirate Subsidiary spiral between periphery and lower suture Axials about 16 per whorl percarinata Powell Axials about 22 per whorl . . nexilis nexilis ( Hutton ) Shoulder without lirae nexilis recens ( Fleming ) Peripheral flange undulated by distant broad low axials Subsidiary spirals sparse and weak . . . alio (Jousseaume) Plate 309. Fig. 1, Taranis morchi Malm, West Norway. 3 mm. Fig. 2, T. ticaonica Powell new species, 2.3 mm., Ticao Island, 226 fms., Philippines, holotype. Fig. 3, T . turritispira (E. A. Smith), 6 mm., Japan, co-type in British Museum. Fig. 4, T. bicarinata (Suter), 23 fms. off Ahipara, New Zealand. Fig. 5, T. nexilis (Hutton), Castlecliff, New Zealand. Pleistocene. Fig. 6, T. percarinata Powell new spe- cies, 3 mm., east of Masbate, 135 fms., Philippines, holotype. [23 - 146] May 15, 1967 INDO-PACIFIC MOLLUSCA, vol. 1, no. 7 Taranis 427 Taranis alio Jousseaume, 1934 (PI. 308) Range — Known only from Djibouti, Red Sea. Remarks — This species appears insufficiently dis- tinct from the type of Taranis to warrant even sub- generic segregation. Jousseaume’s genus Alio and species alio were described and published post- humously. Authorship by Jousseaume is clearly in- tended by the editor, Ed. Lamy, particularly since the names appear in the index to that particular volume of the Journal de Conchyliologie as belong- ing to Jousseaume. Lamy added additional remarks deriding the name Alio alio, and illegally proposed the substitute names Feliciella jousseaumei. I con- sider them synonyms. Description — Shell very small, 4 mm. in height, rather broadly ovate-biconic, with a moderately tall spire and truncated body-whorl. Protoconch eroded but evidently paucispiral. Spire-whorls strongly angulated just above the middle by an irregularly nodulose raised keel. Four or five rather indistinct spiral cords on the body-whorl, from below the peripheral carina and over the base. Axial sculp- ture of weak, rather irregular and distant lamellae. Sinus broadly open, V-shaped, very shallow, its apex on the peripheral carina. The anterior canal is very short, its shallowly-notched extremity being very oblique to the axis. Measurements (mm.) — height width 4.0 3.0 Synonymy — 1934 Alio alio Jousseaume, Journ. de Conch., 78, p. 67. 1934 Feliciella jousseaumei Lamy, Journ. de Conch., 78, p. 67 (unnecessary substitute name for Alio alio Jous- seaume ) . Records — Known only from the type locality, Djibouti, French Somalia. Taranis percarinata Powell, new species (PI. 309, fig. 6) Range — Philippines in 80 to 135 fathoms. Remarks — This species, with its prominent sharp peripheral keel, more closely resembles the New Zealand nexilis (Hutton, 1885) than it does the European type of the genus, morchi Malm, which has semi-inflated weakly carinated whorls. From the New Zealand species the Philippine shells differ in having a still more prominent peripheral keel and fewer, stronger axial lamellae. Description — Shell very small, 3 mm. in height. with prominently carinated whorls, overridden by lamellate axials and with a truncated anterior end. Spire 1/2 times height of aperture plus canal. Whorls 5/2, including a papillate protoconch of 1/2 whorls, densely sculptured with minute, stippled lirae. Post-nuclear whorls strongly and bluntly carinated below, and a second carina, almost as strong, is emergent over the last half-whorl, followed by two much weaker spirals on the base and five oblique lirae on the anterior end. Prominent lamellate axials, about 16 per whorl, cross the whole surface; these are retractively slanting and very broadly V-shaped over the spire- whorls, but are recurved arcuately over the base. They crenulate the main carinae and render the secondary spirals gemmate. The sinus is weak, very broadly V-shaped, as defined by the axial lamellae and has its apex on the peripheral carina. The aperture is obliquely subquadrate and is produced below into a very short but strongly flexed, unnotched anterior canal. Colour dull-white. Occurring in this species is a form which differs in having a sharp, relatively strong, subsutural spiral and another on the body-whorl between the two main carinae. However, there is insufficient material to determine the status of this form. Measurements (mm.) — height width 3.0 1.5 Types — The holotype is in the United States National Museum, Washington (USNM 281739L). Records — PHILIPPINES: south of Adyagan Island, east Masbate, 135 fathoms (holotype; Albatross Sta. 5392). Off Destacado Island, east Masbate, 80 fathoms (Albatross Sta. 5213). Taranis ticaonica Powell, new species (PI. 309, fig. 2) Range — Off Ticao Island, Philippines, in 226 fathoms. Remarks — This species is very similar to the European morchi Malm, both in shape and in sculp- ture. In fact, the only differences in the Philippine shell are a slightly more pronounced peripheral carina, more numerous basal spirals, fewer and slightly stronger axial lamellae, and a thicker, less excavated, anterior end. Description — Shell very small, 2.3 mm. in height, white, biconic, with a tabulated spire, slightly taller than the height of the aperture plus the canal. Whorls 5, including a subglobose protoconch of 1/2 whorls, delicately sculptured with closely spaced, minute, stippled lines. Post-nuclear whorls medially [23-151] 428 Taranis A. W. B. Powell Turridae strongly carinate. Suture channelled and submar- gined by a moderately strong spiral cord; two slightly weaker cords between the subsutural cord and the periphery, and 1 or 2 between the periph- ery and the lower suture; about 8 cords on the body-whorl, plus 5 weaker and more closely spaced spirals on the anterior end. The whole crossed by crisp, axial, chevron-shaped lamellae, which follow the outline of the sinus which is a widely open V, with its apex at the peripheral carina. Measurements (mm.) — height width 2.3 1.3 holotype Types — The holotype is from Albatross Sta. 5388 and is in the United States National Museum, Washington (USNM, no. 285237). Taranis turritispira (E. A. Smith, 1882) (PI. 309, fig. 3) Range — Japan. Description — Shell small, of light build, 6 mm. in height, elongately ovate, with rounded whorls, weakly angulated at the shoulder. Height of spire slightly greater than height of aperture plus canal. Sculpture clathrate, not very prominent, the spirals dominant, and the axials tending to become sub- obsolete over the last whorl. Three spiral cords are on the upper whorls, with a fourth emergent over the last whorl. Protoconch paucispiral and with about two whorls (not examined for micro-sculp- ture). Sinus and other apertural features typical. Measurements (mm.) — height width 6.0 2.0 Synonymy — 1882 Pleurotoma (Taranis?) turritispira E. A. Smith, Ann. Mag. Nat. Hist., Ser. 5, 10 (not figured), p. 306. 1952 Taranis turritispira (Smith), Kuroda and Habe, Check List and Bibliogr. Rec. Mar. Moll. Japan, p. 88. T ypes — Three co-types in the British Museum (Natural History). The specimen illustrated here is the lectotype. The type locality is Japan. [23 - 152] May 15, 1967 INDO-PACIFIC MOLLUSCA, vol. 1, no. 7 Taranis 429 RECENT AND PLEISTOCENE SPECIES FROM SOUTHERN AUSTRALIA AND NEW ZEALAND Taranis benthicola (Dell, 1956) Range — New Zealand, off Eastern Otago, 300 fathoms (type locality); Chatham Rise, 200 fathoms, and Pitt Island, 155 fathoms. Synonymy — 1956 Fenestrosyrinx benthicola Dell, Domin. Mus. Bull., No. 18, p. 141, fig. 197. Types — Holotype and paratypes in the Dominion Museum, Wellington. Taranis bicarinata (Suter, 1915) (PI. 309, fig. 4) Range — New Zealand, Wanganui ( Castlecliffian Pleistocene) ( bicarinata ); Recent, off Hen and Chicken Islands (thomsoni); off Ahipara, 23 fathoms. Synonymy — 1885 Clatliurella nexilis Hutton (partim), Trans. N. Z. Inst., 17, p. 317 ( = 5 of Hutton’s syntypes of nexilis). 1915 Tunis (Hemipleurotoma) nexilis bicarinatus Suter, N. Z. Geol. Surv. Pal. Bull., No. 3, p. 34. 1919 Leucosyrinx thomsoni Mestayer, Trans. N. Z. Inst., 51, p. 133, pi. 8, fig. 5. 1942 Fenestrosyrinx nexilis bicarinata (Suter), Powell, Bull. No. 2, Auck. Inst. Mus., p. 57. Types — The holotype of bicarinata is in the Canterbury Museum and the holotype of thomsoni is in the Dominion Museum, Wellington. Taranis gratiosa (Suter, 1908) Range — New Zealand, Port Pegasus, 18 fathoms, Stewart Island (type locality) to Dusky Sound. Synonymy — 1908 Bathytoma gratiosa Suter, Proc. Malac. Soc., 8, p. 186, pi. 7, fig. 19. 1942 Fenestrosyrinx gratiosa (Suter), Powell, Bull. No. 2, Auck. Inst. Mus., p. 57. Types — The holotype is in the New Zealand Geological Survey, Wellington. Taranis imporcata (Dell, 1962) Range — New Zealand, off Taiaroa Head, 300 fathoms. Synonymy — 1962 Fenestrosyrinx imporcata Dell, Rec. Domin. Mus., 4(15), p. 73, fig. 4. Types — Holotype and paratypes in the Dominion Museum, Wellington. Taranis mayi (Verco, 1909) Range — South Australia, 35 miles south-west of Neptune Islands, 104 fathoms. Remarks — This very small shell, 4.6 mm. in height, is biconical, with medially sharply angled whorls, sculptured with a dense clathrate pattern of numerous thin but sharply raised spirals, crossed by equally strong and numerous axials. Protoconch typical, of two convex apparently smooth but microscopically finely spirally lirate and intersti- tially punctate. Known only from the type locality. Synonymy — 1909 Hemipleurotoma mayi Verco, Trans. Boyal. Soc. S. Aust, 33, p. 295, pi. 25, fig. 2. Taranis nexilis subspecies nexilis (Hutton, 1885) (PI. 309, fig. 5) Range — New Zealand, Wanganui ( Castlecliffian Pleistocene ) . Synonymy — 1885 Clathurella? nexilis Hutton, Trans. N. Z. Inst., 17, p. 317, pi. 18, fig. 9. 1942 Fenestrosyrinx nexilis (Hutton), Powell, Bull. No. 2, Auck. Inst. Mus., p. 57. Types — The holotype is in the Canterbury Mu- seum, Christchurch, New Zealand. Taranis nexilis subspecies recens (Fleming, 1948) Range — New Zealand, Dusky Sound, 22 fathoms (type locality), and Edwardson Sound, 58 fathoms. Synonymy — 1948 Fenestrosyrinx nexilis recens Fleming, Trans. Royal Soc. N. Z., 77, p. 91, pi. 8, fig. 7. Types — The holotype is in the New Zealand Geological Survey, Wellington. [23 - 163] 430 Taranis A. W. B. Powell Turridae Taranis spirulata (Dell, 1962) Range — New Zealand, off Taiaroa Head, 300 fathoms. Synonymy — 1962 Fenestrosyrinx spirulata Dell, Rec. Domin. Mus., 4(15), p. 73, fig. 5. T types — Holotype and paratypes in the Dominion Museum, Wellington. [23 - 164] May 15, 1967 INDO-PACIFIC MOLLUSCA, vol. 1, no. 7 Micropleurotoma 431 Genus Micropleurotoma Thiele, 1929 Type: Pleurotoma spirotropoides Thiele, 1925 The type species of this genus is a small, rather featureless shell (4.5 mm.), from deep water off South Africa. The protoconch is relatively large, globose, dome- shaped, smooth and paucispiral. The adult whorls are smooth except for a prominent rounded smooth peripheral keel. The sinus is shallow, its apex on the peripheral keel. Anterior canal short, spout-like and unnotched. The genus may be near to Taranis, which differs in the protoconch being microscopically granulose- lirate and in having the anterior canal shallowly notched. I have not seen material relevant to this genus, so have no remarks to add to the rather meagre original description. An alleged second species of the genus, M. ashi- yaensis Shuto and Uedo, 1963, from the Ashiya group, upper Oligocene of Japan is very doubtfully a member of the genus. The incomplete holotype is very much larger (23.4 mm.) than spirotropoides, and in addition to the smooth rounded peripheral carina, has a dense surface pattern of spiral lira- tions. Except for the lack of peripheral nodes, the Japanese species shows possible affinity with Ln- cerapex. Synonymy — 1929 Micropleurotoma Thiele, Handbuch der systematischen Weichtierkunde, 1, p. 362. Type by original desig- nation: Pleurotoma spirotropoides Thiele, 1925. Micropleurotoma spirotropoides (Thiele, 1925) (PI. 310, fig. 1) Range — Off South Africa, 126 and 2750 metres. Measurements (mm.) — height width 4.5 2.3 Synonymy — 1925 Pleurotoma spirotropoides Thiele, Wissenschaft Ergebn. Deutschen Tiefsee-Exped., 17, Gast. 2, p. 213( 179), pi. 23, fig. 18. 1929 Epideira (Micropleurotoma) spirotropoides (Thiele), Handb. system. Weichtierkunde, 1, p. 362. Records — SOUTH AFRICA: 35° 19' S; 20° 12' E, 126 metres. Agulhas Bank, 35° 32.8' S; 18° 20.1' E, 2750 metres. ? Micropleurotoma ashiyaensis Shuto and Ueda, 1983 (PI. 310, figs. 2, 3) Range — Japan, coastal cliff north of Taya, Ashiya machi, Onga gun, Fukuoka Prefecture (Ashiya group, Nishisonogian, upper Oligocene). Remarks — Very doubtfully a member of the genus. Measurements (mm.) — height width 23.4 mm. (incomplete) 13.5 Synonymy — 1963 Micropleurotoma ashiyaensis Shuto and Ueda, Jap. Joum. Geol. and Geogr., 34(1), p. 2, pi. 1, figs. Types — The holotype is in the Department of Geology, Kyushu University. Plate 310. Fig. 1, Micropleurotoma spirotropoides (Thiele), 4.5 mm., South Africa, 2750 metres (from Thiele, 1925, Gast. deutsch. Tiefsee-Exped., 17, pi. 23, fig. 18). Figs, 2, 3, PMicropleurotoma ashiyaensis Shuto and Ueda, 23.4 mm., Japan, upper Oligocene (from Shuto and Ueda, 1963, Jap. Journ. Geol. and Geogr., 34(1), pi. 1, figs. 1, 2). [23 - 175] 432 Micropleurotoma A. W. B. Powell Turridae [These occasional blank areas occur genera and subgenera to permit the of new material and future sections proper systematic sequence.] between insertion in their [23 - 176] May 15, 1967 INDO-PACIFIC MOLLUSCA, vol. 1, no. 7 Turrinae 433 ADDITIONS AND CORRECTIONS TO GEMMULA AND XENUROTURRIS [looseleaf subscribers should place the next 12 pages in their proper places by following the pagination at the bottom of the pages.] [22 - 662a] 434 Turrinae A. W. B. Powell Turridae Figs. Plate 175 (opposite page) Corrections made on May 15, 1967 (see stars). 1, 6 Gemmula ( Unedogemmula ) 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 ( Unedogemmula ) deshay esii (Dou- met). Hongkong (7), Japan (8) (p. 22-762). 9 Lophiotoma (Lophioturris) indica (Roding). Philippines (p. 22-931). 10, 1 1 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 Powell. Hawaiian Is- lands (p. 22-964). 16 Lophiotoma ( Lophioturris ) polytropa (Hel- bling). Moluccas. 17,18 Xenuroturris millepunctata (Sowerby). New Caledonia (p. 22-963). 19,20 Xenuroturris cingulifera (Lamarck). Japan (p. 22-962). 21, 22 Xenuroturris castanella Powell. Hawaiian Is- lands (p. 22-964). ^ (all 2/3 natural size) [22 - 662b] * * May 15, 1967 INDO-PACIFIC MOLLUSCA, vol. 1, no. 7 uda 435 Gemmula interpolata Powell, new species (PI. 311, figs. 1-3) Range — Known only from the Hawaiian Islands; 10 to 355 fathoms. Remarks — This one- to two-inch Gemmula ap- pears to be allied to G. gilchristi ( Sowerby ) of the Indian Ocean and Japan [p. 22-701]. This Ha- waiian species is also broadly fusiform, but has a more regular outline due to the subsutural fold being more prominent than the peripheral carina. The colours consist of sparse but bright brown maculations. Description — Shell moderately broadly fusiform, 25 to 52 mm. (about 1 to 2 inches) in height, with a tall narrow spire with an angle of 34 to 36 de- grees, and a long, almost straight, unnotched ante- rior canal. Spire height a little less than that of the aperture plus the canal. Protoconch narrowly coni- cal, of 3M dark-brown whorls, smooth at first but strongly and closely axially costate over the last 1/2 whorls. Post-nuclear whorls 8. Spire- whorls sculp- tured firstly with a prominent, narrowly-crested, smooth, subsutural fold, followed by a straight, steeply descending shoulder slope, which bears three narrow, crisp, smooth spiral threads; then ap- pears the low-set peripheral carina, which is com- posed of two linear-spaced spiral cords, which are rendered cog-like by vertically fused nodes. Below the carina there is one smooth cord just emergent at the lower suture. On the body-whorl, below the peripheral carina, there are 8 or 9 distant, smooth, prominent cords, a single thread in each interspace, and 12 to 14 closely-spaced threads on the anterior end. Peripheral nodes 26 to 28 per whorl. Sinus deep, U-shaped, its apex occupying the whole width of the peripheral keel. Colour ivory-white, sparsely maculated in dark reddish brown under a thin pale-buff periostracum. The maculations are large and squarish, and occurring in a peripheral series regularly alternating with the nodes. Other spots are sparsely disposed, both on the subsutural fold and on the primary spirals. Inside the outer lip there are distinct spiral lirations. Average-sized specimens are about 25 mm. in height but occa- sional ones from deep water reach a height of over 50 mm. In such specimens the maculations are paler and there is a tendency for the peripheral keel to become less prominent over the later whorls. Measurements (mm.) — height width 52.4 16.1 paratype 26.5 9.2 holotype; off Laysan Id. Types — The holotype, from Laysan Island (USNM 190417) and other paratypes are in the United States National Museum, Washington. Records — HAWAII: near Laysan Id., 59-70 fins., bottom temperature 70° F. (Albatross Sta. 3940; USNM 190417) (holotype); Near Laysan Id., 130-148 fms., bottom tem- perature 63° F (Albatross Sta. 3937; USNM 335269); Oahu, off Waikiki, 100-190 fms.; Keehi Lagoon, 20-30 fms.; off Ala Aesava, 10-40 fms. (C. M. Burgess); off N.E. coast of Maui Id., 143-178 fms., bottom temperature 60.8° F. (Albatross Sta. 4079; USNM 335271); Pailolo Channel, Maui, 128 fms., bottom temperature 62.5° F. (Albatross Sta. 3857; USNM 206003); Kaiwi Channel, 350-355 fms., bottom temperature 41.6° F. (Albatross Sta. 4107; USNM 338272); off S. coast of Molokai Id., 169-182 fms., bottom temperature 55° F. (Albatross Sta. 3835; USNM 335275); near Ranai Id., 233-240 fms., bottom temperature 48.5° F. (Albatross Sta. 3982; USNM 335270). Plate 311. Gemmula interpolata Powell, new species. Fig. 1, holotype, USNM 190417, 59-70 fms., off Laysan Island, Hawaii; 26.5 mm. Fig. 2, paratype, 143-178 fms., off Maui Island, Hawaii; C. M. Burgess; 52.4 mm. Fig. 3, paratype, 20-30 fms., Keehi Lagoon, Oahu Island, Hawaii; C. M. Burgess; 25.4 mm. [22 - 703] 436 Gemmula A. W. B. Powell Turridae Gemmula pseudomonilifera Powell, new species (PI. 312) Range — Known only from the Hawaiian Islands; 20 to 67 fathoms. Remarks — This attractive, ?4-inch turrid resem- bles monilifera (Pease, 1860) from Hawaii and Fiji [p. 22-701], but has a buff colored shell with light- brown, peripheral nodes in contrast to monilifera which is always broadly zoned in brown and with a white anterior end. G. pseudomonilifera also has a more gradually contracted base, more widely spaced basal spirals, and a shorter anterior canal. This new species also shows close affinities to hom- broni (Hedley, 1922) from the southwest Pacific [p. 22-734], A specimen from 253 fathoms off Maui Island exhibits a tubular labial outgrowth of a basal spiral, as is sometimes found in specimens of Pinguigemmula. Description — Shell small (17 mm.), narrowly fusiform, with a tall spire and a moderately long, very slightly flexed anterior canal. Spire-height slightly greater than half the length of the entire shell. Protoconch conical and with about 4 whorls, the first two smooth, the remainder closely and strongly axially costate. Post-nuclear whorls 6. Spire- whorls sculptured firstly with a prominent sharply carinated subsutural fold, followed by a deep shoulder concavity bearing two fine smooth spiral threads, then by the low-set peripheral keel, which is composed of two gemmate cords. Over the last two whorls, between the peripheral keel and the lower suture, a strong plain spiral cord becomes emergent. On the base and neck the primary spirals are few, strong, plain and wide-spaced, with occa- sional fine interstitial threads, after which there are closely spaced, but indistinct, spiral lirae on the an- terior end. Sinus moderately deep; rather broadly U-shaped, its apex on the peripheral keel. Colour buff, with the protoconch and peripheral nodes tinged with light-brown and most of the body- whorl diffused with the same colour. Measurements (mm.) — height width 17.7 6.3 holotype 16.0 5.9 paratype Records — HAWAII: Oahu, Keehi Lagoon, 20-30 fms. (type locality); off Waikiki, 100-190 fms. (P. Burgess); off N.E. coast of Maui Id., 253-267 fms., bottom temperature 46° F. (Albatross Sta. 4084; USNM 338283); S. coast of Molokai Id., 64-60 fms., bottom temperature 71.5° F. (Alba- tross Sta. 3846; USNM 338282); near Ranai Id., 233-240 fms., bottom temperature 48.5° F. (Albatross Sta. 3982; USNM 338284). Plate 312. Gemmula pseudomonilifera Powell, new species. Holotype, Auckland Museum. 20-30 fms., Keehi Lagoon, Oahu Island, Hawaii; 17.7 mm. [22 - 704] May 15, 1967 INDO-PACIFIC MOLLUSCA, vol. 1, no. 7 Gemmula 437 Gemmula congener subspecies unilineata Powell, new subspecies (PL 313) Range — Known only from the Hawaiian Islands; 100 to 312 fathoms. Remarks — This subspecies differs from typical congener as well as from the subspecies cosmoi and diomedea, in that the spiral lirations of the shoulder- slope are knotted or beaded, not plain. In the strong development of the subsutural fold and peripheral carina, i inilirata is nearest to the typical subspecies, but the shoulder sulcus is wider, bearing three spirals. In typical congener the shoulder sulcus is deep and narrow, with only one or two smooth spirals. In cosmoi and diomedea there is a very wide shoulder slope that bears 4 to 6 smooth spiral threads. The only other subspecies that has a brown-banded subsutural fold is cosmoi, but it is unicarinate and smooth, not tricarinate and beaded. Description — Shell rather broadly fusiform, with a tall spire, with an angle of 34 or 35 degrees, and a moderately long, straight and unnotched anterior canal. Spire height a little more than that of the aperture plus the canal. Protoconch narrowly coni- cal of about three whorls, the first two smooth and the last closely axially costate. Post-nuclear whorls 8. Spire-whorls sculptured firstly with a massive subsutural fold, which is composed of three gem- mate spirals, the middle one strongest, followed by a moderately wide shoulder slope, which is sculp- tured with three knotted to gemmate sub-cords; then the low-set strong peripheral carina, which is composed of two closely-spaced cords, studded with about 32 closely-spaced series of nodes which are vertically fused; from the carina to the lower suture there is a single strong smooth cord. From the carina to the anterior end there are about six- teen cords, which are strong above, then gradually diminishing, the uppermost undulating, the median ones more or less gemmate, the lower ones smooth, with a thread in most interspaces. Sinus moderately deep, with a rather broadly V-shaped entrance, its U-shaped apex occupying the full width of the pe- ripheral carina. Interior of outer lip finely lirate. Ground colour ivory-white under a pale straw- coloured periostracum; subsutural fold dark red- dish brown, and pale reddish brown between the peripheral nodes. Measurements (mm.) — height width 31.0 10.7 holotype 28.0 10.2 off Diamond Head, 240-260 fms. Types — The holotype is in the B. P. Bishop Mu- seum, Honolulu. The type locality is 200 fathoms, mud bottom, off Waikiki, Oahu Island, Hawaii. Pele Expedition, June 13, 1964, C. Weaver, collec- tor. Paratypes are in the U. S. National Museum from other Albatross stations ( see records ) . Records — HAWAII: 100-190 fms., off Waikiki; 240-260 fms., off Diamond Head (both Oahu Island; Pat Burgess); 147-198 fms., bottom temperature 49° F. (Albatross Sta. 4045; USNM 338280); near Kauai Id., 309-257 fms., 43.7° F. (Albatross Sta. 4131; USNM 173042); near Kauai Id., 283-309 fms., 46.1° F. (Albatross Sta. 4130; USNM 338273); near Kauai Id., 257-312 fms., 46.8° F. (Albatross Sta. 4132; USNM 338274); south coast of Molokai Id., 238- 255 fms., 48° F. (Albatross Sta. 3836; USNM 338278); south coast of Oahu Id., 211 fms., 47.7° F. (Albatross Sta. 3810; USNM 338277). (Most of the Albatross specimens are dead bleached shells, but one from Sta. 4045 exhibits the characteristic brown subsutural band. All temperatures were taken at the bottom. ) Plate 313. Gemmula congener new subspecies unilineata Powell. Holotype, B. P. Bishop Museum. 200 fms., off Wai- kiki, Oahu Island, Hawaii; 31.0 mm. [22 - 716a] 438 Gemmula A. W. B. Powell Turridae [These occasional blank areas occur genera and subgenera to permit the of new material and future sections proper systematic sequence.] between insertion in their [22 - 716b] May 15, 1967 INDO-PACIFIC MOLLUSCA, vol. 1, no. 7 Gemmula 439 Gemmula hombroni (Hedley, 1922) (PI. 200, fig. 4; pi. 201, fig. 8; pi. 314) Remarks — I add new locality records and new illustrations of unusual specimens with a double sinus on the outer lip. Three specimens of this species taken at Batangas Bay, Philippines, and forwarded for inspection by Mr. Donald Dan of Manila, exhibit an unusual fea- ture, in that there is a second, properly formed sinus, situated in the lower part of the outer lip. This sinus is of the same character as the anal sinus but is not quite so deep; it is, however, slightly rim- margined and is certainly not just an accidental cleft due to injury. This extra sinus does not seem to be quite com- parable with the irregular flutings often found in Gemmula and the related genera Pinguigemmuhi and Ptychosyrinx. In all such cases the flutings have a projecting rounded or spout-like termina- tion, and in most instances they are filled with callus. The solid nature of most of these flutings negates the suggestion made by MacNeil (1960), that they may represent a response to an oxygen- poor environment into which one or more incur- rent siphons are developed. Unfortunately there are no animals available from specimens with either the flutings or the extra sinus. Only an examination of these animals associated with these shell abnor- malities would elucidate the problem. New records — PHILIPPINES: Batangas Bay, Luzon Is- land (Donald Dan, Auck. Mus.). FIJI: dredged in 80 to 90 ft., muddy sand, Momi Bay, Viti Levu Id. (Bill Jennings, 1962, ANSP). Plate 314. Gemmula hombroni Hedley, Batangas Bay, Philippines; showing a second sinus in the outer lip. 24.0 mm. Gemmula tessellata Powell, new species (PI. 315) Range — Known only from the Hawaiian Islands. Remarks — This small, attractive species most closely resembles the more slender Gemmula amabi- lis ( Weinkauff ) from the Bed Sea, from which it is distinguished by its stouter outline, tuberculate sub- sutural fold and its shorter, straighter anterior canal. This species belongs to Gemmula sensu stricto. Description — Shell less than an inch in size, fusi- form, with a tall spire which is slightly more than half the length of the entire shell. Protoconch tall, narrowly conical, of 4/2 whorls, the tip smooth, the remaining whorls sculptured with closely spaced stout axials, the whole crossed by delicate spiral threads, more prominent on the axials than in the interspaces. Post-nuclear whorls 6. Spire-whorls bi- carinate, sculptured with a prominent, narrowly crested and tuberculate subsutural fold, which is separated by a narrow, deep shoulder sulcus from the massive peripheral keel, composed of two al- most coalescent cords and bearing prominent, verti- cally fused, cog-like nodes. Below the low-set pe- ripheral keel there is a single smooth cord emergent Plate 315. Gemmula tessellata Powell, new species. Holo- type, Auckland Museum. 100-120 fms., off Waikiki, Oahu Island, Hawaii. 15.8 mm. [22 - 734a] 440 Gemmula A. W. B. Powell Turridae at the lower suture of the penultimate whorl, and a second one appears over the last half-whorl. Body- whorl with rather widely spaced, flat-topped cords above, four weaker cords, the upper two with inter- mediate threads, over the neck, and closely spaced fine threads over the anterior end. Sinus moder- ately deep, U-shaped, its apex occupying the whole width of the peripheral carina. Interior of outer lip strongly lirate. Colour ivory white with the proto- conch tinted light reddish brown, and a regular tessellation in the same colour over the remainder of the shell. The tessellations alternate with the tubercles on the subsutural fold, and are regularly spaced on the primary spirals. Inner-lip callus and interior of the aperture white. Measurements (mm.) — height width whorls 15.7 9.5 10.5 holotype Types — The type locality is 100 to 120 fathoms, in mud and sand, off Waikiki, Oahu Island, Hawaii. Collected by Dr. Pat Burgess. The holotype, and only known specimen, is deposited in the Auckland Museum and Institute, New Zealand. Gemmula microscelida (Dali, 1895) (PI. 316, figs. 1-3) Range — Hawaiian Islands, deep water from 253 to 528 fathoms. Remarks — This deep water, rather common spe- cies bears some resemblance to g raeffei (Wein- kauff) from the southwest Pacific in its form and sculpture, but the canal is much shorter and is de- cidedly twisted. Judging from the poor specimens selected as holotype and paratype, the full range of material listed below could not have been avail- able to Dali when he wrote his 1895 paper. A much better specimen from Albatross station 4083 is now figured along with the type specimens. Description — Shell about an inch in size, rather broadly fusiform, with the spire being more than half the entire length of the shell. Protoconch usu- ally missing, but consists of about 4 narrowly-coni- cal whorls, the last two of which are closely and axially costate. Post-nuclear whorls about 7. Sculp- ture of spire whorls consisting firstly of a rather prominent, rounded, smooth subsutural cord, then a rather wide, moderately concave shoulder slope, bearing 1-3 weak spiral threads, followed by the low-set prominent peripheral keel, which is com- posed of two linear-spaced cords, rendered cog-like by numerous vertically fused nodes. These nodes frequently extend a little beyond the keel, both above and below, where they coalesce with the axial growth lines. The nodes number between 20 and 22 per whorl. Between the periphery and the lower suture there is a single smooth spiral cord. On the body whorl, from the periphery downward, there are about 11 narrow smooth cords, extending to the bottom of the neck, after which there are weak indefinite closely-spaced threads over the an- terior end. Surface smooth except for numerous, fine, crisp axial growth lines; surface covered by a pale to moderately dark olivaceous periostracum. Sinus rather broadly V-shaped at its entrance, the apex U-shaped and occupying the full width of the peripheral carina. Measurements (mm.) — height width 22.0 8.5 holotype 22.2 10.0 paratype 23.5 9.0 paratype Types — The holotype is in USNM 127122. The type locality is 351 fathoms, sand bottom, off the Hawaiian Islands, Albatross station 3475. Synonymy — 1895 Pleurotoma microscelida Dali, Proc. U. S. Nat. Mus., 17, p. 677 (not figured). Records — (All off the Hawaiian Islands, all dredged by the Albatross, and all lots in the United States National Mu- seum. All temperatures mentioned were taken at the bot- tom.) Station 4041, 382-253 fms., 41.6° F., west coast of Hawaii Id.; sta. 4131, 309-257 fms., 43.7° F., off Kauai Id.; sta. 4083, 238-253 fms., off N.E. Maui Id.; sta. 3992, 528 fms., 39.6° F., off Ranai Id.; sta. 3883, 277-284 fms., 45.2° F., Pailolo Channel; sta. 3839, 259-266 fms., 46.3° F., off south Molokai Id. Also from Albatross stations 4130. 4133, 4090, 3866, 3865, 3867, and 4027. Plate 316. Gemmula microscelida (Dali, 1895). Fig. 1, holotype, USNM 127122, 351 fms., off the Hawaiian Islands; 22.0 mm. Fig. 2, paratype, Albatross station 3475, off Ha- waiian Islands; 22.3 mm. Fig. 3, 238-253 fms., off N.E. coast of Maui Island, Albatross station 4083, USNM 173038; 23.5 mm. [22 - 734b] May 15, 1967 INDO-PACIFIC MOLLUSCA, vol. 1, no. 7 [replacing vol. 1, no. 5, pp. 323-324; see stars] Xenuroturris 441 Plate 253. Geographical distribution of Viridoturris corona Laseron and members of the genus Xenuroturris. Xenuroturris cingulifera subspecies erythraea (Weinkauff, 1875) (PI. 252, fig. 1) Range — Red Sea. Remarks — It is likely that the Red Sea popula- tions may represent a good regional subspecies dif- fering from cingulifera typical mainly in the much stronger and fewer spiral cords and a characteristic purple staining in adults of the anterior canal and pillar. Unfortunately, I have seen insufficient ma- terial to determine if the above mentioned charac- ters are constant and if cingulifera typical is absent from the area. Description — Adult shell 23 to 25 mm. ( about 1 inch) in height. Whorls about 8, plus protoconch (missing in material examined). Spire twice the height of the aperture plus canal, angle 18° to 20°. Spire whorls with three heavy spiral keels and a fourth emergent over the latter part of the body whorl. Spiral cords on the base, 4 or 5 and strong, interspaces each with 3 or 4 crisp, spiral threads and 4 or 5 rounded cords on the fasciole. In two of the three specimens I have examined all the keels including the sinus one are narrowly arched but in the third specimen the sinus rib is composed, as in cingulifera, of two closely spaced cords. Col- our pattern of small brown speckles on the primary cords and large regularly spaced, rectangular patches of dark-brown on the sinus rib, or paired dashes in one example. Pillar and anterior canal stained bright violet but this staining absent in the example with the bifid sinus rib. Despite these variations, the Red Sea form would appear on the scant material available to represent a regional subspecies on the basis of stronger and fewer spirals and a tendency to violet staining of the anterior end. Measurements (mm.) height width 25.0 9.0 23.5 8.5 23.5 8.0 19.0 7.5 Synonymy — 1875 Pleurotoma erythraea Weinkauff, Conch. Cab., p. 22, pi. 4, fig. 10. 1884 Pleurotoma erythraea Weinkauff, Tryon, Manual of Conch. 6, p. 166, pi. 3, fig. 24. Records — RED SEA: Massaua and Dahlack (Weinkauff) (type locality); Port Sudan (MCZ). Fossil Records — Somali coast (Pleistocene) (Abrard, 1942, p. 86, recorded and figured as cingulifera) . Port Sudan Type, Massaua Port Sudan Port Sudan Xenuroturris millepunctata (Sowerby, 1908) (Color pi. 175, figs. 13, 17, 18; pi. 252) Range — New Caledonia, Fiji, Kermadec Islands, Ryukyu Island, Japan and Australia. ^ Description — Adult shell 25 to 50 mm. ( 1 to 2 inches ) in height. Whorls 9 or 10, plus protoconch of 4 to 4M axially costate whorls, as in cingulifera. Spire about l/i times the height of the aperture plus canal, angle 22° to 24°. The differentiating charac- ters from those of cingulifera are the colour pattern of rather sparse brown speckles, minus maculations on the sinus area, the more prominent sinus ribs and a distinct angulation of the base. [22 - 963a] 442 Xenuroturris [replacement page] A. W. B. Powell Turridae Measurements (mm.) — height width 50.0 15.0 Kii, Japan 34.7 11.0 New Caledonia 25.0 9.0 New Caledonia Synonymy — 1908 Pleurotoma millepunctata Sowerby, Proc. Malac. Soc. London 8, p. 198, text fig. 1914 Pleurotoma cingulifera zonifera Bouge and Dautzen- berg, Joum. de Conchyl. 61, p. 128 (not figured). Records — JAPAN : Kii (A. W. B. Powell coll.); Ikenodan, Izu (D. Thaanum). RYUKYU ISLANDS: Gima, Kumejima Id., Okinawa (D. Thaanum). LOYALTY ISLANDS: (D. Thaanum). NEW CALEDONIA: lie Monac (type local- ity); Voh, on sea-grass tidal flat. Miss V. Orr, ANSP). KERMADEC ISLANDS: Raoul Id. (W. R. B. Oliver, Aus- tralian Mus.). FIJI ISLANDS: Namotu Id., 13-15 fms. (W. ★ Jennings). AUSTRALIA: 30 fms., off Southport, Queens- land (T. Garrard, coll. 1965). -fa Xenuroturris cerithiformis Powell, 1964 (Color pi. 175, figs. 14, 15; pi. 252) Range — Hawaiian Islands. Remarks — This subspecies has long been known from distributed material bearing Dali’s manuscript name, cerithiformis, which was quoted along with figures and a description by Tinker (1952). Tinker did not intend to describe this and several other of Dali’s manuscript species in his book. His . action does not measure up to the requirements of ^ the International Rules and Powell, 1964, not Tinker, 1952, must be the author. Tins Hawaiian species is close to millepunctata but differs consistently in having more rounded whorls, neither sharply angled on the base nor deeply excavated at the neck. Description — Adult shell 33 to 53 mm. ( 1J£ to 2 inches) in height, whorls 10 or 11, plus a multi- spiral narrowly conic protoconch of 4 to 4/2 whorls, brown, densely sculptured with slightly curved strong rounded axials crossed at right angles by weak spiral lirae. Spire tall, almost twice the height of the aperture plus canal, 24° to 27°, outlines lightly convex except for bulging subsutural fold and sinus rib. Sculpture of spue whorls consisting of a broadly-convex subsutural fold bearing 3 to 5 weak spiral threads, separated by a narrow deeply channeled concavity from the sinus rib, situated at about middle whorl height and composed of two moderately strong, rounded, spiral cords with a concavity between. Below the sinus area there are 2 to 4 crisp cords of varying strength. About 10 primary cords and occasional interstitial threads on the base from below the sinus area plus 4 cords on the weak fasciole. Sinus deep and narrow. Col- our white, rather evenly speckled with reddish brown dots and dashes; larger maculations not present. Measurements (mm.) — height 53.0 width 18.0 Pearl and Hermes Reefs, Hawaii 45.0 14.5 Kalaekiki, Hawaii 38.5 11.4 Honolulu, 6-8 fms. (type) 33.4 11.0 Honolulu, 6-8 fms. 32.0 10.2 Honolulu, 6-8 fms. Types — l hereby select the specimen in USNM 338601 as the holotype. It is figured in our plate 175, fig. 14. The type locality, here designated, is 8-10 fms., entrance to Honolulu Harbor, Oahu Id., Hawaii. Synonymy — 1869 Pleurotoma lirata Pease, Amer. Jour. Concli., Philadel- phia, 5, p. 68 (non PI. lirata Reeve, 1845). 1952 Tunis cerithiformis (Dali, ms.) Tinker, Pacific Sea Shells, Honolulu, p. 46, pi. 47, fig. (upper row). Records — HAWAIIAN ISLANDS: entrance to Honolulu Harbour, 6-8 fms., Oahu (holotype, USNM) (D. Thaanum coll.) (ANSP); off Fort Armstrong, Honolulu, 15-20 ft. (ANSP); Ka Lae Kiki Koloa, Kauai (ANSP); Pearl and Hermes Reef, Hawaii (ANSP); off Kaanapali, West Maui, 25-75 ft.; Keaukaha Ponds, Hawaii; off Kewalo, Oahu, 20 fms. (D. Thaanum coll.). Xenuroturris castanella Powell, 1964 (Color pi. 175, figs. 21, 22; pi. 252) Range — Hawaiian Islands. Remarks — Again this species has long been known from distributed material bearing Dali’s manuscript name. ( See remarks concerning Tinker’s use of Dali’s manuscript names, above.) This is a member of the cingulifera series but an endemic Hawaiian one sufficiently distinct in its sculpture and coloration to warrant full specific status. Description — Adult shell, 35 to 45 mm. ( lfi to 1 % inches) in height. Whorls about 10, plus a multi- spiral, narrowly conic protoconch of 4 to 4M whorls, brown, densely sculptured with slightly curved, strong, rounded axials crossed at right angles by weak spiral lirae. Spire tall, 1M to 1/1 times the height of the aperture plus canal, 22° to 24°, out- lines lightly convex. Sculpture of spire whorls of closely spaced, rather evenly developed, moder- ately strong but narrow, crisp, spiral cords. Three cords on a slightly raised subsutural fold, 3 forming the sinus rib, situated above middle of whorl height and 2 primary cords with 3 threads in each inter- [22 - 964a] May 15, 1967 INDO-PACIFIC MOLLUSCA, vol. 1, no. 7 Xenuroturris 443 Xenuroturris gemmuloides Powell, new species (PI. 317) Range — Known only from the Hawaiian Islands from 16 to 267 fathoms. Remarks — Although this species has the general characters of the genus Xenuroturris, it has the gemmate or beaded peripheral keel of a Gemmula. It has the high spire and sinus of a Xenuroturris. This species differs from X. cerithiformis Powell, 1964 [p. 22 -964a] in having the peripheral keel minutely gemmate. Description — Shell about 19-25 mm. ( about one inch) in length, with a very tall spire and a trun- cated, decidedly flexed anterior canal, but with a gemmulate carina. Spire height almost twice that of the aperture plus the canal. Protoconch narrowly conical of about 3/2 whorls, the first two smooth, the remainder strongly and closely axially costate. Post- nuclear whorls 10. Spire-whorls sculptured, firstly with a broad but low subsutural fold, bearing three closely-spaced crisp cords, then a narrow shallow shoulder sulcus bearing two more cords, followed by a broad but rather weakly projecting sinus-keel, which is composed of two rather weak gemmate cords and an intermediate thread, the gemmules vertically fused, and closely spaced in cog-like fashion. From the peripheral carina to the lower suture there are two smooth spiral cords and sev- eral intermediate threads. From the peripheral keel to the neck there is a regular alternation of rela- tively strong smooth cords and weak threads; 10-12 closely-spaced smooth threads are on the anterior end. The anterior canal is strongly twisted and has an oblique shallowly-notched termination. The sinus is very deep, narrow and parallel-sided, its squarish apex occupying the full width of the peripheral keel. Colour of protoconch creamy white; remain- der of shell dull-white, irregularly maculated and streaked with reddish brown. Odd squarish spots are disposed upon the subsutural fold and upon the peripheral carina, and there is a broad ill-defined zone on the base. Measurements (mm.) — height width 24.7 7.7 holotype 19.2 6.5 paratype Types — The holotype is in the B. P. Bishop Mu- seum, Honolulu. The type locality is 16.5 fathoms, off Waianae, Oahu Island, Hawaii. Clifton Weaver, collector. Records — HAWAII: off Waianae, 16M fms., Oahu (C. Weaver) (type); off Honolulu, 60 fms. (“Pele II”; Mrs. M. King); off Waikiki, 100-120 fms., mud and sand (C. M. Burgess); off N.E. coast of Maui Island, 253-267 fms., bottom temperature 46° F. (Albatross Sta. 4084; USNM 338284). w Plate 317. Xenuroturris gemmuloides Powell, new species. Holotype, B. P. Bishop Museum. 16.5 fms., off Oahu Island, Hawaii; 24.8 mm. 444 A. W. B. Powell Turridae Xenuroturris ( Published by The Department of Mollusks Academy of Natural Sciences of Philadelphia 19th and the Parkway Philadelphia, Pennsylvania 19103 [22 - 968] May 15, 1967 INDO-PACIFIC MOLLUSCA, vol. 1, no. 7 T erebellum 445 THE GENUS TEREBELLUM (GASTROPODA: STROMBIDAE) by Peter Jung Naturliistorisches Museum, Basel, Switzerland and R. Tucker Abbott Pilsbry Chair of Malacology Academy of Natural Sciences of Philadelphia The strombid genus T erebellum is represented in the Recent fauna by a single species which is confined to the shallow waters of the Indo-Pacific region. Both the shell and the soft parts are highly modified. The animal is very active and is capable of darting rapidly through the water or rolling over sideways with great alacrity. It normally lives buried in coral sand in waters from low tide to a depth of 50 meters. Its egg laying and feeding habits are unknown, although from the nature of the stomach contents and the delicate radula it may be assumed that it is an herbivore like other strombids. Its manner of travelling under the surface of the sand while still maintaining contact with the watery world above has been described by D. P. Abbott (1962): “As each animal ploughed down into the sand, one of its eyestalks was extended upward and back over the shell. As this raised eyestalk was contacted by the sand which came to cover the anterior dor- sal region of the shell, the organ was moved in such a way that the terminal blue eyeball was placed just above the sand surface. The animals continued to move forward and down, burying themselves, but each one ‘left behind itself’ one eye protruding above the surface. Since the exposed eye remained stationary relative to the sand around it, the eye- stalk hidden below the sand was clearly elongating at a rate which matched the forward movement of the animal. When the shell was largely buried and Plate 318. Terebellum digs into the sand and travels for- ward, keeping one or the other of its eyeballs at the surface its anterior end was judged to be approximately one inch ahead of the exposed eye, the siphon was extended upward through the sand at this point, the siphonal folds closely appressed to form a closed cone. Once at the surface, the siphonal folds flared open terminally, and a swift current of water was drawn down into the mantle cavity. Following this inhalation, the second eye, thus far concealed below the surface of the sand with the rest of the animal, passed upward through the lumen of the siphon. With the second eye now exposed, the siphon folds unrolled and the siphon was pulled down out of sight, leaving the blue eyeball just at the surface of the sand. Simultaneously, the first eye, an inch to the rear, was withdrawn below the surface and disappeared. These actions were ob- served to be repeated with only minor variations as the animals burrowed along, their shells covered by a layer of sand perhaps a centimeter in depth. For- ward progression below the sand was nearly con- tinuous, but the eyes were ‘walked’ forward, with one of them always stationary and exposed at the surface like a periscope during the burrowing.” The gross anatomy of Terebellum is very un- usual in several respects. The foot is extremely compressed laterally and only the small anterior portion has a flat sole. The four-toothed operculum is almost entirely encased in a cape-like opercu- ligerous lobe. The ommatophores are very long, and the tentacles (not previously noticed by other workers) are extremely small and located just be- hind the distally-located eyeball. In S trombus, which rarely burrows, the tentacles are much larger and face downwards to prevent the ommatophore from bumping against the bottom. This function would not be required in the sand-burrowing Tere- bellum. at all times. ( Hypothetical drawing based upon an account by D. P. Abbott, 1962, and upon our anatomical studies. ) [09 - 801] 446 Terebellum P. Jung and R. T. Abbott Strombidac Plate 319. Gross anatomy of Terebellum terebellum Linne. Fig. 1, front view showing upheld siphonal appendage (s), eye (e) and foot (f). Fig. 2, right side of animal with shell removed, showing anus (a), eye (e), ommatophores (om), operculum (op), penis (pe), proboscis (pr), siphonal ap- The posterior end of the mantle edge is modified into three appendages, two of them being short filaments. The third is extremely long and forms the deep sutural gutter almost all the way to the apex of the shell. We presume that this exposed filament is tactile in function and serves to inform the animal whether the shell is fully buried or not. A similar gutter is present in the suture of the shell of the sand-burrowing Olividae. The sexes are separate. The female has a thin- walled flap bordering the egg channel which runs anteriorly from the oviduct to the right anterior side of the foot. It is similar to that in female Strombus, and it is likely that the egg strands of Terebellum are long, thin, coiling and sand-encrusted like those of Strombus. The penis is half the length of the entire animal. It has a narrow, open gutter for the passage of sperm running up to the two-lobed dis- tal end. The sides of the distal end of the penis are studded with about 20 to 30 proportionately large, chitinous hooks which are turned backwards. Pre- sumably, these barbs prevent the active female from escaping during copulation. A somewhat com- Plate 320. Operculum of T. terebellum from New Cale- donia. Dautzenberg collection, Institut Royal des Sciences Naturelles de Belgique, Brussels. 4 x. pendage ( s ) , sutural filament ( sf ) , and its cross-section ( x ) , and the minute tentacle (t). Fig. 3, distal end of penis showing chitinous hooks (h). Fig, 4, radula teeth (greatly enlarged ) . parable penial armament is found in the pyramidel- lid snail, Eulimella laevis ( Brown ) of northwestern Europe (D. Maas, 1964, Zool. Anzeiger, 173, pt. 2, p. 143 ) . The only other prosobranch genus bearing a chitinous, hooked armament is Stenothyra, a fresh-water hydrobiid (R. T. Abbott, 1951, Jour. Wash. Acad. Sci., 41, p. 14). The radula is small, delicate, and located at the anterior end of the pharynx. It is very similar to that of some Strombus and Lambis, and has 35 rows of teeth. Both the inner and the outer marginal teeth are long and narrow. The stomach and intes- tine contain detritus, coral sand, foraminifera and microscopic bits of shell. The shell of Terebellum is elongate and smooth, features which would permit rapid darting through the water. There is no stromboid notch in the edge of the shell for the protuberance of the right eye- stalk. Evidently, the eyestalks are protruded up through the lumen of a very specialized siphonal extension of the anterior portion of the mantle edge. This “eyestalk sheath” is a separate organ, much like the siphonal flaps developed in the man- tle of the Volutidae. Plate 321 (color plate on opposite page). Terebellum tere- bellum ( Linne ) showing the variability in color patterns in one species. Figs. 1-6, 13 and 15 from near Noumea, New Caledonia. Figs. 1, 2 and 14 are forma lineatum Roding. Figs. 3, 5 and 6 are forma punctulorum Roding. Figs. 9, 10 are forma delicatum Kuroda and Kawamoto from Honshu Id., Japan. Fig. 11 is forma nebulosum Roding from Oman, S. E. Arabia. Fig. 14, Upolu Id., Samoa. All natural size. Lower figure is a living Terebellum terebellum (Linne) from the Palau Islands attempting to dig into the sand. ( x 3. ) [09 - 802] May 15, 1967 INDO-PACIFIC MOLLUSCA, vol. 1, no. 7 Terebellum 447 Plate 321. Terebellum (see opposite page for explanation). [09 - 803] 448 Terebellum P. Jung and R. T. Abbott S trombidae Geological History The genus Terebellum came into existence dur- ing the early Tertiary. It represents a predominantly Tethyan group with the center of its evolution lying in the Indo-European region. The earliest records date back to the Paleocene. The interval from the Paleocene to the middle Eocene was characterized by a rapid morphological development and a geo- graphic expansion which was mainly directed to- ward the west. During that period the genus split into what we recognize today as three subgenera: Terebellum s.str.. Seraphs Montfort (type species: Bulla sopita Solander), and Mauryna de Gregorio (type spe- cies: Terebellum (Mauryna) plicatum d’Archiac and Haime). Seraphs, which is mainly characterized by its involute shell, is known from the Paleocene, but Mauryna, which carries more or less accentuated axial plicae appeared in the Eocene only. Seraphs was apparently more successful than Mauryna, as it is known from many species distributed almost all over the tropical zones of the world, whereas Mauryna, which did not reach the New World, is represented by a few species restricted to Europe, India and Java. In the middle Eocene the genus reached the Caribbean (Jamaica) and Central America (Pan- ama), and during the late Eocene times it spread over the entire Caribbean, Florida, and as far as California. Towards the east the genus did not get farther than Java during the late Eocene. Mauryna became extinct before the Oligocene, whereas Seraphs is known from a few early Oligo- cene localities, but did not survive the Oligocene. The transition from the Paleogene to the Neo- gene was not only marked by a rapid decline in morphological variety and number of species, but also by a considerable restriction of the geographi- cal range of Terebellum s.str., the only surviving subgenus. In the Western Hemisphere, Terebellum is absent in post-oligocene deposits with the excep- tion of an unconfirmed record from the early Mio- cene of Florida. In Europe there are no Miocene records at all. In late Miocene to Pliocene times the Recent species came into existence. Although fossil records of T. terebellum are rare, it seems that its distribu- tion was about the same as that of today. From these facts it appears that the Recent Tere- bellum represents a highly specialized, dwindling group compared with the morphologically rich and widely distributed fossil group of over 50 described taxa. Acknowledgments We are indebted to the following persons for the loan of or information on material: W. Adam, Institut Royal des Sciences Naturelles, Brussels, Belgium; C. O. van Regteren Altena, Leiden, Neth- erlands; W. J. Clench, Museum of Comparative Zoology, Cambridge, Mass.; B. R. Wilson, Western Australian Museum, Perth. The drawings of the opercula and shells were made by O. Garraux, Basel, Switzerland. Selected Bibliography Abbott, D. P. 1962. Observations on the Gastropod Tere- bellum terebellum (Linnaeus), with particular reference to the behavior of the eyes during burrowing. Veliger, vol. 5, no. 1, pp. 1-3. Beets, C. 1950. On Quaternary Mollusca from the Islands of Boenjoe and Tarakan, E. Borneo. Leidsche Geol. Meded., deel 15, pp. 241-264. Bergh, R. 1895. Beitrage zur Kenntniss der Strombiden, be- sonders der Gattung Terebellum Klein. Zool. Jahrbiicher, Abt. Anatomie und Ontogenie, vol. 8, pp. 342-378, pis. 22-23. Boettger, O. 1908. Liste der tertiaren und jiingeren Ver- steinerungen. In: Verbeek: Molukken-verslag. Jaarb. Mijnwezen Nederl. O. -Indie, Jg. 37, pp. 668-675. Cox, L. R. 1931. The Geology of the Farsan Islands, Gizan and Kamaran Island, Red Sea. Part 2. Molluscan Palae- ontology. Geol. Mag., vol. 68, pp. 1-13. Cox, L. R. 1948. Neogene Mollusca from tire Dent Penin- sula, British North Borneo. Schweiz. Pal. Abh., vol. 66, pp. 1-70, pis. 1-6, 3 text figs. Dey, A. K. 1962. The Miocene Mollusca from Quilon, Kerala (India). Palaeont. Indica, N.S., vol. 36, pp. 1-129, pis. 1-10, text figs. 1-2. Dodge, Henry. 1955. A historical review of the mollusks of Linnaeus, pt. 3. Bull. Amer. Mus. Nat. Hist., vol. 107, pp. 1-157. Koert, W. and F. Tornau. 1910. Zur Geologie und Hydrol- ogie von Daressalam und Tanga ( Deutsch-Ostaf rika ) . Abh. K. Preuss. Geol. Landesanstalt, N.F., Heft 63, 77 pp., pis. 1-10. Martin, K. 1880. On a posttertiary fauna from tire stream- tin-deposits of Blitong (Billiton). Notes from the Leyden Museum, vol. 3, Note 7, pp. 17-22. Martin, K. 1928. Mollusken aus dem Neogen von Atjeh in Sumatra. Mijnbouw in Nederlandsch-Indie, Wetensch. Meded., no. 10, pp. 1-36, 1 pi. Nuttall, C. P. 1965. Report on the Haile collection of fossil Mollusca from the Plio-Pleistocene Togopi formation, Dent Peninsula, Sabah, Malaysia. Geol. Survey, Borneo Region, Malaysia, Mem. 16, pp. 155-192. Oostingh, C. H. 1935. Die Mollusken des Pliozans von Boemiajoe (Java). Dienst van Mijnbouw in Nederlandsch. Indie, Wetensch. Meded., no. 26, pp. 1-247, pis. 1-17, 31 text figs., 1 map. Wissema, G. G. 1947. Young Tertiary and Quaternary Gas- tropoda from the Island of Nias (Malay Archipelago). Thesis. Leiden, 212 pp., 7 tables, 6 pis., 1 map. [09 - 804] May 15, 1967 INDO-PACIFIC MOLLUSCA, vol. 1, no. T erebellum 449 Genus Terebellum Roding, 1798 Type: Terebellum terebellum Linne, 1758 Subgenus Terebellum Roding, 1798 Although several Recent species have been de- scribed, it is our opinion that there is only one living species, namely terebellum Linne, 1758. The subgenus Terebellum sensu stricto is characterized by its elongate, narrow, strong shell, lacking a stromboid notch, and having no axial sculpture. The type by indirect tautonomy is based on both Terebellum nebulosum Roding, 1798, and lineatum Roding, 1798, both of which are color forms of Bulla terebellum Linne, 1758. If legalistic doubt exists, we hereby designate T. nebulosum Roding, 1798, as the type of the genus. Synonymy — 1798 Terebellum Roding, Museum Boltenianum, Hamburg, pt. 2, p. 135. 1799 Terebellum Lamarck, Memoires Soc. d’Hist. Naturelle de Paris, 1, p. 69, no. 4 (type by tautonomy: Bulla terebellum Linne, 1758). 1810 Terebellum Montfort, Conchyliologie Systematique, 2, p. 379 (type: subulatum Lamarck). 1815 Terebrina Rafinesque, Analyse de la Nature, Palermo, p. 145 (emendation for Terebellum Lam.). 1848 Lucis Gistel, Naturgeschichte des Thierreichs fur hohere Schulen bearbeitet, Stuttgart, p. 170 ( type by monotypy: L. subulatus ). 1848 Artopoia Gistel, ibidem, explanation to pi. 7, fig. 8 (type by monotypy: A. subulata). 1887 Artopoia Gistel, Paetel, Catalog der Conchyl.-Samml., Berlin, 1, p. 315. Plate 322. The original specimen of “Terebellum punc- tatum Chemnitz” illustrated by Chemnitz in Neues Sys- tematisches Conchylien-Cabinet, vol. 10, pi. 146, figs. 1362, 1363. 43.1 mm. in length. Zool. Mus. Copenhagen. Terebellum terebellum (Linne, 1758) (Pis. 318-327) Range — East Africa to Samoa; Japan to Aus- tralia. Remarks — This is a colorful, moderately common species found in shallow water throughout most of the tropical Indo-Pacific. The color patterns are very variable, even on the same specimen. This has led to a large number of synonyms, some of which we list below as mere color forms: typical form ( Linnaeus’ type ) : ( pi. 323 ) . Un- fortunately the type specimen is an old beach- worn and almost colorless specimen. Linnaeus described it as “white.” The name album Link, 1807, is based on a similar form. Lamarck’s subulatum is merely a substitute name for this form. forma nebulosum Roding, 1798: (pi. 321, fig. 11). Refers to Conchyl. Cab., vol. 2, figs. 568, 569, the “banded borer.” Characterized by three spiral, brown bands of color. forma lineatum Roding, 1798: (pi. 321, figs. 1, 14). Refers to Lister, pi. 736, fig. 31, which is char- acterized by strong, numerous, narrow, zigzag brownish lines. forma punctulorum Roding, 1798: (pi. 321, fig. 6). Refers to Lister, pi. 737, fig. 32. Charac- terized by numerous, small, red-brown dots over the entire surface. This is the same as the shell illustrated in Martini and Chemnitz, vol. 10, figs. 1362, 1363, later referred to as “punc- tatum ” Chemnitz by Reeve, K. Martin, La- marck, and others. forma delicatum Kuroda and Kawamoto, 1961, in Habe: (pi. 321, figs. 9, 10). Characterized by its soft-brown background which is covered with brown dots which have a white half-moon surrounding the anterior side of each dot. Most common in Japan and Madagascar. Other forms, including the large. New Cale- donian and Arabian shells, with a brown line on the suture and on the anterior edge of the body whorl, have not had names applied to them. In our opinion, they are not valid subspecies. Dodge ( 1955, pp. 41-45) has given a very lengthy account of the history of the nomenclature of Tere- bellum terebellum Linne, with which we agree in the details he sets forth. There is little worth re- peating, except perhaps to say that Linnaeus’ 1764 re-description is an accidental mixture of terebel- lum and a species of Conus. [09 - 809] 450 Terebellm P. Jung and R. T. Abbott Strombidae Plate 323. Bulla terebellum (Linne). Specimen from Lin- naean collection. Lectotype. Length 37.5 mm., maximum diameter 10.6 mm. Courtesy of the British Museum (Natu- ral History ) . The general shape of the shell is also variable to some degree. Usually the spire is flat-sided and the body whorl but little convex. Somewhat stouter shells with slightly convex spire whorls occur mainly in the areas of New Guinea and the Samoan Islands. The height of the spire is about one third, and the maximum width about one fourth of the total height. These proportions are disturbed in stouter forms. Immature specimens have different proportions. The height of the spire is much smaller compared to the total height. In addition, they have a num- ber of incised, spiral lines near the base which are lacking in full-grown shells (see under ontogeny). It must be remembered that adult T. terebellum is much more resistant mechanically which means that immature shells are not likely to be preserved on the shore, i.e., in the zone of heavy wave action or other mechanical influences. Thus it is not aston- ishing to find lots consisting only of adult shells in the collections. A glimpse at the depth data shows that the majority of immature specimens has been dredged at depths greater than 15 meters. Habitat — T. terebellum seems to prefer a bottom of silt, sand or sandy mud between some coral, rock, and patches of algae. It has been collected dead from a few feet down to 50 fathoms, and live specimens have been recovered from a water depth of a few inches to 18 fathoms. Ontogeny — The ontogenetic development of T. terebellum can be subdivided into 4 stages, namely a protoconch stage, a first terebelloid stage, seraph- oid stage, and a second terebelloid stage. The limits betwen these stages are not sharp, of course, but transitional. The protoconch (see pi. 326, figs. 1, 2; pi. 325, fig. 2) consists of about one volution of glassy shell material. The two characteristic features are its almost planispiral coiling, and its subcircular cross section. The inner walls of the protoconch touch to form a “protoconchal columella.” The beginning of the first terebelloid stage (see pi. 326, figs. 1, 2; pi. 325, fig. 2) is marked by an accelerated growth in height, i.e., the tube gradu- ally becomes more than twice as high as wide. This stage is evolute with the suture descending at a low angle. The shell material is still entirely glassy and translucent, but two structural elements appear: faint growth lines, and a thin callus on the inner lip. This stage comprises almost two volutions. The seraphoid stage (see pi. 326, figs. 3, 4; pi. 325, fig. 2) is produced by the appearance of a spiral band of callus just above the suture which soon ascends spirally toward the apex. The suture of the first terebelloid stage then is covered, and the shell becomes entirely involute. The seraphoid stage con- sists of 1 to lM volutions, and is further character- ized by the appearance of the first color patterns which are more or less pronounced, however. In Plate 324. Variability of general outline. Fig. 1. Immature. Bawi Id., W. Zanzibar. ANSP 213721. Fig. 2. Immature. Honshu Id., Tapan. ANSP 234910. Fig. 3. Almost adult. Tutuila Id., Samoa Ids. ANSP 209255. Fig. 4. Adult. Sem- porna, North Borneo. ANSP 295409. Fig. 5. Adult. Noumea, New Caledonia. ANSP 237367. Slightly enlarged. [09 - 810] May 15, 1967 INDO-PACIFIC MOLLUSCA, vol. 1, no. 7 Terebellum 451 Plate 325. Fig. 1. Cross section through shell at suture showing prismatic and nacreous layer, sutural callus and organic matter (black). 50 x. Fig. 2. Cross section through early ontogenetic stages showing protoconch, the first tere- belloid stage, and the involute coiling of the seraphoid stage. The cross section of the tube changes with growth. 50 x. addition, two sculptural elements are introduced: faint, somewhat irregular, axial lines on the spire, and fine incised, spiral lines near the base. The interspaces between these lines may be regular or irregular in width. Some of the upper lines may be crossed by growth lines to form small rhombs. Specimens from the seraphoid stage look like mi- nute members of the Paleocene to Oligocene sub- genus Seraphs. The second terebelloid stage (see pi. 326, figs. 5, 6; pi. 325, fig. 2) starts when the shell becomes evolute again. The angle between the axis and the suture is smaller than in the first terebelloid stage, i.e., the suture now descends more rapidly. The spiral band of callus above the suture is conspicu- ous. The incised lines near the base, which appear in the seraphoid stage, continue into this stage, but they gradually disappear as the shell grows larger. In full-grown specimens this stage consists of about 2M volutions. The above remarks on the ontogeny are based on a rich sample from near Nossi Be, N.W. Mada- gascar (ANSP 261971), containing about 70 speci- mens representing all growth stages. Description — Shell usually between 40 and 50 mm. in length (about 1.5 to 2 inches), slender, cylindrical, moderately heavy, and without a strom- boid notch. Immature shells delicate. Aperture nar- row, pointed above, regularly widening toward the base. Spire usually almost flat-sided, pointed, about half as high as the rest of the shell. Base of body whorl higher than base of columella. Outer lip slightly thickened. Inner lip with a callus which continues as a spiral band above the suture. Suture with a very deep, narrow channel running from the apex to the outer lip. Surface of adult shell smooth with the exception of faint growth lines. Adults with about 7 whorls (see under ontogeny). The predominant color is brown to yellowish brown. The variation of the intensity of the color and the color patterns is considerable (see under remarks). [09-811] 452 Terebellum P. Jung and R. T. Abbott Strombidae Measurements ( mm.) — length 57.6 width 14.2 spire height 19.0 Oman, S.E. Arabia 49.1 12.5 16.6 Mindanao Id., Philippines 44.0 11.0 14.3 Sanga Sanga Id., Sulu 41.7 12.7 11.6 Archipelago Schouten Ids., New Guine; The largest specimen seen has a length of 77.1 mm. (New Caledonia: ANSP 30215). Synonymy — 1758 Conus terebellum Linne, Systema Naturae, ed. 10, p. 718, no. 284 (in Asia); 1764, Museum Ludovicae Ulricae, p. 564, no. 178. 1767 Bulla terebellum Linne, Systema Naturae, ed. 12, p. 1185. [1788 Terebellum punctatum Chemnitz, Neues systema- tisches Conchylien-Cabinet, 10, p. 124, pi. 146, figs. 1362, 1363. [Non-binomial.] 1791 Conus terebellum Gmelin, Systema Naturae, ed. 13, 1, pt. 6, p. 3390. 1798 Terebellum nebulosum Roding, Museum Boltenia- num, pt. 2, p. 135 (refers to Bulla terebellum and to Lister, pi. 736, fig. 30). 1798 Terebellum lineatum Roding, Museum Boltenianum, pt. 2, p. 135 ( refers to Bulla terebellum and to Lister, pi. 736, fig. 31). 1798 Terebellum punctulorum Roding, Museum Boltenia- num, pt. 2, p. 135 (refers to Lister, pi. 737, fig. 32). 1799 Terebellum terebellum Linne, Lamarck, Mem. Soc. Hist. Nat. Paris, 1, p. 69. 1801 Terebellum terebra Bose. Histoire naturelle des coquilles, tome 5, p. 72, pi. 38, fig. 7 ( refers to Bulla terebellum Linne and to Lister, pi. 736, figs. 30, 31). 1807 Terebellum variegatum Link, Beschreibung der Na- turalien-Sammlung der Universitat zu Rostock, p. 99 (refers to Conus terebellum Linne). 1807 Terebellum album Link, ibidem, p. 99. 1810 Terebellum sabulatum (sic) Lamarck, Montfort, Conchy 1. Syst., 2, p. 379, in text. 1811 Terebellum subulatum Lamarck, Ann. Mus. Hist. Nat. Paris, 16, p. 301 (refers to Lister, pi. 736, figs. 30, 31; pi. 737, fig. 32, and to others); 1822, Lamarck, Histoire naturelle des animaux sans vertebres, 7, p. 410. 1811 Terebellum lineatum Perry, Conchology, London, pi. 37, fig. 1 (no locality). 1811 Terebellum spirale Perry, ibidem, pi. 37, fig. 2 (Eastern Ocean). 1848 Terebellum subulatum Lamarck, Adams and Reeve, Mollusca. In: Adams, A.; The zoology of the voy- age of H.M.S. Samarang, p. 36, pi. 9, fig. 6; 1859, Sowerby, Thesaurus Conchyliorum. Terebellum, pi. 218, figs. 1-5; 1874, Lischke, Japanische Meeres- Conchylien, 3. Theil, p. 21; 1880, K. Martin, Notes from the Leyden Museum, 3, Note 7, p. 19. 1863 Terebellum punctatum Reeve, Conch. Icon., 14, Terebellum, pi. 1, figs. la-g. 1886 Terebellum punctatum Chemnitz, Watson, Report . . . Challenger, 15, p. 421; 1899, K. Martin, Samml. geol. Reichsmus. Leiden, N.F., Bd. 1, p. 195, pi. 31, figs. 452, 452a; 1908, Boettger, in: Plate 326. Ontogenetic development of shell. Two left figs. right figs. Beginning of second terebelloid stage and final Protoconch and first terebelloid stage. Middle two figs. Be- terebelloid stage. All specimens from near Nossi Be, N.W. ginning of seraphoid stage and full seraphoid stage. Two Madagascar (ANSP 261971). 4 x. [09 - 812] May 15, 1967 INDO-PACIFIC MOLLUSCA, vol. 1, no. 7 Terebellum 453 Plate 327. Geographical distribution of T. terebellum. Solid circles: Recent occurrences. Crosses: fossil occurrences (partly according to literature). Verbeek: Molukkenverslag. Jaarb. Mijnwezen Nederl. O.-Indie, Jg. 37, p. 674. 1910 Terebellum subulatum Lamarck, Koert and Tornau, Abh. K. Preuss. Geol. Landes-anstalt, N.F., Heft 63, p. 10. 1911 Terebellum punctatum Chemnitz, Martin-Icke, in: Salenka and Blanckenhorn : Die Pithecanthropus- Schichten auf Java, pp. 47, 49; 1919, K. Martin, Unsere palaezoologische Kenntnis von Java, pp. 92, 133, 141; 1928, K. Martin, Wet.' Meded. Dienst Mijnbouw, No. 10, pp. 8, 25. 1931 Terebellum terebellum (Linne) ( subulatum La- marck), Cox, Geol. Mag., 68, p. 7. 1931 Terebellum punctatum Chemnitz, Van der Vlerk, Leidsche Geol. Meded., deel 5, p. 248. 1931 Terebellum subulatum Lamarck, Van der Vlerk, ibidem, p. 248. 1931 Terebellum punctatum Chemnitz, van Es, The age of Pithecanthropus. Thesis, Delft, p. 51. 1935 Terebellum terebellum (Linne), Oostingh, Wet. Me- ded. Dienst Mijnbouw Nederl. Indie, No. 26, p. 58; 1947, Wissema, Young Tertiary and Quater- nary Gastropoda from the Island of Nias ( Malay Archipelago). Thesis, Leiden, p. 92; 1948, Cox, Schweiz. Pal. Abh. 66, p. 31, pi. 1, figs. 6a, b; 1950, Beets, Leidse, Geol. Meded., deel 15, p. 244; 1959, Kira, Coloured illustrations of the shells of Japan, p. 34, pi. 15, fig. 1; 1961, Nuttall, Brit. Bor- neo Geol. Survey, Ann. Rep. 1960, p. 92; 1962, D. P. Abbott, Veliger, 5, No. 1, pp. 1-3; 1962, Dey, Palaeont. Indica, N.S., 36, p. 67; 1962, Kira, Shells of the Western Pacific in color, p. 34, pi. 16, fig. 1; 1965, Nuttall, Geol. Survey, Borneo Region, Malaysia, Memoir 16, p. 170. 1961 Terebellum terebellum delicatum Kuroda and Kawa- moto, Habe, Coloured illustrations of the shells of Japan, 2, p. 38, pi. 17, fig. 1 (manuscript name?); 1964, Habe, Shells of the Western Pacific in color, 2, p. 58, pi. 17, fig. 1. Lectotype selection — The specimen from the Lin- naean collection at present at the British Museum (Natural History), marked with the remnants of the number ‘388,’ is here selected as lectotype. The number ‘388’ corresponds to that with the species name in the 12th edition of the Systema Naturae according to written communication from S. P. Dance. The lectotype measures 37.5 mm. in length, and has a maximum diameter of 10.6 mm. There is no locality information. Selected records (see map) — RED SEA: Eilet, Gulf of Aqaba, Israel (USNM). MOZAMBIQUE: off dock Nacala (Franz Steiner, ANSP). ZANZIBAR: 1 mi. N. of Bawi Id.: 1/2 mi. W.S.W. Ras Mungwe; between Mwamba Ukombi and Chumbe Ids. (all ANSP). COMORE IDS.: Lagon de Mayotte (ANSP). MADAGASCAR: 2-3 mi. S. of Nosy Iranja, 32 mi. S.W. of Nossi Be (ANSP). MAURITIUS: ( MCZ, USNM). SEYCHELLES: 15-40 ft., sand and algae, Baie Temay, West Mahe Id., Feb. 9, 1966, R. E. Ostheimer and Minerva Buerk (ANSP). ARABIA: Oman (ANSP, USNM). MALDIVE IDS.: Fadiffolu Atoll; Addu Atoll (both ANSP). THAILAND: 25 mi. N.N.W. of Phuket Id., Andaman Sea ( Anton Bruun Cruise 1, ANSP). BORNEO: Kudat Bay, North Borneo; Sempoma, North Borneo (both Mary Saul, ANSP, USNM). PHILIPPINES: frequent throughout the archipelago on Luzon, Cebu, Leyte, Minda- nao, etc. (ANSP, USNM). JAPAN: Baso Peninsula, Chiba Pref., Honshu (ANSP); Tatsugahama, Wakayama Pref., Honshu (ANSP); Southern Kyushu (ANSP). RYUKYU IDS.: Okinawa Id. (ANSP, MCZ, USNM). MARIANAS: N. side of Apra Harbor, Guam Id. (ANSP, USNM). ENI- WETOK ATOLL: (USNM). CAROLINE IDS.: frequent throughout the archipelago. MARSHALL IDS.: Bikini Atoll (USNM); Kwajalein Atoll (USNM). INDONESIA: Ambon (Amboina) Id. (MCZ); 35 meters, Sunda Straits (Copen- hagen Mus.). NEW GUINEA: Aoeri Ids., Geelvink Bay (ANSP, MCZ); Japen Id. (ANSP); Schouten Id. (ANSP, MCZ); Finschhafen Bay (MCZ). NEW BRITAIN: Matapi Id., 5 mi. from Rabaul (ANSP); Rabaul Harbor (ANSP); Kumbun, near Kandrian (ANSP). SOLOMON IDS.: Atao District, Malaita Id. (Rev. J. van der Riet, ANSP); Roviana Lagoon, New Georgia (MCZ). AUSTRALIA: Broome, Western Australia (MCZ); Cape Leveque, Western Aus- tralia (USNM); Lizard Id., 80 mi. N.E. of Cooktown, Queensland (ANSP). NEW CALEDONIA: Noumea (ANSP, MCZ, USNM); lie des Pins (MCZ). LOYALTY IDS.: Ouvea Id., D. Getz, 1949 ( BPBM ) . NEW HEBRIDES: Aore Id. (MCZ). FITI IDS.: (ANSP, MCZ). SAMOA IDS.: Tutuila Id. (ANSP, MCZ, USNM). TONGA IDS.: Tonga- tapu Id. (USNM). [A record of “Marquesas Islands” in the Dautzenberg collection in Brussels is open to question.] [09 - 813] 454 Terebellum P. Jung and R. T. Abbott Strombidae Fossil records- MIOCENE: Java (Martin, 1928, p. 25); Quilon, Kerala, India (Dey, 1962, p. 67). NEOGENE: Java (Oostingh, 1935, p. 59); Saonek besar near New Guinea (Boettger, 1908, p. 674; Oostingh, 1935, p. 59). PLIO- CENE: Java (Oostingh, 1935, p. 59, and others); Sumatra (Martin, 1928, p. 8); North Borneo, Dent Peninsula (Cox, 1948, p. 31; Nuttall, 1965, p. 170); Borneo, North Kutei (collections of the Natural History Museum Basel); Timor (Cox, 1948, p. 31); East Ceram (collections of the Natural History Museum Basel). PLIO-PLEISTOCENE: Nias (Wis- sema, 1947, p. 93). QUATERNARY: Farsan Kebir, Red Sea (Cox, 1931, p. 7); Dar es Salaam, Tanganyika (Koert and Tornau, 1910, p. 10); Belitung Id. (Billiton Id.) (Mar- tin, 1880, p. 19); Bunju Id., East Borneo (Beets, 1950 p. 244). Published by The Department of Mollusks Academy of Natural Sciences of Philadelphia 19th and the Parkway Philadelphia, Pennsylvania 19103 [09 - 814] May 15, 1967 INDO-PACIFIC MOLLUSCA, vol. 1, no. 7 S trombus 455 STROMBUS (CANARIUM) WILSONI NEW SPECIES FROM THE INDO-PACIFIC by R. Tucker Abbott Pilsbry Chair of Malacology Academy of Natural Sciences of Philadelphia Strombus wilsoni Abbott, new species (PI. 328, figs. 1-3) Range — East Africa and Australia to Fiji. Remarks — A small, inch-long Strombus has been recently collected in several widely scattered parts of the Indo-Pacific, which appears to be a new sympatric relative of the well-known S. mutabilis Swainson [p. 09-900] and S. microurceus (Kira, 1959) [p. 09-899]. This species is rare in Zanzibar and Fiji, but is moderately common off the coast of Western Australia in water from 1 to 23 fathoms in depth. Strombus wilsoni is characterized by 5 to 7 equal- sized, elongate, short, rounded nodules on the shoulder of the last half of the body-whorl, by the squarish penultimate whorl which bears about 20 neat, elongate, smooth riblets, and by a columella which is smooth and whitish at the center. The base of the columella bears 7 to 10 minute brown- ish to purple-brown teeth. S. microurceus differs in having the interior half of the columella colored purple-brown, and in having a blue-green splotch on the shoulder of the last whorl just below the suture and just behind the apertural varix. S. mu- tabilis differs in having no nodules immediately behind the upper part of the apertural varix, and in having a white-striped columella. This species is named after the Australian mala- cologist, Dr. Barry Wilson, and also after Mr. Wil- son Darwin of Philadelphia. Description — Adult shell 18 to 28 mm. ( about an inch) in length, solid, elongate-quadrate, nod- uled and mottled with light yellow-brown. Whorls 8 or 9. Nuclear whorls 3, glassy, smooth, translucent lavender or whitish and bulimoid in shape. Post- nuclear whorls glossy, minutely malleated and with a small but distinct subsutural cord. The spire has about 9 to 11 tan, former varices. Penultimate whorl square-shouldered, with about 15 to 22 mi- nute, elongate, axial riblets. Body- whorl rounded and smooth on the parietal side, but bears 5 to 7 Plate 328. Strombus wilsoni Abbott, new species , Figs. 1, 2, paratype, 23 fathoms, off Legendre Island, Dampier Archi- pelago, Western Australia, 1960. W.A. Museum no. 847-66, 22.1 mm. Fig. 3, apertural view of holotype from Zanzibar. ANSP no. 253088. Total length: 22.2 mm. Fig. 4, Strombus mutabilis Swainson, showing columellar sculpturing. Fig. 5, S trombus microurceus ( Kira ) , showing columellar sculp- turing. [09 - 919] 456 Canarium R. T. Abbott Strombidae equal-sized, short, rounded, elongate nodules at the shoulder on the last half of the whorl. Varix behind the outer lip is swollen and spirally threaded. Colu- mella whitish to tan, smooth at the center and bounded above and below by minute tan to purp- lish teeth. Interior of aperture with numerous, raised, wavy, brown to purple lirae or threads. The aperture is narrowly constricted at its posterior end. Stromboid notch rather shallow and near the base of the outer lip. Color of shell whitish to cream, with red-brown to tan maculations and spiral rows of small, sparse, brown or white dots. Periostracum very thin and translucent. Operculum stromboid, barely arching, light-brown, and with 10 to 12 small, sharp serrations. Measurements (mm.) — length width no. who; 22.3 10.1 8 28.4 13.5 9 23.7 11.5 8 22.2 10.5 8 19.5 9.0 7 18.4 9.5 7 holotype; Zanzibar paratype; Zanzibar paratype; Legendre Id. paratype; Legendre Id. paratype; Legendre Id. paratype; Nandi Bay, Fiji Types and Records — The type locality is Zanzibar. The holotype is in ANSP no. 253088. Paratypes come from: ZANZIBAR: 8 fms., fine grass and shell bottom, 1.5 miles W.S.W. of Ras Mungwe. Feb. 20, 1960; A. J. Ostheimer, 3rd V. Orr (Maes) and Richard Thorington, coll. (ANSP 212695). WESTERN AUSTRALIA: 23 fms., off Legendre Island, Dampier Archipelago (Western Australian Mus. 847-66); low tide, Eaglehawk Island (W.A. Mus. 840-66); 14 fms., 14 miles west of Eaglehawk Island (W.A. Mus. 838-66); 6-10 fms., west of Flat Island, near Onslow (W.A. Mus. 851-66; ANSP). Paratypes also in B. P. Bishop Mus. Strombus rugosus Sowerby, 1825 (PI. 20, figs. 11, 12; p. 09-840) Range — Fiji, the Ellice, Samoan and Tonga Islands. Remarks — I concur with Cernohorsky ( 1965, p. 5 ) that this taxon should be considered as a species, rather than as a subspecies of erythrinus Dillwyn, 1817 (see p. 09-909), since it has now been shown that the two forms co-exist without seeming hy- bridization. New Fijian records of rugosus are added below. I do not think it wise to consider the New Cale- donian populations of erythrinus as subspecifically different from those of Fiji or other neighboring islands, since many New Caledonian species possess darker colors, stronger sculptural features, and greater average sizes, probably due to environ- mental conditions. Since the appearance of my 1962 monograph on Strombus (Indo-Pacific Mollusca, vol. 1, no. 2), three other species have been recorded from Fiji by Cernohorsky (1965); S. labiosus Wood, 1828, S. ther sites Swainson, 1823, and S. haemastoma Sowerby, 1842. Synonymy — 1962 Strombus erythrinus subsp. rugosus Sowerby, Abbott, Indo-Pacific Mollusca, 1, no. 2, p. 81 [09-909], 1965 S trombus rugosus Sowerby, Cernohorsky, Records of the Fiji Museum, 1, no. 1, p. 5 (type locality desig- nated: Viti Levu, Fiji). Records — FIJI : 20-30 ft. Namotu Id., Vanua Levu Id. (A. Jennings, 1962), Wading Id., Viti Levu Id. (A. Jen- nings, 1962); beach, Tivoa Id., N.W. of Viti Levu Id. (A. Jennings, 1961). All ANSP. Common on sand and algae substrate, 0-100 ft.; also sand pockets on coral reef fiats (fide Cernohorsky, 1965, p. 5). Published by The Department of Mollusks Academy of Natural Sciences of Philadelphia 19th and the Parkway Philadelphia, Pennsylvania 19103 [09 - 920] December 8, 1967 MOLL INDO-PACIFIC MOLLUSCA, vol. 1, no. 8 Cypraea 457 THE GENUS CYPRAEA (SUBGENUS ZOILA JOUSSEAUME) by Barry R. Wilson Western Australian Museum, Perth and Jennifer A. McComb University of Western Australia, Perth The generic classification within the family Cy- praeidae is a controversial subject. Several sub- families and many genera and subgenera have been described, primarily on shell characters of living and fossil species, but the diagnostic features sup- posed to distinguish these taxa are difficult to apply with any satisfactory degree of consistency. The result has been a number of confusing and contra- dictory classifications. Most of the taxa have not been clearly diagnosed, and we have found that the only key to the genera so far published ( Steadman & Cotton, 1946) is unworkable. Kay ( 1960a ) has shown that available anatomical data do not confirm the conchological classifications. She proposed to revert to the generic name Cypraea Linnaeus, 1758, for all cowries as a temporary measure until such time as additional anatomical data may permit a new and more comprehensive re-assessment of the problem. This seems to be a reasonable expediency, if not a practical solution. It is true that, if this procedure is widely and per- manently adopted, many nomenclatural difficulties ( particularly homonymy ) will arise ( Schilder, 1963b). However, this is no justification for an un- satisfactory classification and only highlights one unfortunate consequence of premature splitting. In order to expedite solution of the problem of generic classification of the Cypraeidae, anatomical data are urgently needed for many more species. In this paper, observations on the anatomy, shell morphology, habitats and distributions of five cow- Plate 329 Living Cypraea (Zoila) venusta Sowerby from 3 fathoms, collector and photographer, 1962. Natural size, off Sorrento Beach, Perth, Western Australia. B. R. Wilson, 458 Zoila B. R. Wilson and J. A. McComb Cypraeidae ries endemic to Western Australia and South Aus- tralia are presented. Previously these five species have been grouped together to form the genus Zoila Jousseaume, 1884. Zoila has often been allied with Bernaya Jousseaume, 1884 and S iphocypraea Heilprin, 1887. Some authors (e.g. Schilder, 1965) have placed these three genera in the subfamily Cypraeorbinae, thus separating them from other cypraeids; other authors (e.g. Steadman & Cotton, 1946) have used the subfamily name Zoilinae for them. The status and relationships of Zoila are evalu- ated here. It is shown that four of the five species have several apparently distinctive characters in common and may be regarded as a phyletic cluster of species. The relationship of the fifth species re- mains somewhat doubtful. This evidence seems to justify retention of Zoila as a taxon. However, be- cause of the weakness of the diagnostic shell char- acters and the lack of comparative anatomical data for other cowries, we support Kay (1960a) in recognising a single genus, Cypraea, but rank Zoila as a subgenus. This procedure we regard as a tem- porary expediency and hope that our data and dis- cussion may contribute toward a better under- standing of the relationships of species within the Cypraeidae. Anatomy Kay (1960a) observed that the 52 species of cowries she examined could be divided into two groups on the basis of certain characteristics of the radula and the female genital system. With the Plate 330. Female Cypraea (Zoila) friendii. a, semidia- thr grammatic drawing of animal with shell removed and with C. mantle lifted back; b, female reproductive tract; c, section of [10-458] help of senior zoology students at the University of Western Australia we studied the anatomy of friendii, venusta and decipiens in detail, paying particular attention to the reproductive system and the radula. Observations were also made on the anatomy of marginata and rosselli. The anatomy of friendii, type of the subgenus, is described here and comparisons are made with the other species of Zoila. Comparisons are also made with other cy- praeids, notably C. caputserpentis L., the anatomy of which was described in detail by Kay (1960a), and C. (S iphocypraea) mus L., which has been said to be closely related to Zoila. Materials and methods Living specimens of friendii and venusta were collected at Cockburn Sound and Sorrento Beach respectively, narcotized with isotonic magnesium chloride, fixed in formal-saline and preserved in 1 % propylene phenoxetol. Dissections were prepared of preserved specimens. Tissues for histological work were embedded in paraffin; serial sections were cut at 10 to 15 p. and stained with haemotoxy- lin and eosin or Triple Mallory’s. Specimens of de- cipiens were collected by pearl divers off Broome and were fixed and preserved in methylated alco- hol. Dissections and preparations of histological sections of this material were not entirely satisfac- tory due to the contraction and hardening of the tissues. For comparative purposes, dissections were pre- pared of specimens of Cypraea caputserpentis from intertidal reefs at Cape Vlaming, Rottnest Island, nigh mucous gland; d, section through mucous gland of venusta; e, section through vagina and bursa copulatrix 3. venusta. 3020IH2001 December 8, 1967 INDO-PACIFIC MOLLUSCA, vol. 1, no. 8 Cypraea 459 Western Australia. Two specimens of Cypraea mus, a male and a female, from 1-2 ft., Amuay Bay, Venezuela (courtesy of Dr. R. Tucker Abbott) were also examined. The mantle and the mantle cavity Vayssiere (1923, 1927) has described the mantle cavity of several cypraeids and Schilder ( 1936 ) published a condensed survey of the characteristics of the mantle throughout the family. Kay ( 1960b ) describes the mantle complex of caputserpentis L. The mantle cavity and associated organs of friendii (plate 330) is typical of the Cypraeacea and shows the results of regulative detorsion dur- ing development (see Naef, 1911 and Ghiselin and Wilson, 1966 ) . It is elongate with anterior incurrent and posterior excurrent siphons. The respiratory currents flow in a direct path from the incurrent siphon, across the tripartite osphradium, over the ctenidium and hypobranchial gland and out the excurrent siphon. The ctenidium is curved and lies across the path of the incoming respiratory currents with the osphradium nestling in the curve. The anus and genital apertures are situated posteriorly. The organization of the organs within the mantle cavities of venusta and decipiens show no notable differences. The mantles of most cowries bear simple or branched papillae and the margins of the siphons are denticulate, flabellate or serrate ( Schilder, 1936). In mus there are numerous conical or ten- tacular mantle papillae and the siphonal margins are serrate. The mantle of rosselli bears a few sim- ple wart-like or conical papillae but the margins of the siphons are smooth and entire. In friendii the mantle bears a few very small nodule-like papillae, only visible when the mantle is fully extended. Small black flecks on the mantle of decipiens may be similar nodular papillae in a contracted condi- tion ( the mantles of living specimens of that species were not examined ) . In venusta the mantle is thin and smooth, with- out papillae. There is no evidence of papillae on the mantle of preserved marginata or in a photograph of a living specimen taken by Mr. Max Cramer. The margins of the siphons are entire in all 5 species of Zoila. Female genital system The ovary of friendii is orange and forms a large part of the visceral whorl with an anterior extension on the left side as in other cowries. The renal ovi- duct runs from the ovary on the left side, laterally across the body to join the pallial oviduct. It is lined with a ciliated columnar epithelium thrown into deep folds, and enlarges before entering the pallial oviduct. A gono-pericardial duct runs parallel with the renal oviduct and appears to enter the latter a short distance before it enters the pallial oviduct. The gono-pericardial duct has an epithelium similar to that of the renal oviduct. As in other members of the superfamily Cyprae- acea (Fretter, 1941; Kay, 1960b; Ghiselin and Wil- son, 1966 ) , the pallial oviduct in friendii is a swollen tube with glandular walls and some diverticula. It is divided by a constriction into a proximal and a distal lobe. The proximal lobe has been called an albumen gland ( Kay, 1960b ) although Ghiselin and Wilson ( 1966 ) showed that this structure con- tains both albumen and membrane glands in the ovulid Cyphoma. The distal lobe has been called the capsule gland (Kay, 1960b) but Ghiselin and Wilson ( 1966 ) preferred the name mucous gland and this name is used here. Distally the mucous gland leads into a short vagina which enters the mantle cavity and terminates in a genital aperture just anterior to the anus. Plate 331. (left) Male Cypraea (Zoila) friendii showing ing details of prostate gland and seminal groove, position of penis on right side, (right) Enlargement show- [10-459] 460 Zoila B. R. Wilson and J. A. McComb Cypraeidae Plate 332 Cypraea (Zoila) friendii friendii Gray, showing variations in color patterns and shell shapes. Fig. 1, 2 fms., off Sor- rento Beach; 2, 6 fms., Eagle Bay, Cape Naturaliste; 3 and 4, Quindalup jetty, Geographe Bay; 5, Woodman’s Point, Cockburn Sound, on pilings; 6, Parmelia, Cockburn Sound, on pilings; 7, 4 fms., Easter Saturday Reef, Geographe Bay ( ANSP ) ; 8, 20-30 fms., 10 mi. south of False Entrance; 9, Esperance, on jetty pilings; 10 and 11, trawled in 60 fms., west of Dorre Island by Poole Bros., 1965. All, except fig. 7, in Western Australian Museum, (p. 10-474.) (About 2/3 natural size.) [10-460] December 8, 1967 INDO-PACIFIC MOLLUSCA, vol. 1, no. 8 Cypraea 461 Plate 333 Figs. 1-3. Cypraea ( Zoila ) friendii friendii forma contraria Iredale. 1, 80-100 fins., off Fremantle; 2, 90 fins., off Eyre (Delaware Natural History Museum); 3, 80-100 fins., 50 miles S.E. of Esperance, W.A. (p. 10-475.) Figs. 4-6. Cypraea (Zoila) friendii subspecies thersites Gaskoin. 4, Sir Joseph Banks Group South Australia; 5, Stansbury, St. Vincent Gulf, South Australia; 6, 20 ft., Lusby Rocks, Sir Joseph Banks Group, South Australia, (p. 10-476.) Figs. 7-10. Cypraea (Zoila) venusta Sowerby. 7, 6 fins.. Eagle Bay, Cape Naturaliste, W.A.; 8-10, 2-5 fms., V% mi. off Sorrento Beach, W.A. All, except fig. 2, in Western Austra- lian Museum, (p. 10-477.) (All 2/3 natural size.) [10-461] 462 Zoila B. R. Wilson and J. A. McComb Cypraeidae In cross section the albumen gland is oval with the dorsal, ventral and postero-lateral walls thick- ened by gland cells, but the antero-lateral wall is formed by a single layer which is an extension of the epithelium lining the lumen of the gland. This epithelium consists of ciliated columnar cells with basal nuclei. Beneath these columnar cells lie large, irregularly developed glandular cells from which ducts run to the lumen. The lumen of the albumen gland is simple with no major ciliated grooves or channels running along its length. The cream-colored mucous gland is contiguous with the albumen gland but is readily distinguished from the latter by both macroscopic appearance and histological detail of the epithelial lining. The mucous gland consists of thickened dorsal and ven- tral lobes, fused at their postero-lateral edges. In friendii the thin antero-lateral wall is formed by a continuation of the epithelium lining the lumen of the gland. This thin wall is folded so as to form a deep, strongly ciliated groove (the capsule duct of Kay, 1960b) running along the antero-lateral edge of the mucous gland between its dorsal and ventral lobes (plate 330, fig. c). The lumen of the mucous gland is lined with epithelium similar to that of the albumen gland; the wall is traversed by several longitudinal grooves. Beneath the epithelium are small gland cells forming strands a few cells wide alternating with ducts running to the lumen. The respective functions of the folded antero-lateral duct and the longitudinal grooves along the inner walls of the mucous glands are unknown. There are two sperm-containing diverticula of the pallial oviduct. Proximally there is a receptaculum seminis closely appressed to the dorsal side of the albumen gland and the proximal end of the mu- cous gland. Distally there is a large bursa copula- trix, a diverticulum of the vaginal duct within the mantle cavity. Macroscopically, the receptaculum seminis looks like a yellow granular mass. Microscopically, it is a mass of tubules embedded in a connective tissue. The epithelium of the tubules consists of tall, glandular, columnar cells with median nuclei. When stained with “Triple Mallory’s,” these cells are packed with yellow granules. The wide main duct of the receptaculum seminis enters the postero- lateral wall of the albumen gland at its extreme end distal to the ovary. The epithelium of this duct is much folded and composed of non-glandular, densely ciliated cells. Many tubules of the recep- taculum seminis contained spermatozoa; it could not be determined whether resorption was taking place there. The bursa copulatrix is a thick-walled, muscular sac attached along its whole length to the wall of the mantle cavity. It is situated very close to the genital aperture. The vagina enters and leaves the bursa copulatrix at its antero-ventral end (plate 330, fig. b). The female genital tracts of venusta and decipiens are similar to those in friendii but several minor differences were observed. The mature ovary of venusta is yellow and in decipiens it is bright- orange. The receptaculum seminis of venusta is a relatively large black, granular structure . In de- cipiens the passageway through the albumen gland is not simple but a channel is formed in the antero- lateral wall similar to that in the mucous gland. In venusta the fold of the thin antero-lateral wall of the mucous gland is displaced ventrally so that the duct comes to lie along the lateral edges of the ventral lobe (plate 330, fig. d) instead of medially between the two lobes as in friendii ( plate 330, fig. c) and decipiens. We have no data on the detailed structure of the female genital tract of marginata and rosselli. In venusta, decipiens and marginata there is a bursa copulatrix very like that of friendii. The bodies of only two specimens of rosselli have been examined; both were in a poor condition. No penis was present in either specimen and it is assumed they were both females. However, due to decompo- sition and damage, neither the female genital aper- ture nor the bursa copulatrix could be located. The bursa copulatrix at least should have been visible in spite of the damage; it seems likely that a bursa is absent in rosselli. Male genital system The mature testes are cream-colored and are situ- ated in the visceral whorl. In friendii, the coiled ampulla (seminal vesicle of Kay, 1960b) abuts broadly onto the anterior left side of the testes. The thin vas deferens leaves the medial regions of the ampulla and crosses the body just behind and be- low the pericardium emerging in the mantle cavity of the right side. In the mantle cavity it forms an open seminal groove which turns sharply anteriorly and runs forward to the penis on the right side of the head. The groove continues along the ventral surface of the penis to its pointed tip. In most respects the male genital tract is similar to that of caputserpentis. However, in caputserpen- tis there is a short duct from the testes to the am- pulla and the relative size and structure of the prostate gland is different. Kay ( 1960b ) stated that the prostate of caputserpentis is “closed.” By this [10-462] December 8, 1967 INDO-PACIFIC MOLLUSCA, vol. 1, no. 8 Cypraea 463 she means closed off from the mantle cavity by an expansion of the posterior wall of the seminal groove so as to cover the body wall (Kay, personal com- munication). We have checked this in male caput- serpentis from Rottnest Island, Western Australia. At the bend where the vas deferens opens into the seminal groove there is an extension and expansion of the posterior wall of the groove forming a wide flap covering the distal end of the vas deferens. Beneath the flap (i.e. between the flap and the body wall) is a cavity lined with ciliated, glandu- lar epithelium. The cavity, which appears to be homologous with the prostate gland of Trivia ( Fret- ter, 1941), is open along its anterior edge. In friendii the prostate, which is much smaller, is similarly formed by a flap-like extension of the posterior wall of the seminal groove ( shown in sec- tion, plate 331). Also in friendii the vas deferens projects as a short papilla into the prostate cavity (plate 331). The male genital tracts of venusta and decipiens were also examined in detail; that of marginata was examined superficially. No notable differences were observed between the males of three species and friendii. Radula The radula of the friendii species complex has been described and illustrated in detail by Wilson and Summers ( 1966 ) and the following description is taken from that paper. The radula is typical of Cpyraeidae in bearing 7 tricuspid teeth per row with 80 to 150 rows. The median tooth is slightly wider than it is high with one prominent medial cusp bordered by a smaller cusp on either side. An internal dumb-bell-shaped bract extending across the base of the tooth stains more densely in Chloro- zol-black than the remainder of the tooth and bears a pair of small denticles, one in each lower corner. A prominent semicircular bract, which does not take up the stain, projects downward from the base of the medial tooth. This latter element appears to be the structure Kay ( 1960a ) calls the “subtending bract.” The lateral tooth ( “admedian” of Kay ) is slightly higher than it is wide. It possesses a prominent medial cusp which is large and pointed, while the lateral cusps are relatively small. A densely staining internal bract also occurs in this tooth. Large denti- cles are present on the lower corners of the tooth, the one on the inner corner being particularly prominent and projecting toward the median tooth. The inner marginal tooth is similar to the lateral but is narrower with a more prominent central cusp and does not possess basal denticles. The outer marginal is still narrower, its lateral cusps are very reduced, no internal bract is visible, and there are no basal denticles. The shape and proportions of the median tooth and the inner marginal tooth vary slightly. In South Australian and Esperance specimens, the outer cusp on the inner marginal tooth forms a distinct shoulder. This is present but less distinct in Geog- raphe Bay and Sorrento samples. The specimens from Cockburn Sound ( plate 334, fig. b ) are excep- tional in that the inner marginal tooth is relatively narrow and there is no shoulder at the outer lateral cusp. This is consistent in specimens from this lo- cality but appears to be a local feature of that small population, not a regional characteristic. ( Cockburn Sound is only about twenty miles from Sorrento MEDIAN Plate 334. Radulae in the subgenus Zoila: a, marginata ; b, friendii; c, venusta (from Sorrento); d, decipiens ; e, ros- selli; and the subgenus Siphocypraea: f, mus. [10 - 463] 464 Z oila B. R. Wilson and J. A. McComb Cypraeidae Plate 335 Figs. 1-6. Cypraea (Zoila) decipiens E. A. Smith. 1-3, Port Hedland, Western Australia; 4-6, northern Western Australia (ANSP). (p. 10-482.) Figs. 7-9. Cypraea (Zoila) marginata Sowerby. 7, Abrolhos Islands, Western Australia (Delaware Natural History Mu- seum); 8, “The Flats,” south of the Abrolhos Islands, W.A.; 9, 28-30 fms., off Jurien Bay, W.A. (p. 10-483.) Figs. 10-12. Cypraea (Zoila) rosselli (Cotton). 10, Leigh- ton Beach, Western Australia (part of type lot from Rossell); 11, off Rottnest Island, W.A. (Delaware Natural History Museum); 12, 30-40 fms., west of Dirk Hartogs Island, W.A. In Western Australian Museum, (p. 10-485.) (All natural size. ) [10-464] December 8, 19 6' INDO-PACIFIC MOLLUSCA, vol. 1, no. 8 Cypraea 465 where animals occur with normal second lateral teeth. ) The radulae of venusta, decipiens and marginata resemble that of friendii in general; all have a sub- tending bract, basal denticles and an internal bract on the median tooth. However, diagnostic differ- ences in shape and relative size of the parts exist. These differences are given as diagnostic characters in the descriptions of the species and are evident in plate 334. The radula of rosselli is significantly distinctive in that there is no subtending bract on the median tooth. The basal denticles on the median and lateral teeth are very small, and the marginal teeth are stout with short medial cusps. The median tooth of mus also lacks a subtending bract, but in that spe- cies the basal denticles on the median and first lateral teeth are prominent and the outer laterals are particularly long and sickle-shaped (plate 334, fig. f). List of recognized taxa The fossil species are not considered in this paper. For a review of them, consult F. A. Schilder, 1935, Proc. Mai. Soc. London, vol. 21, pt. 6, pp. 336- 339 and in Archiv fur Molluskenkunde, 1941, vol. 73, p. 81. Genus Cypraea Linne, 1758 Subgenus Zoila Jousseaume, 1884 Recent: friendii Gray, 1831. Type subsp. thersites Gaskoin, 1849 marginata Gaskoin, 1849 venusta Sowerby, 1846 decipiens E. A. Smith, 1880 rosselli (Cotton, 1948) Fossil: jgigas McCoy, 1867 f mulderi Tate, 1892 f dorsata Tate, 1890 f gabrieli Chapman, 1912 f platypyga McCoy, 1876 f consobrina McCoy, 1877 f toxorhyncha Tate, 1890 f gendinganensis Martin, 1899 ) Uchilderi (Dey, 1962) f simplicior (Schilder, 1935) Affinity between the five living species of Zoila The living species of Zoila have many distinctive morphological characters in common, although some of the characters common to friendii, marginata, venusta and decipiens are not shared by rosselli. Females of the former four species possess a large bursa copulatrix and a glandular seminal receptacle (although this is not confirmed for marginata ); in males the prostate gland is small (relative to that of C. caputserpentis); the ampulla abuts directly onto the anterior end of the testis (not confirmed for marginata ); the median tooth of the radula bears a subtending bract, an internal bract and basal 'denticles and the first lateral tooth bears a prominent basal denticle on the inner side; margins of the siphons are entire and the mantle is thin and may be quite smooth ( venusta, marginata ) or bear very small, simple papillae ( friendii and possibly decipiens). In shells of all four species the aper- ture dentition is weak compared to that of many other cowries, especially on the columellar side; the fossula is quite deep, but the anterior columellar teeth do not extend up onto it; the terminal ridge is obsolete, the right side of the anterior channel (ventrally) being a flat or sloping surface with a simple cornered inner edge. The spire is elevated in friendii and marginata, but only slightly elevated in venusta and almost depressed in decipiens. Thus this characteristic, often used as a diagnostic feature of Zoila, has only limited application. In addition to these morphological characters it has been observed that friendii, venusta and de- cipiens feed on sponges and commonly may be found perched on the sponges in the open; none show any tendency to bury themselves within the sponges. There is evidence that marginata may be- have and feed in the same way. As far as we are aware this kind of feeding and behavior is unique among cypraeids. No other cypraeid is known to have such a direct association with sponges. Most cowries are secretive and hide under stones, among corals or in crevices (at least during daytime). Some, like C. tigris, may be found in the open in coral pools, but are more often hidden. This aggregate of anatomical, behavioural and shell characters suggests a close evolutionary affinity between friendii, venusta, marginata and decipiens. Therefore we propose to retain the name Zoila as a distinct taxon. However, we do not believe that the distinctive characters of the taxon are sufficient to justify generic rank. We rank Zoila as a sub- genus of Cypraea. [10-465] 466 Z oila B. R. Wilson and J. A. McComb Cypraeidae Whether rosselli should be included in Zoila is uncertain. In common with the other species the siphonal margins are smooth, the animal probably lives and feeds on sponges, and the anterior colu- mellar teeth of the shell do not extend onto the fossula. However, rosselli differs from the other species by the presence of small but distinct pa- pillae on the mantle; the absence of a subtending bract and basal denticles on the median tooth of the radula; the small size of the basal denticle on the inner side of the first lateral radula tooth; and by the more strongly denticulated shell. There is doubt whether females of rosselli pos- sess a bursa copulatrix. If, when well-preserved ani- mals of rosselli become available for study, a bursa copulatrix is shown to be present, then it may be concluded that a relationship with the other four living species of Zoila probably does exist, even though rosselli is the most outstanding of the group. If the females of rosselli do not possess a bursa copulatrix, retention of this species within the same subgenus would be difficult to justify. Relationships of the subgenus Zoila While a close affinity between the living species of Zoila is strongly indicated by the data ( rosselli being a possible exception ) , the relationship of this species group to other cowries is yet to be deter- mined. Information on the anatomy and behaviour of many living cowries is still wanted. One interesting result of this study has been the discovery that the living species of Zoila do not conform to either of the two categories recognised on anatomical ground by Kay ( 1960a ) . Kay recog- nised four radula types (R1 to R4) among the spe- cies she studied. Those species in which the fe- males possessed a bursa copulatrix and a glandular receptaculum seminis invariably had radula types R2, R3, or R4; those species without a bursa and with a saccate receptaculum seminis had radula type Rl; this is the dichotomy shown by Kay to cross previous concho logical classifications. Evi- dence presented in this paper demonstrates that, with the exception of rosselli, females of Zoila spe- cies possess a bursa, a glandular receptaculum seminis, and a radula resembling Kay’s type Rl (i.e. with a subtending bract and an internal bract on the median tooth). This contradiction indicates that a simple two- way split of the Cypraeidae on the basis of the fe- male genital system and radula type cannot be maintained as a taxonomic division of the family. However, the fact that four of the five species of Zoila differ in the same ways from the fifty-two species studied by Kay, is additional evidence sup- porting the phyletic distinctness of Zoila. The subgenus Zoila has generally been consid- ered closely related to Bernaya Jousseaume, 1884 (Upper Jurassic to Recent) and more distantly to Siphocypraea Heilprin, 1887 (Miocene to Recent). The taxonomic arrangement of these groups and other fossil relatives has varied (see Schilder and Schilder, 1939; Wenz, 1941; Schilder, 1941). Re- cently Schilder ( 1965 ) ranked Zoila, Bernaya and Siphocypraea as equal genera within the subfamily Cypraeorbinae. We propose to treat these groups as subgenera of Cypraea. The anatomy of Siphocypraea mus does not indi- cate a very close affinity with Zoila. The mantle bears numerous, simple papillae and the margins of the siphons are filamented. The female has a large bursa copulatrix but this does not adhere along its whole length to the wall of the mantle cavity as in Zoila. In mus the median tooth of the radula has an internal dumb-bell-shaped bract, but no subtending bract and the basal denticles are prominent and pointed (pi. 334, fig. f). The lateral tooth bears a prominent basal denticle on the inner corner simi- lar to Zoila but the marginals are very long and sickle-shaped. Comparative anatomical data on the two living species of Bernaya (i.e. fultoni Sowerby, 1903 and teuleri Cazenavette, 1846) would be most useful, but these species are seldom collected, and living or preserved specimens are not available for study. The diagnostic shell characters separating Zoila from Bernaya are poorly defined in the literature. The descriptions given by Wenz ( 1941, pp. 969 and 971) for the two genera contain only a few rather subjective differences. In Bernaya the sides are said to be rounded, the ventral surface convex, concave at the front, the spire short, the teeth somewhat protracted, the outer lip sloping concavely at the front and the fossula broad, concave, smooth. In Zoila the sides are said to be angular, the base flat, the spire “usually visible,” the teeth coarser, the outer lip less sloping and the fossula fairly shallow but distinct. In these circumstances, Schilder ’s (1963a, p. 127) proposal to attribute the unique Abrolhos Island shell he named catei to the genus Bernaya is full of interest. He adopted this procedure because “. . . the symmetrical profile in lateral view, the less angular margins, the straight aperture, the more primitive outlets, the well developed columellar teeth sloping inward, etc., seem to point rather to [10-466] December 8, 1967 INDO-PACIFIC MOLLUSCA, vol. 1, no. 8 Cypraea 467 the genus Bernaya than to Zoila ( Schilder, 1941, pp. 80-81).” We consider the specimen to which Schilder referred to be an aberrant form of vemista and synonymize it as such. It would be possible to take exception to all the definitive characters listed by Schilder as diagnostic of Bernaya, on the grounds that they occur inconsistently among living and fossil Zoila as well. Nevertheless the point is made that the species of Zoila and the species of Bernaya overlap with regard to many shell characters. In fact, Schilder ( 1963a, p. 128 ) concludes that “The recently discovered Bernaya catei seems to repre- sent another link connecting Bernaya with Zoila, so that one should consider degrading Zoila to a sub- genus of Bernaya .” In this paper Zoila and Bernaya are each considered to be subgenera of Cypraea so that such a procedure could not be implemented without eliminating Zoila altogether ( Bernaya has priority). While appreciating the probable close affinity of these two groups we feel that no nomen- clatural step of this nature should be made at pres- ent, because the diagnostic shell characters remain too ambiguous and poorly defined. A thorough and critical review of the fossil species of both sub- genera, and data on the anatomy of living Bernaya are needed. Better understanding of the relationships between fossil and living species of these subgenera would be particularly interesting from the palaeogeo- graphic point of view. Bernaya is known from the Upper Jurassic to the Recent (“Subgenus” Proto- cypraea, see Schilder, 1963a, p. 127) and species of Zoila are known from Australian specimens as old as the Miocene and from the Upper Miocene to Pleistocene of India and the Indo-Malay Archi- pelago. Schilder (1963a, p. 128) notes that the upper Tertiary and Pleistocene species of Zoila in these tropical Asian regions connect the Australasian Zoila with the Tethyan Bernaya. Whether the two subgenera are considered to be distinct or synony- mous, a close relationship seems highly probable and the living Australian species of Zoila may be regarded as surviving relics of a once-flourishing group of Tethyan- Australasian cowries. Links of this kind between the early Tertiary molluscan fauna of the Tethyan region and the Tertiary and Recent fauna of southern Australasia have been demonstrated in a number of molluscan groups (see Fleming, 1957; Ludbrook, 1954). Additional documentation of such links may greatly improve our understanding of the origin and high degree of endemism in the southern Australian and New Zea- land marine faunas. Gigantocypraea Schilder, 1927, has been treated as a subgenus of Zoila. We cannot follow this pro- cedure, because we treat Zoila itself as a subgenus of Cypraea. The status of Gigantocypraea must re- main in doubt until the fossil species are revised. In may eventually be placed into synonymy with Zoila or accepted as a subgenus of equal rank with Zoila, Bernaya and Siphocyprasa. Acknowledgments This study began in 1963 at the University of Western Australia as a class exercise in applied morphology. The exercise was supervised by Dr. E. P. Hodgkin and to him and senior zoology stu- dents of the 1963 class, we extend our thanks for their interest and help. The scope of the investiga- tion has since been considerably extended by the authors. Most of the nomenclatural and literature research was done by the senior author while Re- search Fellow in Malacology at the Museum of Comparative Zoology, Harvard University. We are grateful to Dr. W. J. Clench and Dr. R. D. Turner for advice and assistance in these matters. Mr. C. N. Cate and Mr. Ray Summers also contributed help- ful suggestions. For information on types we are grateful to Dr. Anna M. Bidder of Cambridge University, Mr. Norman Tebble of the British Museum (Natural History), Dr. D. F. McMichael of the Australian Museum, Miss J. Hope Macpherson of the National Museum of Victoria and Dr. Helene Laws of the South Australian Museum. We wish also to thank the shell collectors who provided helpful informa- tion and field assistance. Mr. Trevor Sutcliffe of Perth and Mr. Max Cramer of Geraldton gener- ously presented us with the soft parts of C. margi- nata, and Mrs. Phyllis McDaniel of Broome pro- cured a series of preserved C. decipiens for our study. Drs. R. W. George, W. G. Inglis and R. Tucker Abbott read and criticised early drafts of the manu- script. We gratefully acknowledge their assistance, but accept all responsibility for errors of fact ox- judgment that may occur in this paper. The abbreviation WAM in this paper refers to the Western Australian Museum, Perth. [10-467] 468 Zoila B. R. Wilson and J. A. McComb Cypraeidae Selected Bibliography Allan, Joyce. 1956. Cowry shells of world seas. Georgian House, Melbourne, pp. 1-170, pis. 1-15. Brazier, John. 1871. Distribution and geographic range of cowries in Australasia. Sydney Mail, Dec. 2, 1871 [pub- lished in a daily newspaper and subsequently issued as a reprint in 1882— see Gatliff (1916) and Cate (1962, 1964)]. Cate, Crawford N. 1961. Rediscovery of Cypraea marginata Gaskoin, 1848. Veliger, vol. 3, no. 3, pp. 76-78, pi. 14. Cate, Crawford N. 1962. Comparison of two rare cowry spe- cies (Gastropoda). Veliger, vol. 5, no. 1, pp. 6-14, pis. 1-4, 2 text-figs. Cate, Crawford N. 1964. Western Australian cowries (Mol- lusca: Gastropoda). Veliger, vol. 7, no. 1, pp. 7-29, pi. 5, map. Cotton, Bernard C. 1948. Southern Australian Gastropoda. Part III. Trans. Royal Soc. South. Australia, vol. 72, pt. 1, pp. 30-32, pi. 1. Cox, James C. 1889. Note on Cypraea venusta (Sowerby), Proc. Linn. Soc. N.S.W., second series, vol. 4, pt. 1, p. 187, pi. 15. Fleming, Charles. 1957. The genus Pecten in New Zealand. Palaeont. Bull. Wellington, N.Z., vol. 26, pp. 1-6, figs. Fretter, Vera. 1941. The genital ducts of Theodoxus, Lamel- laria, and Trivia, and a discussion of the evolution in the prosobranchs. Jour. Mar. Biol. Assoc. U.K., vol. 26, pp. 312-351. Gaskoin, J. S. 1849. Description of new species of the genus Cypraea. Proc. Zool. Soc. Lond. [for 1848], pp. 90-98. Ghiselin, Michael T. and Barry R. Wilson. 1966. On the anatomy, natural history, and reproduction of Cyphoma, a marine prosobranch gastropod. Bull. Mar. Sci. Gulf Caribb., vol. 16, no. 1, pp. 132-141. Iredale, Tom. 1939. Australian cowries, Part II. Aust. Zool., vol. 9, pt. 3, pp. 297-323, pis. 27-29. Kay, Alison. 1960a. Generic revision of the Cypraeinae. Proc. Malac. Soc. Lond., vol. 33, pp. 278-287. Kay, Alison. 1960b. The functional morphology of Cypraea caputserpentis L. and an interpretation of the relations among the Cypraeacea. Int. Revue Ges. Hydrobiol. Hydrogr., vol. 45, no. 2, pp. 175-196. Laws, H. M. 1966. Zoila marginata (Mollusca, Cypraeidae) in South Australia. Records South Australian Mus., Ade- laide, vol. 15, no. 2, pp. 251-256. Ludbrook, N. H., 1954. The molluscan fauna of the Pliocene strata underlying the Adelaide Plains. Part I. Trans. Royal Soc. South. Australia, vol. 77, pp. 42-64. Naef, A. 1911. Studien zur generellen Morphologie der Mol- lusken. 1. Teil: fiber Torsion und Asymmetrie der Gastro- poden. Ergebn. Fortschr. Zool., vol. 3, pp. 73-164. Schilder, F. A. 1930. Beitrage zur Kenntnis der Cypraeacea (Moll., Gastr. ). III. Zool. Anz., vol. 92, p. 74. Schilder, F. A. 1936. Anatomical characters of the Cyprae- acea which confirm the conchological classification. Proc. Malac. Soc. London, vol. 22, pt. 2, pp. 75-112, 2 pis. Schilder, F. A. 1941. Verwandtschaft und Verbreitung der Cypraeacea. Arch. Molluskenk., vol. 73, pp. 57-120, pi. 8, fig. 10. Schilder, F. A. 1960. Zur Kenntnis der Cypraeidae. Arch. Molluskenk., vol. 89, pp. 185-192. Schilder, F. A. 1961. New Cowries described since 1938. The Cowry, vol. 1, no. 2, pp. 1-5. Schilder, F. A. 1963a. Further remarks on two rare cowry species (Gastropoda). Veliger, vol. 5, no. 4, pp. 125-128. Schilder, F. A. 1963b. Lumpers and splitters. Veliger, vol. 6, no. 2, pp. 104-110. Schilder, F. A. 1965. The geographic distribution of cowries. Veliger, vol. 7, no. 3, pp. 171-183. Schilder, F. A. 1966a. Zoila venusta and Zoila episema. Hawaiian Shell News, for June, 1966, vol. 14, no. 8, p. 5, text-fig. Schilder, F. A. 1966b. Zoila friendii (Gray) in Shark Bay. Hawaiian Shell News, for August, 1966, vol. 14, no. 10, p. 4, text-fig. Schilder, F. A. 1966c. Notes on some species of Cypraeidae. Hawaiian Shell News, for Dec., 1966, vol. 14, no. 14, p. 4, figs. 1-5. Schilder, F. A. and M. Schilder. 1938-1939. Prodrome of a monograph on living Cypraeidae. Proc. Malac. Soc. Lon- don, vol. 23, nos. 3-4, pp. 119-231. Serventy, D. L. 1937. Zoological notes on a trawling cruise in the Great Australian Bight. Jour. Proc. Royal Soc. West Australia, vol. 23, pp. 65-87, pis. 1-4. Steadman, W. R. and B. C. Cotton. 1946. A key to the classification of cowries ( Cypraeidae ) . Records South Australian Mus., vol. 8, no. 3, pp. 503-528, pis. 8-13. Stokes, Alf. 1966. Range of Zoila marginata extended. Ha- waiian Shell News, for August, 1966, vol. 14, no. 10, p. 5, text-fig. Trenberth, Paul. 1962. Observations on Zoila thersites Gas- koin, 1849. Hawaiian Shell News, vol. 11, no. 2, p. 8, Vayssiere, A. 1923. Recherches zoologiques et anatomiques sur les mollusques de la famille des Cypraeides. Ann. Mus. Hist. nat. Marseille, vol. 18, pp. 1-120. Vayssiere, A. 1927. Recherches zoologiques et anatomiques sur les mollusques de la famille des Cypraeides. He Partie. Ann. Mus. Hist. nat. Marseille, vol. 22, pp. 133-184. Verco, Joseph C. 1918. Notes on South Australian marine mollusca, with descriptions of new species. Trans. Royal Soc. South. Australia, vol. 36, pp. 209-210. Wenz, W. 1941. [In]: Schindewolf, O. H., Handbuch der Palaozoologie, vol. 6, Gastropoda (Berlin, 1938-41), xii plus 1639 pp.; 4211 text figures. Wilson, B. R. and Ray Summers. 1966. Variation in the Zoila friendi ( Gray ) species complex ( Gastropoda; Cy- praeidae) in South-Western Australia. Jour. Malac. Soc. Australia, no. 9, pp. 3-24, pis. 1-4, 4 text figures. [10-468] December 8, 1967 INDO-PACIFIC MOLLUSCA, vol. 1, no. 8 Cypraea 469 Subgenus Zoila Jousseaume, 1884 Type: Cypraea friendii Gray, 1831 This subgenus is represented by five living spe- cies on the continental shelf of Western Australia and South Australia. The centre of distribution is the central coast of Western Australia, although two species are known to extend into South Aus- tralian waters and one species is apparently re- stricted to the warm waters of the far northwestern part of Western Australia. Fossil species have been described from the Upper Miocene to the Recent age in southeastern Australia, and from the Upper Miocene to the Pleistocene of India and the Indo- Malay archipelago. The shell characters diagnostic of Zoila are the elevated or at least visible spire, the weak columel- lar teeth which may extend only part of the way along the columellar side of the aperture, and the concave, smooth, rather shallow but distinct fossula. Wenz (1941, pp. 969 and 971) separates Zoila from Bernaya by the more angular sides, flatter base, coarser teeth, and less sloping outer lip. These differences need further study in view of the inter- mediate shell characteristics of venusta (see Schil- der, 1963a, p. 127). A number of anatomical features separate Zoila from other living cowries. In females there is a large bursa copulatrix; in males the ampulla abuts directly onto the anterior end of the testis; there are subtending internal bracts and basal denticles on the median tooth of the radula; the mantle is smooth and bears only very small papillae, if any; the siphonal margins are smooth and entire. Little is known about the anatomy of rosselli, but obser- vations on the mantle and radula of that species place its position in Zoila in doubt. Synonymy — 1883 Zoila Jousseaume, Le Naturaliste, Paris, 1st series, vol. 2, 6th year, no. 52, p. 414 (Feb. 15, 1883). No type given; 1883 (April?), Bull. Soc. Zool. France, vol. 9, p. 89. Type by original designation: Cypraea scottii Brod. Key to the living species of Zoila based on shell characters a. Sides spotted b a. Sides unspotted c b. Medial columellar teeth well-developed . . marginata Gaskoin b. Medial columellar teeth absent or obsolete d c. Margins and siphonal canals sharp rosselli Cotton c. Margins and siphonal canals rounded . . decipiens E. A. Smith d. Siphonal canals elevated and sharp friendii Gray d. Siphonal canals low and rounded venusta Sowerby [10 - 473] 470 Z oila B. R. Wilson and J. A. McComb Cypraeidae Cypraea friendii subspecies friendii Gray, 1831 (Pis. 332 and 333) Range — Western Australia, from Done Island to the western part of the Great Australian Bight. Remarks - The variation of the friendii complex has been studied and described by Wilson and Summers (1966) who concluded that this is a highly variable polytypic species with two allo- patric populations worthy of recognition as sub- species. The Western Australian population (friendii friendii) occurs across the whole width of the conti- nental shelf. Shells from the outer parts of the shelf tend to be much wider than shells from more shal- low adjacent water, and there is also a geographic ( clinal ) increase in width from north to south. This results in a complex situation where the width of the shells of any given population will be deter- mined by both the geographic position and the depth. It was suggested that the causal factor may be water temperature, i.e. the lower the environ- mental mean water temperature the more swollen the last shell whorl becomes. In addition, beyond 70 fathoms or so, the shells lose then- color and may be entirely white, or white with brown or orange spots. At the northern end of the range, shells tend to have faint crenulations or teeth along the colu- mellar side of the aperture and are more pyriform than southern shells. Other characters such as shell size, elevation of the spire, width of the aperture and the degree of elevation of the anterior and pos- terior channels vary either clinally or at random between local populations and thus have no taxo- nomic value. Schilder ( 1966b ) referred to a “unique” speci- men from off False Entrance, recorded by Wilson and Summers (1966), apparently overlooking the fact that ten further specimens from off Dorre Is- land ( Shark Bay ) were recorded in an addendum of the same paper. Schilder suggested that these northern end-of-cline variants may deserve a new racial name. We oppose this suggestion. In view of the complex variation of so many shell characters within the Western Australian popula- tion as a whole, there is little to be gained from sub-dividing it into subspecies. The extreme variability of friendii makes it diffi- cult to give an adequate description. Nevertheless, other species possess sufficient diagnostic charac- ters so that confusion of any of them with friendii is unlikely. Habitat — The full width of the continental shelf from depths of a few feet to at least 100 fathoms. The animal feeds on sponges and they are com- monly taken from sponges growing on rocks, jetty pilings or on weed flats. They do not hide within the sponges, but are found on top or on the sides of the sponges, or sometimes crawling on sand or among weeds on the ocean floor. Description — Shell ovate ( southern end of range and deeper water ) to elongately ovate-pyriform ( in shallow water near northern end of range). Dorsal hump relatively low ( except in extreme deep-water south-coast populations) and behind the centre of the shell. Margins more or less rounded and margi- nated only at the extremities ( more strongly margi- nated in wider shells ) ; right posterior margins sometimes irregularly indented. Sides of the ante- rior and posterior channels sharp and usually greatly elevated. Spire elevated and often produced beyond the posterior channel lips; the spire may project into the posterior channel or lie to the left side of it. Aperture narrow, slightly curved posteriorly (more strongly so in wider south-coast shells ) , and dilated anteriorly. Aperture dentition of shell — Labial teeth num- ber from 14 to 30 depending on shell size; usually rounded but short; bifid labial teeth are not uncom- mon, especially in the Esperance population. Colu- mellar teeth greatly reduced; only 4 to 8 columellar teeth present anteriorly, although in shells from the northern end of the range there are usually addi- tional faint crenulations or teeth all along the colu- mellar side of the aperture; anterior columellar teeth short and do not extend up onto the deep and concave fossula. Anterior terminal ridge obsolete. Color of shell — Extremely variable. Dorsally, shallow-water shells have a white ground color with four radial bands of pale ash-blue blocks over- lain by varying degrees of large, irregular and un- usually confluent chocolate-brown blotches. Sides bear large, more or less round and discrete spots on a uniform dark chocolate-brown ground color. Base uniformly dark chocolate-brown. In shells from depths greater than 70 fathoms the brown markings become greatly reduced in number and color in- tensity; usually only a few pale brown spots or flecks remain on the dorsum and the base and sides [10-474] December 8, 1967 INDO-PACIFIC MOLLUSCA, vol. 1, no. 8 Cypraea 471 are pale brown; some shells are entirely off-white, without markings. Measurements of shell — Individual size varies greatly between populations. The population with the smallest known individuals is in very shallow water at Quindalup in Geographe Bay ( mean length 50.8 mm.; range 48.4 mm. to 52.9 mm.) and the largest individuals have been found at Sorrento about 20 miles north of Fremantle (mean length 91 mm.; range 85 mm. to 99 mm.). Relative width is a clinal character, increasing both horizontally from north to south and east, and vertically with increas- ing depth. For details of shell dimensions and their varia- tions see Wilson and Summers ( 1966 ) . Soft parts — Mantle, anterior and posterior siphons, foot, head and eye-tentacles are all black; a trans- verse groove across the truncated anterior end of the foot is yellow and a yellow streak may extend a short distance posteriorly down the antero-ventral edges of the foot. The radula and reproductive anatomy of this species are described in detail in a preceding section (p. 458). See plates 330 and 331. Synonymy — 1831 Cypraea friendii Gray, Zool. Miscell, p. 35. New Hol- land, near Swan River [type locality here restricted to Cockburn Sound and Owen Anchorage between Garden and Carnac Islands and the mainland]. Lectotype, selected by S. P. Dance, Brit. Mus. (Nat. Hist.), no. 1964777. 1831 [July, 1832] Cypraea scoitii Broderip, Zool. Jour., vol. 5, p. 330 (Strait of Sunda, near Angia, Java). Sowerby, 1837, corrected the locality to Garden Is- land, Swan River. Type not located. 1912 Cypraea thersites Gaskoin, Verco, Trans. Roy. Soc. So. Australia, vol. 36, pp. 209-210 (in part, i.e., three specimens dredged from 78-100 fms. west of Eucla, Great Australian Bight). 1918 Cypraea friendii friendii Gray, Verco, Trans. Roy. Soc. So. Australia, vol. 42, pp. 144-145. 1930 Zoila friendii vercoi Schilder, Zool. Anzeiger, Leipzig, vol. 92, p. 74 (“westliche Siidkiiste von Australien; Typus: das mittelgrosse [South Aust. Mus. no. D14124], von Verco 1918, Trans. Roy. Soc. So. Australia, vol. 42, p. 147, erwahnte Stuck von Esperance ) . 1935 Zoila thersites contraria Iredale, Australian Zool., vol. 8, p. 107 ( 72-100 fms. west of Eucla, Great Aus- tralian Bight). The types are the specimens col- lected by Verco (Verco, 1912). Lectotype, here selected, Australian Mus. Sydney no. C35580; also two paralectotypes Australian Mus. no. E3848 and E3839. 1938 Zoila friendii friendii Gray, Schilder and Schilder, Proc. Malac. Soc. Lond., vol. 23, p. 174. 1938 Zoila friendii contraria Iredale, Schilder and Schilder, Proc. Malac. Soc. Lond., vol. 23, p. 174. 1962 Cypraea contraria Iredale, Griffiths, The Cowry, vol. 1, pp. 36-38. Types and type localities — Gray gave as type locality “New Holland near Swan River [Swan River Colony, not the Swan River itself].” This has been further restricted by Wilson and Summers ( 1966 ) to Cockburn Sound between Owen Anchor- age and Garden Island, where a large population of C. (Z. ) friendii friendii exists. The lectotype, identified by S. P. Dance, British Museum (Nat. Hist. ) has no. 1964777. The type of Schilder s southern race, Z. friendii vercoi, is one of the specimens described by Verco ( 1918 ) and is now in the South Australian Museum, no. D14124, type locality Esperance. The type locality for scottii was given as Angia, Java. Is., but this is evidently an error. Three specimens in the Australian Museum repre- sent the type series of Iredale’s subspecies Z. ther- sites contraria; lectotype no. C 35580; type locality 72 to 100 fathoms west of Eucla, Great Australian Bight. Plate 336. Geographical distribution of Cypraea (Zoila) southwest Australia. friendii friendii (solid dots) and C. (Z. ) friendii thersites in [10-475] 472 Zoila B. R. Wilson and J. A. McComb Cypraeidae N omenclature — In older texts this species ap- pears under the name C. scottii Broderip, 1832, but Gray’s name has priority by a few months and is now generally used. Schilder ( 1930 ) proposed the subspecies name Z. friendii vercoi for specimens from Esperance on the south coast of Western Aus- tralia, described by Verco (1918). Wilson and Sum- mers ( 1966 ) have rejected this name after an analy- sis of the diagnostic characters on which it was based. The white or pale form from the outer edge of the continental shelf in the western part of the Great Australian Bight was given the subspecies name Z. thersites contraria by Iredale, 1935. How- ever, Wilson and Summers ( 1966 ) have shown that these pale shells are deep-water ecotypes of the south-coast friendii and that similarly pale shells (but not as wide) occur in similar depths off the west coast of Western Australia. For these reasons deep-water pale forms can have no taxonomic recog- nition and the name contraria must be rejected. Specimens dissected — Fifteen animals from Par- melia Bank, Cockburn Sound. The radulae of twenty specimens were examined from the same sample; other radulae were examined from specimens from Sorrento, Geographe Bay and Esperance. Records — WESTERN AUSTRALIA: W. of Done Id. ( trawled, 60 fins., Poole Bros, on L.F.B. “Bluefin,” see Wil- son and Summers, 1966 ) ; off Geraldton ( craypot, 20-25 fms.); off Dongara (craypot, 32 fms.); between Fremantle and Geraldton ( pale form, trawled “Endeavour,” 1912, 80- 100 fms.); 20 mi. N.W. of Rottnest Id. (pale form, pieces only, dredged B. R. Wilson on L.F.B. “Bluefin,” 85-110 fms.); 1 mi. N. Bathurst Light, Rottnest Id. (17 fms.); off Sorrento Beach (2-3 fms., on sponge, B. R. Wilson and R. W. George); Parmelia Bank, Cockburn Sound (1-4 fms., on sponge, jetty pilings and among sea-grass, B. R. Wilson, leg); Quindalup, Geographe Bay (small form, 0-1 fm., B. R. Wilson); off Quindalup, Geographe Bay (2-8 fms., B. R. Wilson); Busselton, Geographe Bay (jetty piles, 1-3 fms., B. R. Wilson); King George Sound; Bremer Bay; Esperance (jetty piles, 1-4 fms., B. R. Wilson) (all West. Aust. Mus.); 60 mi. W. of Eucla [probably S.W.], Great Australian Bight (pale form, trawled 72-100 fms., Aust. Mus.); 75 mi. S. of Eyre, Lat. 33°15'0" S., Long. 126°22T5" E., Great Australian Bight (pale form, trawled 93 fms., S.S. “Bonthorpe,” see Serventy, 1937, p. 77, pi. 4); S. of Eyre, Lat. 33°20' S., Long. 126° E., Great Australian Bight (pale form, trawled, Ray Summers collection ) . A complete list of specimens examined is given by Wilson and Summers (1966). Cypraea friendii subspecies thersites Gaskoin, 1849 (PI. 333, figs. 4-6) Range — Streaky Bay, eastward to Beachport in South Australia. Remarks — South Australian populations seem to be isolated from those of southern Western Aus- tralia, although insufficient collecting has been done along the shores of the Great Australian Bight to be sure of this. South Australian shells are very similar to shallow-water Esperance shells in dorsal color, but are wider. On the ventral surface they differ in having a white patch along each side of the aperture and usually in having two to four weak posterior columellar teeth. No specimens are recorded from the outer edge of the continental shelf in South Australian waters. No differences were observed in the soft parts of typical friendii and its eastern subspecies, thersites. Habitat — Mr. Paul Trenberth ( 1962 and per- sonal communication) reports that the habitat is much the same as that described above for friendii friendii. The deepest South Australian record is 40 fathoms. Description — Shell ovate. Roundly humped, with the maximum height behind the centre. Base flat- tened; margins slightly angular at the extremities. Sides of the anterior and posterior channels sharp but only slightly elevated (like deep-water speci- mens from the south coast of Western Australia). Spire evident but only slightly projecting and usu- ally lying within the posterior channel. Aperture curved and dilated anteriorly. Aperture dentition — same as friendii friendii, except that two, three, or four weak columellar teeth are often present at the posterior end. Color of shell — same as shallow- water friendii friendii, except that very dark shells are rather more common and the base has a white patch along each side of the aperture. Measurements of shell — Average length 73.3 mm.; range 63 6 to 86 mm. For detailed analysis of di- mensions, see Wilson and Summers ( 1966 ) . Synonymy — 1849 Cypraea thersites Gaskoin, Proc. Zool. Soc. London, for 1848, vol. 16, p. 90. Hab. ? [Roberts, 1885 (in) Tryon, Man. of Conch, gives South Australia]. Lectotype, selected by S. P. Dance, Brit. Mus. (Nat. Hist.), no. 18465151, bears no locality. 1918 Cypraea friendii thersites Gaskoin, Verco, Trans. Roy. Soc. So. Australia, vol. 42, pp. 144-148. 1935 Zoila thersites thersites Gaskoin, Iredale, Australian Zool, vol. 8, p. 107. 1938 Zoila friendii thersites Gaskoin, Schilder and Schilder, Proc. Malac. Soc. London, vol. 23, p. 174. 1961 Zoila thersites Gaskoin, Schilder, The Veliger, vol. 4, p. 110. Types and localities — G askoin (1849) gave no locality, but Roberts, 1885, gave South Australia. The lectotype selected by S. P. Dance, British Mu- seum (Nat. Hist.) no 1846.5.15.1 bears no locality. [10 - 476] December 8, 1967 INDO-PACIFIC MOLLUSCA, vol. 1, no. 8 Cypraea 473 Nomenclature — Several authors (Verco 1918; Schilder and Schilder 1939) regarded South Aus- tralian populations as a subspecies of the western frienclii, but in recent years others have regarded them as a distinct species. The reasons for this were considered to be the much greater relative width of the shells, the lower spire, less upturned sides of the siphonal channels and the white base. Wilson and Summers ( 1966 ) reverted to the earlier proce- dure and recognised only subspecific rank for the South Australian populations. They showed that relative width is not a suitable diagnostic character because of the two-dimensional clinal variation it exhibits throughout the range of the species com- plex. Most other shell characters are either incon- sistent or are determined by shell width. Only the white patch on the base and the frequent presence of posterior columellar teeth are useful diagnostic characters. Specimens dissected — One male and two female specimens from Sir Joseph Banks Group, Spencer Gulf, South Australia. Records - SOUTH AUSTRALIA: Lusby Rocks, Sir Joseph Banks Group, Spencer Gulf, South Australia (P. Trenberth, leg; WAM); Thistle Id., Spencer Gulf (Nat. Mus. Viet.); Corny Pt., Yorke Peninsula, Spencer Gulf (NMV); Stans- bury, Gulf St. Vincent (WAM); Ardrossan, Gulf St. Vincent (WAM). Cypraea venusta Sowerby, 1848 (Pis. 333, 337-340) Range — Abrolhos Islands to the western part of the Great Australian Bight, Western Australia. Remarks — This is a variable species, once con- sidered among the world’s rarest cowries. From the beginning there has been controversy and confu- sion about the identity of the species and its distri- bution, because two apparently distinct color forms occur: one has a pale-yellow or fleshy-pink shell with yellow or pale-brown spots, and the other has a darker shell with dark-brown or tan markings. The holotype of venusta and the British Museum (Natural History) paratype of thatcheri have been compared by Dr. Anna Bidder of Cambridge Uni- versity (personal communication) who pronounced them very similar in all respects. From previous illustrations, they and the holotype of thatcheri in the Dautzenberg collection, typify the pale form, which Schilder ( 1963a, 1966a ) regards as the “typi- cal” form of venusta. The specimen described and figured by Cox, 1889, and later made the type of episema by Iredale, 1939, typifies the dark form. The dark form is known from localities between Hamelin Bay (just north of Cape Leeuwin) to the Abrolhos Islands. The pale form has been collected at several localities within this range and also in the western part of the Great Australian Bight. Thus, they are sympatric, but it is not yet known whether they ever occur in close proximity. Very little is known about the habitat of the pale form although it is known that it ranges from 5 fathoms (recently collected shell at Cervantes Id. ) to 100 fathoms Plate 337. Cypraea venusta Sowerby. Paratype of C. thatch- eri Cox in the British Museum (Natural History). Length: 78 mm. Courtesy of Norman Tebble. [10-477] 474 Zoila B. R. Wilson and J. A. McComb Cypraeidae ( Great Australian Bight ) . Therefore, the pale form cannot be regarded as a deep-water variant (cf. pale, deep-water friendii). Schilder (1966a) has concluded that these two forms are color morphs of the one species because they conform in all shell characters except color. We agree with Schilder’s conclusion (but not with the taxonomic procedures derived from it — see nomenclatural section). A less likely alternative is that the two forms are distinct sympatric species differing only in shell color (a notoriously bad character for separating species). Pale shells may have strong columellar teeth more often than dark shells, but this needs confirmation. Live specimens of the pale form are needed for comparison with the soft parts and radula of the better known dark form. The shells of the dark form might be mistaken for friendii thersites (see Iredale, 1939) but they are more pyriform, the extremities are more rounded and the whole base is white, pale-tan or pale-pink. In shape the shell of venusta resembles decipiens but is more swollen and less acutely humped than the latter species. The dark chocolate base and sides of the shell of decipiens at once distinguish that species from venusta. Habitat — The depth range is from 2 to about 100 fathoms. At Sorrento Beach living animals are found on yellow-sponge-encrusted limestone rocks, and on the walls and ceilings of underwater cav- erns at depths of 2 to 5 fathoms ( sometimes found in the company of the large friendii described by Wilson and Summers, 1966). Two live specimens collected by the senior author at Cape Naturaliste and Yallingup, at depths of 6 to 12 fathoms, were found on sponge- and weed-covered rocks. At Sorrento during the early summer months (November to February), females may occasionally be found on egg clutches. These consist of several hundred egg capsules embedded in a gelatinous matrix and are laid in upturned, dead limpet, fis- surellid or haliotid shells, or in suitably shaped de- pressions in sponge. It is certain that the animals feed on sponges. Frequently an animal may be found in a depression in the smooth encrusting sponge which is clearly a result of its own browsing activity. Sponge spicules form a large proportion of the gut contents. Description — Shell subpyriform; maximum height posterior to the centre; base flattened; lateral mar- gins tend to be angular, especially at the anterior end where marginal wings are produced; sides of posterior and anterior channels rounded and not elevated; spire evident but depressed and not pro- jecting (compared to friendii and marginata); aperture anteriorly dilated and usually curved pos- teriorly; anterior siphon canal usually oblique. Aperture dentition of shell — Labial teeth short and strong. In our Sorrento sample, mean number of labial teeth is 21, range 18 to 23; number of labial teeth in larger shells from other localities varies from 19 to 27. Anterior columellar teeth dis- tinct and extend into the aperture but not on to the fossula; posterior columellar teeth relatively short although distinct, but the medial teeth on the colu- mella are frequently absent, and when present they are obsolete or weak. In the Sorrento sample most shells have no visible medial columellar teeth, but some have faint medial crenulations and a small percentage have a full set of 13 or 14 teeth on that side. One specimen from Yallingup, Cape Natu- raliste (WAM 2/1966), a specimen from the Ab- rolhos Islands (J. Seabrook collection) and the types of venusta thatcheri and catei, all possess a full set of 15 to 16 columellar teeth, although the medial ones are weak. It is possible that a full set of columellar teeth may be more common in the pale form, but not enough specimens are known to confirm this. Color of shell is polymorphic; a pale and a dark morph occur. In both morphs the base is pale (pale-tan, white or pale-pink), but there may be light stripes corresponding to the position of the teeth, especially the anterior labial teeth. The sides have a characteristic greyish granular texture, which tends to become darker toward the ends and extends dorsally over the anterior and posterior channels, usually covering the lower part of the spire in adult specimens, and thus encircles the shell. The dorsal background color is white, yellow or pale-tan with varying degrees of irregular darker blotches. The fossula is white but the darker tint of juvenile shells may sometimes be seen within the aperture. The dorsum of the pale morph may be white, pale-yellow or pale-pink and may bear no blotches (e.g., the type of bakeri which is white with two pale distinct bands), or bear a few pale, orange- brown blotches or spots. The sides (i.e., overlaying the pale-grey granular texture ) have only a few in- distinct, pale orange-brown spots or none at all. The base is white, pale-yellow or pale-pink. The dorsum of the dark form is usually pale-tan with darker tan or chocolate-brown blotches which may be confluent or more or less discrete and may be sparse or so dense that they cover almost the [10-478] December 8, 1967 INDO-PACIFIC MOLLUSCA, vol. 1, Cypraea 475 no. 8 whole dorsum. The sides usually, but not always, have large brown spots overlaying the marginal granular texture which is darker than in the pale morph and is often more blue than grey. Some specimens may have a mantle line, e.g., the type of catei, see Cate, 1962, pi. 1, a specimen from Yal- lingup (WAM 2-62) and specimens from Sorrento (WAM). Measurements of shell — There is considerable variation in shell size between local populations. Shells from relatively deep water (3 to 20 fms.) in the Cape Naturaliste-Geographe Bay area and the Abrolhos Islands area are consistently larger than shells from the small shallow-water population around the reefs off Sorrento Beach. Mean length of our Sorrento sample is 58.3 mm.; range 52.0 to 65.2 mm. Mr. J. Seabrook’s specimen from the Ab- rolhos Islands is 79.7 mm. long. Specimens from Geographe Bay are usually longer than 70 mm. Several large shells have been taken by divers in 17 to 20 fathoms off Rottnest Island ( unfortunately none are yet available for measurement) and one specimen from shallow reefs off Garden Island is small (60.5 mm.), like Sorrento shells. For meas- urements of additional specimens see Cate, 1962, table 1. Soft parts — This description is for the dark form only, as we have not yet seen soft parts of pale specimens. The mantle is grey, thin, and the dorsal shell markings are just visible through it; it has a velvety texture and there are no papillae. Eye ten- tacles and the smooth edges of the anterior and posterior siphons are black. The rest of the head and the sides of the foot are fleshy-grey. Radula — This description is for the dark form only. Radula dentition of this species is distinctive. The basal denticles on the median tooth are much larger than in any other Zoila and may project be- low or level with the base of the subtending bract (plate 334, fig. c). The central cusps of the lateral teeth are very long and curved compared to other species. Radulae of 33 specimens from the Sorrento population were examined and showed surprisingly little variation. The radula of one specimen from Yallingup, Cape Naturaliste (WAM 2-1965) was examined and was indistinguishable from those of Sorrento specimens. One aberrant Sorrento speci- men was noticed with rows containing four lateral teeth on one side; the extra tooth was intermediate in shape between the 2nd and 3rd lateral teeth of normal specimens. Nomenclature — The exact origin of the very early specimens is still subject to doubt. The first positive record was made by Iredale ( 1939 ) , who described a shell from Cape Naturaliste, South- western Australia, as a western relative of Zoila thersites (Gaskoin, 1849), giving it the name Z. t. episema. Schilder (1960) separated epissma from thersites and later (1961) allied it with venusta. Subsequently, specimens have been collected at other localities and the species has been treated taxonomically in several different ways ( Cate, 1962, 1964; Schilder, 1963a, 1966a). Plate 338. Cypraea venusta Sowerby. Eastern part of the Great Australian Bight. B. Bardwell, coll., National Museum of Victoria, Melbourne. Length: 71 mm. [10-479] 476 Zoila B. R. Wilson and J. A. McComb Cypraeidae Cate (1962) argued that venusta and episema are distinct species, although he has since changed that view (personal communication; see also Cate, 1964, p. 22). The stimulus for his 1962 paper was the discovery of the West Wallabi Island, Abrolhos, shell (later nominated type of catei by Schilder) which Cate then believed to represent the typical, pale form of venusta. He argued that this shell and the types of venusta and thatcheri were conspecific but that they were a different species from dark episema as represented by his specimens from Ge- ographe Bay and Sorrento. He drew evidence for this argument from the fact that his sample from Sorrento consisted of glossy dark shells like those from Geographe Bay, but unlike his “pale” speci- men from the Abrolhos. Cate also pointed out that the type of thatcheri and his Abrolhos specimen each had a full set of columellar teeth while his samples from Geographe Bay and Sorrento did not include any specimens with this characteristic. This evidence it not substantiated by our data and ap- pears to be the result of a too small sample. Some shells from all localities may possess a full set of columellar teeth. Although Cape Naturaliste and Geographe Bay shells are characteristically dark, the Sorrento population is highly variable in tliis respect. We observed that, at Sorrento, speci- mens found in the underwater caverns are usually dark and glossy while specimens found on sponges in the open shallow water are frequently damaged, deformed and lighter in color. It is likely that Cate’s sample from this population was biased to- ward the “better quality” darker shells by the col- lectors who sent him his specimens. Cate’s speci- men from the Abrolhos Islands is not unlike many light tan shells from Sorrento (albeit other differ- ences ) but it is not a truly pale shell like the types of venusta and thatcheri. Schilder ( 1963a ) recognized the conspecificity of the Geographe Bay and Sorrento populations with the types of venusta and thatcheri. However, he recognised three geographic subspecies within the species. For the “southern” subspecies (Geographe Bay) he retained the name episema. For the Sor- rento population he introduced the new subspecies name sorrentoensis, supposing this to be a “west- ern” subspecies. The typical subspecies venusta he supposed to be the “northern” representative on the basis of the type locality which he “preferred” to restrict to the Dampier Archipelago. Schilder (1963a, p. 126) states: “The differences of geographically restricted races concern size and color only, but there is no constant difference in shape and in the structural features of the dorsum, extremities, margins, base, aperture, dentition and fossula; the posterior columellar teeth possibly are more developed in Zoila venusta venusta than in the other two races.” Because of the very restricted area and shallow conditions occupied by the Sorrento populations and the occurrence of similarly small shells in simi- lar habitats (Garden Island) and large shells in adjacent deep water (off Rottnest Island), we re- gard size as a purely local ecotypic variant having no taxonomic relevance. Color is variable and there are no discrete differences in dorsal ground color and spotting between Geographe Bay and Sorrento populations as indicated by Schilder ( 1963a, table 2). The western subspecies (“dwarf race”) sor- rentoensis, therefore is not here substantiated, par- ticularly since it occurs within the range of the larger specimens. Furthermore, in proposing venusta as a pale “northern (probably Dampier Archipelago)” race, Schilder (1963a, p. 125) committed a circular argu- ment. The only grounds he had for his “preference” for retaining Dampier Archipelago as the type lo- cality for venusta, in spite of Brazier’s (1871) evi- dence to the contrary, is that “Cervantes Island is too close to the area inhabited by Z. episema .” This very doubtful argument cannot then be used to support the separation of these taxa. It seems un- likely that this species occurs in northern Western Australia at all and the corrected type locality (Cervantes Island) is much more in conformity with the known distribution of the species. Subsequently, Schilder ( 1966a ) appears to have changed his view because he records a “typical” pale venusta from Geographe Bay, i.e., within the range of his southern subspecies venusta episema. The new arrangement of the taxa within the spe- cies implied by Schilder ( 1966a ) is totally unac- ceptable in terms of modern taxonomy. The “dwarf race” sorrentoensis is apparently still accepted as a subspecies of episema but episema itself is stated to be a “morph of subspecific rank,” presumably of venusta with which it is sympatric. Sympatric sub- species is a contradiction of terms. It is proposed here that subspecific division of venusta be discontinued and that the characters on which subdivision was based should be considered as inconsistent or variables dependent upon eco- logical conditions. The procedure adopted here of including Ber- naya catei Schilder as a synonym of venusta is a tentative one. The type of this species is the unique INDO-PACIFIC MOLLUSCA, vol. 1, no. 8 December 8, 1967 Cypraea 477 specimen from the Wallibi Group, Abrolhos Islands, described and figured by Cate (1962) as venusta. Schilder (1963a) correctly pointed out that the shell differs just as much from the types of tliatcheri and venusta as it does from Sorrento and Geog- raphe Bay shells (e.g., Cate, 1962, pi. 4 and text- fig. 2). The most significant differences are the globular shape with the maximum height central, the straight aperture with relatively well developed inward sloping teeth and the presence of a dorsal line. Because of these characters Schilder placed the specimen in the genus Bernaya rather than Zoila and gave it a new species name. The senior author had the privilege of examining this shell before it left Australia (by courtesy Mr. Max Cramer), and concluded then that it was an interesting aberrant form of venusta. Aberrant shells of similar form (although not identical) are quite common at Sorrento. Other large shells, more typi- cal of venusta, have since been collected at the Abrolhos Islands. Presence of a dorsal line is not confined to Bernaya, as supposed by Schilder. A dorsal line is common in friendii friendii, friendii thersites and also occasionally occurs in decipiens and rosselli, as well as in venusta. We do not con- sider the presence of a dorsal line in the type of catei to be of anything more than passing interest. Synonymy — 1846 Cypraea venusta Sowerby, Proc. Linn. Soc. London, vol. 1, p. 314. 1869 Cypraea tliatcheri Cox, Proc. Zool. Soc. London, [for 1869], p. 358, pi. 26, figs. 1, la. 1888 Cypraea roseopunctata Sby., Melvill, Mem. Proc. Man- chester Lit. Soc., ser. 4, vol. 1, no. 5, p. 239 (nomen nudum ) . 1916 Cypraea venusta bakeri Gatliff, Victorian Nat., Mel- bourne, vol. 32, p. 147, figs. 1-2. 1939 Zoila thersites episema Iredale, Aust. Zool., vol. 9, no. 3, pp. 300-301, pi. 27, figs. 3-4. 1963 Zoila venusta sorrentoensis Schilder, Veliger, vol. 5, no. 4, p. 126. 1963 Bernaya catei Schilder, Veliger, vol. 5, no. 4, p. 127. Types and localities — Type localities of venusta and its synonyms are as follows: venusta Sowerby: no locality given, but Cate ( 1962 ) restricted the type locality to Cervantes Is- lond [approx. 65 miles north of Perth], Western Australia; holotype in the Saul collection, Cam- bridge University, England; not figured. thatcheri Cox: the original description and the labels with the types give the locality as Dampier Archipelago, west coast of Australia, but that was subsequently corrected to “Cervantes Id.” by Brazier (1871), see Gatliff (1916), Cate (1962, 1964); holotype in Dautzenberg collection, Inst. Roy. Sci. Nat. de Belgique, Brussels, figured by Cox, 1869, pi. 26, figs. 1, la (copied by Roberts, 1885, pi. 10, figs. 44, 45, and Cate, 1962, pi. 4); paratype in British Museum (Natural History), no. 19042147, figured by Sowerby, 1869 (copied by Weinkauff, 1881); Allan, 1956, pi. 13, figs. 12-13; and our plate 337. bakeri Gatliff: “Western Australia”; holotype in National Museum of Victoria, Melbourne, no. F616, Plate 339. White form of Cypraea venusta Sowerby. Holo- type of C. bakeri Gatliff in the National Museum of Victoria, Melbourne. [10-481] 478 Zoila B. R. Wilson and J. A. McComb Cypraeidae figured by Gatliff, 1916, p. 148, figs. 1-2. See our plate 339. epissma Iredale: “Cape Naturaliste, south-west- ern Australia”; holotype in South Australian Mu- seum, Adelaide, no. D3980, figured by Cox, 1889, pi. 15, figs. 1-2, Iredale, 1939, pi. 27, figs. 3-4 (copied by Cate, 1962, pi. 3, figs, la, lb). sorrentoensis Schilder: “Sorrento Beach near Fre- mantle [Western Australia]” leg. B. B. Wilson, Jan. 7, 1962; holotype in Schilder collection, no. 13344, not figured; paratypes are the “hypotypes” numbers 3, 4 and 8 to 13 listed by Cate, 1962, p. 7 (numbers 3 and 4 figured, pi. 3, figs. 2a, 2b, 3a, 3b). catei Schilder: “West Wallaby 1.. Abrolhos Is. [Western Australia]” leg. Max Cramer; holotype in C. M. Cate collection, no. 563, figured by Cate, 1962, pi. 1 and pi. 2, figs, la, lb, and text-fig. 2a. Specimens dissected — Fifteen specimens from Sorrento Beach dissected and radulae mounted. Observations made on external anatomy and radula of one live-collected specimen from Eagle Bay, Cape Naturaliste (WAM 115-65) and one from Canal Rocks, Yallingup (WAM 2-62). Records — West Wallabi Id., Abrolhos Is. ( dark form, beach shell, see Cate, 1962); Rat Id., Abrolhos (dark form, J. Seabrook coll., Perth); Cervantes Id. (pale form, see Brazier, 1871, and Cate, 1962); Cervantes Id. (pale form, another specimen, 6 fms., craypot, N. Harold coll., Perth); Sorrento Beach, Perth metropolitan area ( small dark form, 2 to 5 fms., sponge covered rocks and walls of underwater limestone caves, leg. B. R. Wilson and R. W. George, WAM, see also Cate, 1962, 1964 and Schilder, 1963); west side Garden Id. (6 fms., craypot, R. Swan, WAM); off Binningup Beach, approx. 15 mi. north of Bunbury (pale form, 65 ft., see Cate, 1964); Busselton, Geographe Bay (pale form, see Schilder, 1966a); off Dunsborough, Geog- raphe Bay (dark form, 9 fms., limestone reef, leg. B. R. Wilson, WAM ) ; Eagle Bay, Cape Naturaliste ( dark form, 6 fms., gneissic reef, leg. B. R. Wilson, WAM ) ; Canal Rocks,, Yallingup (dark form, 12 fms., gneissic reef, leg. B. R. Wilson, WAM); west part Great Australian Bight (pal? form, trawled 80-100 fms., T. W. Marwick coll., Perth ) . Plate 340. Geographical distribution of Cypraea (Zoila) venusta in Western Australia. Cypraea decipiens E. A. Smith (PI. 335, figs. 1-6) Range — North-west Cape to Buccaneer Archi- pelago, Western Australia. The geographic range of this species is difficult to determine exactly. It is not always possible to obtain reliable locality data because of the circumstances in which most speci- mens are obtained. At one time pearl divers brought specimens into Broome, Port Hedland, Port Wal- cott ( old port for Roebourne ) and Onslow, but the pearling luggers ranged some distance from these ports. However, there appear to be no records from the Northern Territory or from south of North-west Cape. Remarks — C. (Z.) decipiens is a well-known and distinctive species from the northwestern coast of Western Australia. Apparently it is not encountered on intertidal reefs and is chiefly brought in by pearl divers from the pearling grounds. The unusually humped shell, rounded margins and extremities, and the unspotted chocolate-brown base and sides at once distinguish decipiens from other species. Habitat — Mrs. Phyllis McDaniel, part owner of pearling luggers at Broome, writes (personal com- munication) that the pearl divers take living speci- mens from branching sponges in 10 to 40 fathoms. Judging from the numbers brought in on request, the species must be abundant, although occurring in restricted areas and habitats. Description — Shell proportionately short and sub- pyriform. Dorsal hump high and steep; maximum height just behind centre. Base flattened with lateral margins only slightly angular, even at the anterior end (i.e., there are no expanded anterior marginal wings as in venusta). Sides of anterior and pos- terior channels rounded and low. Spire evident and slightly projecting with the apex dorsal to the pos- terior channel (but rarely projecting into the chan- nel). Suture of body whorl slightly impressed. Aperture rather straight and only slightly dilated at the extreme anterior end. Dentition of shell — in- side edge of labial lip thickened in such a way that the labial teeth are unusually deep (vertically), although they do not extend laterally across the base (except at the anterior end where they tend to be declivous). Weak but relatively (to other Zoila ) long teeth along the whole length of the columella; anterior columellar teeth only slightly stronger than the midde and posterior teeth and do not extend up onto the fossula or across the base. [10-482] December 8, 1967 INDO-PACIFIC MOLLUSCA, vol. 1, no. 8 Cypraea 479 Plate 341. Geographical distribution of Cypraea ( Zoila ) decipiens in Western Australia. Color of shell — Base and sides evenly colored deep chocolate-brown ( no basal or lateral spots ) ; basal coloring extends between the teeth on both sides of the aperture and covers the ventral ends of the teeth, the remainder of which are white. There is usually a patch of brown on the fossula. Dorsum has a unique pattern of numerous fine, white re- volving and longitudinal lines (slightly raised) with small rectangular facets between ( slightly de- pressed); each facet contains a more or less cres- cent-shaped blotch of brown, the overall appear- ance being a texture like that of a coarsely woven cloth (but smooth to touch). Overlaying this basic pattern there are varying amounts of darker, larger and confluent brown blotches extending upwards from the sides. Interior white, rarely with any coloration on that part of the body whorl visible within the aperture. Measurements of shell — Average length 57 mm.; range 48 to 70 mm. Average ratio of width: length 0.67; range 0.5 to 0.75. Soft parts — Mantle thin and orange with black flecks which are most dense near the margins. Smooth margins of siphons black. Eye tentacles and dorsal rim of the mouth black, head otherwise orange. Foot orange with black lateral margins and orange-yellow transverse anterior groove. Radula — Median tooth rectangular and higher than it is wide ( H : W ratio approx. 6:5). A narrow subtending bract is present but the basal denticles are small and ill-defined. Lateral tooth relatively higher than that in friendii. See plate 334, fig. d. Synonymy — 1880 Cypraea decipiens E. A. Smith, Proc. Zool. Soc. Lon- don [for 1880], p. 482, pi. 48, figs. 8, 8a. Types and localities — Smith gave only “North Australia” as the type locality for decipiens. The types are in the British Museum (Natural History), no. 18805272. Specimens dissected — Twelve animals collected by pearl divers off Broome (courtesy Mrs. Phyllis McDaniel); fixed, and preserved in methyl alcohol. Six radulae examined. Records — WESTERN AUSTRALIA: Yampi Sound, Buc- caneer Archipelago ( Geo. Robinson, leg. ) ; Sunday Id., King Sound (W. H. Matthews, leg.); Broome; Eighty Mile Bench (V. Proudfoot, leg.); Port Hedland; off Legendre Id., Dampier Archipelago (dredged 22 fms., "Hawaiian-W.A. Exp. I960”; Onslow ( Rossell Coll.). All West. Aust. Mus. Cypraea marginata Gaskoin, 1848 (PI. 335, figs. 7-9; pi. 342) Range — Dorre Island, Western Australia, to Cape Jervis, South Australia. Remarks — Until a few years ago marginata was known only from a unique and unlocated specimen Plate 342. Cypraea marginata Gaskoin. Holotype from the British Museum (Natural History). Length: 57 mm. [10-483] 480 Zoila B. R. Wilson and J. A. McComb Cypraeidae in the British Museum (Natural History), thought by some ( e.g., Allan, 1956 ) to be a juvenile of ther- sites Gaskoin, 1849. In recent years a number of specimens, including some alive, have been col- lected and the range of the species established as Western and South Australia (Laws, 1966). Cypraea marginata seems to be the closest living relative of friendii. The radulae of the two species differ only in very minor ways and from the dorsal view some dark specimens of marginata might be easily mistaken for friendii. However, the pale base with brown impressed spots toward the margins is characteristic of marginata. When live animals are compared, the two species may be easily distin- guished by differences in animal color. The mantle, eye-tentacles, head and foot are black in friendii, but only the eye-tentacles are black in marginata, the rest of the body being white or without pigments. Habitat — The depth range is known to be at least from 6 fathoms to 80 fathoms. The majority of the live specimens collected have come from pots (“craypots”) set to catch the commercial crayfish or spiny lobster of Western Australia on rocky (limestone) bottom at depths of 15 to 40 fathoms. A number of live specimens have been collected by skindivers recently. Mr. Kevin Morgan of Perth col- lected one (J. E. Norton collection) on a sponge- covered rock at about 6 fathoms off Carnac Islands, Western Australia. Dr. H. Laws (1966) records three specimens recently collected by skindivers in South Australia at depths of 7 to 13 fathoms. The species is also reported from South Australia, by Stokes, 1966. Description — Shell ovate. Dorsal hump steep pos- teriorly; maximum height just behind centre. Base broad and flattened with strongly marginated sides which are characteristically irregularly indented, especially near the posterior end; marginal indenta- tions correspond with irregular impressions on the base. Sides of anterior and posterior channels sharp and elevated, although not as high as in friendii. Spire elevated and usually projecting on left side of the posterior channel, sometimes even beyond the extremities. Aperture narrow and only slightly curved. Relative width and height are clinal variants in this species (cf. friendii, Wilson and Summers, 1966). The few specimens from the south coast of Australia are significantly wider and higher relative to shell length than specimens from the mid-west coast of Western Australia (see illustrations by Cate, 196.1; Stokes, 1966, and Laws, 1966). Aperture dentition of shell — Teeth present along whole length of both sides of the aperture. Labial teeth well developed but rather small and sharp and extend about a quarter of the way across the outer lip. Columellar teeth small, short and do not extend up on to the fossula. Number of labial teeth 24 to 29; columellar teeth 16 to 23. Color of shell variable. Dorsum may be entirely white, or, over a white ground color, bear scattered spots and flecks of brown or occasionally large semi-confluent blotches of brown as in friendii. Teeth, fossula and sides of the aperture white; the rest of the base spotted and colored with brown in varying degrees, the larger brown spots usually occupying irregular depressions on the base of the shell. Basal spots extend to the lateral margins. Measurements — Average length 60 mm.; range 46.0 mm. to 62.4 mm. Range of the ratio width : length 0.62 to 0.65. For additional data see Schilder, 1966c. Soft parts — Mantle and siphons very thin, smooth, colorless and transparent. The shell markings and [10-484] December 8, 1967 INDO-PACIFIC MOLLUSCA, vol. 1, no. 8 Cypraea 481 teeth are easily visible beneath the extended man- tle. The foot and head are translucent- white; eye- tentacles black. ( Data from color photograph cour- tesy Max Cramer of Geraldton. ) Radula — The radula teeth are similar to those of friendii but there is a curious indentation in the top of the median tooth, and the central cusp of the median tooth is more projecting and pointed. The subtending bract also tends to be pointed rather than rounded as in friendii. See plate 334, fig. a. Synonymy — 1849 Cypraea marginata Gaskoin, Proc. Zool. Soc. London [for 1848], p. 91. 1964 Zoila marginata (Gaskoin, 1849), C. N. Cate, The Veliger, vol. 7, no. 1, p. 23; 1967, Schilder, Ha- waiian Shell News, new series, no. 88, April, p. 5 (radula illus. ); 1967, ibid., no. 92, Aug., p. 4. Types and localities — Gaskoin gave no data with the original type specimen which is now located in the collection of the British Museum (Nat. Hist.), no. 42777. Cate (1961, pp. 76-78) purchased the second known specimen from the late Lloyd Berry collection and restricted the type locality on the basis of data on the label with the specimen to “Albany Bay, Western Australia (34° 57' South Lat., 117° 58' East Long.)” giving the depth as approxi- mately 80 fathoms. There is no “Albany Bay” (Al- bany is a small city at the head of King George Sound) and Cate apparently intended the locality to be King George Sound, although the depth “within the perimeter” of the Sound nowhere ex- ceeds 40 fathoms. As Cate points out in a subsequent paper ( 1964, p. 23 ) the locality itself is questionable. Mr. Angelo was not a fisherman, as concluded by Cate, but an amateur shell collector who bought this and speci- mens of other deep water cowries from trawlers operating from Albany (personal communication from the late Senator T. W. Marwick who accom- panied Mr. Angelo). The trawlers were working at depths of 80 to 100 fathoms in the western part of the Great Australian Bight between Eucla and Esperance some 400 miles to the east of Albany, and there is every reason to believe the specimen was trawled in that area. Cate ( 1964, p. 23 ) “corrected” the type locality of marginata to “Pelsart Is., Houtman Abrolhos Group” (which lie off the mid-west coast of West- ern Australia) on the basis of another specimen in his collection from that locality. Although doubt remains about the exact locality from which the type specimen came, the identity and range of the species is now well established and there is little point in arbitrarily restricting the type locality. Specimens dissected (cursory examination only) — Bodies of one male and one female from off Jurien Bay (courtesy T. Sutcliffe) and one male from off Geraldton (courtesy Max Cramer). Radulae were examined from all three specimens. See plate 334, fig. a. Records — WESTERN AUSTRALIA: W. of Dorre Id. ( trawled 60 fins., Poole Bros, on “Bluefin” ) ; 12 mi. off Geraldton (craypot, 20 fms., T. Merindino, leg.); S. of Southern Group, Abrolhos Ids. (craypot, 35 fms., K. Gus- tafsson, leg.); off Jurien Bay (craypot, 25-30 fms., K. Gus- tafsson ) (all West. Aust. Mus. j; Garden Id., near Fremantle, 30 ft. depth (Schilder, 1967, Hawaiian Shell News, new series, no. 88, p. 5); Lancelin Id. (see Cate, 1964); off Carnac Id. (diver, 6 fms., K. Morgan, on sponge-covered wall of limestone reef, J. E. Norton coll.); Robert Point, Mandurah (see Cate, 1964); “Albany Bay” [Kang George. Sound], Lat. 34°57' S., Long. 117°58' E. [almost certainly- erroneous— probably trawled on fishing grounds south of Eyre] (trawled 80 fms., see Cate, 1961, Veliger, vol. 3, no. 3, pp. 76-78, pi. 14). SOUTH AUSTRALIA: near Taylor’s Id., 20 mi. from Port Lincoln, 40 ft. (see Stokes, 1966, and Laws, 1966); Cape Jervis, 70-80 ft. (see Laws, 1966). Cypraea rosselli (Cotton, 1948) (PI. 335, figs. 10-12) Range — Central coast of Western Australia from Binningup Beach north of Bunbury to Dirk Hartogs Island. Remarks — Superficially, rosselli resembles margi- nata, especially in size, dentition and the margi- nated sides, although, as Schilder (1960) suggests, this is probably due to parallel evolution, not neces- sarily indicating close genetic relationship. The papillae on the mantle, the lack of a subtending bract and basal denticles on the medial radula tooth, the relatively small size of the inner basal denticle of the first lateral tooth, the relatively well developed aperture dentition and low spire indi- cate that rosselli is probably only distantly related, if at all, to other species of Zoila. This relationship will need to be re-evaluated when it becomes pos- sible to examine well-preserved female specimens. [10-485] 482 Zoila B. R. Wilson and J. A. McComb Cypraeidae This species was discovered relatively recently and is still a rare shell in collections. The first speci- mens were collected on the beach at Leighton near North Wharf, Fremantle, by the late Mr. Harold Rossell of Subiaco ( a suburb of Perth ) in the 1920’s, but it was not until 1947 that his specimens came to the attention of Mr. B. C. Cotton of the South Australian Museum, who described them in 1948. Rossell sent four shells to Mr. W. R. Steadman who presented one of them ( the holotype ) to the South Australian Museum. Rossell himself presented three other specimens to the Western Australian Museum in May 1947 (apparently unknown to Cotton) and retained several in his own collection, which was disposed of by his widow some years after his death. Unfortunately, Rossell dipped all his speci- mens in hydrochloric acid, so removing all trace of color from their dorsal surfaces and giving them an unnatural white finish. Rossell (see Cotton, 1948, pp. 30-31) suggested that the shells may all have come from dredgings taken in Fremantle Harbour and dumped just off the beach between North Wharf and Leighton. However, at least one additional dead specimen has since been taken at Cottesloe Reach a few miles farther north (1955, WAM no. N874) and frag- ments have been dredged in 20 fathoms north of Rottnest Island. It seems at least as likely that the type series came from a living population some- where off the mainland or north of Rottnest Island. Mr. E. Nicholls of Mandurah picked up a dead shell on Rinningup Beach, about 15 miles north of Bunbury ( Hawaiian Shell News, vol. 10, no. 2, p. 7, December 1961 ) . The first live specimen was a juvenile collected by Mr. Max Shaw at Perth while SCUBA diving in 35 fathoms west of Rottnest Is- land. Shaw and Mr. Kevin Morgan had dived to this great depth and found a beautiful reef with many sponges and other growths. Shaw found the young rosselli, which did not have its mantle ex- tended, nestling in a crevice of a large fan-shaped sponge. The shell, sponges and other interesting objects were stuffed into a sack. On reaching the .surface Shaw complained of severe cramps and was rushed back to Fremantle and to hospital where his difficulty was diagnosed as divers’ paralysis or the “bends.” He was placed in a recompression cham- ber and later recovered. The following day the con- tents of the sack were examined and out rolled the young live rosselli. It was several days before the specimen was brought to the senior author and by that time the animal was in an advanced state of decomposition, but some anatomical information was obtained from it. As a result of publicity given to this story in the local press, the Western Australian Museum was inundated with cowry shells to be identified and within a few weeks two more fully adult and live- collected shells came to light. Both these specimens were taken from “craypots” set in 20 to 40 fathoms off the Western Australian coast. Since then several other specimens have been collected alive, one by Mr. Max Cramer under the wharf in Geraldton Harbour. Undoubtedly this beautiful and most unusual species is rare only in collections because of its habitat in relatively deep water. Habitat — Continental shelf from 3 fathoms to at least 40 fathoms. The specimen collected by Mr. M. Shaw off Rottnest Id. was found on a large fan- shaped sponge ( not on fan-coral as stated by Cate, 1964, p. 23). Plate 344. Geographical distribution of Cypraea (Zoila) rosselli in Western Australia. [10-486] December 8, 1967 B. R. Wilson and J. A. McComb Cypraea 483 Description — Shell ovate-pyriform. Dorsal hump high and rather steep posteriorly, maximum height just behind centre. Base broad and flattened, with sharply angulated margins. At the anterior end the outer lip slopes inward steeply, thereby increasing the angularity of the right extreme anterior margin, so forming a sharp edge; the left anterior and the right posterior margin are flanged and almost as sharp, but the sharpness of the left posterior mar- gin is variable depending on the degree of indenta- tion of the shell beneath the sides of the posterior channel. Anterior channels often almost vertical. Margins on both sides are curiously elevated in the centre, especially on the left side, and often irregu- larly indented, especially posteriorly on the right side. Sides of the anterior and posterior channels sharp and only slightly elevated. Spire low (com- pared to other species of the subgenus Zoila) and not usually visible in adult shells, being covered by depositions of shell. Aperture strongly curved, but only slightly dilated anteriorly. Aperture dentition of shell — Teeth are present along both sides of the aperture. Labial teeth num- ber 25 to 31; columellar teeth 18 to 26. Labial teeth short but quite strong; columellar teeth shorter and weaker; anterior columellar teeth only slightly longer than those on the medial region and do not extend up on to the deep fossula. Columellar termi- nal ridge obsolete. Color — Dorsal ground color cream, sometimes with a faint pale-tan band anteriorly; ground color usually obliterated, except for a cream-colored patch on the dorsal hump, by dark chocolate-brown ex- tending up from the sides. Base similarly brown, and there are no basal, lateral or dorsal spots. In one specimen (WAM no. 114-65) all that remains visible of the dorsal ground color is a pale longi- tudinal mantle line which divides to form a Y at both ends; the remainder of the shell is evenly colored dark chocolate-brown. Teeth brown; inter- stices white. Aperture and fossula white but color sometimes visible on the body whorl posteriorly. Measurements (mm.) — Shell length varies from 42 to 57 mm. ( maximum size teste Mr. Max Cramer, Geraldton ) . Range of the ratio width : length is 0.61 to 0.66. Soft parts — The bodies of only two specimens have been examined, both in a condition from which little conclusive information could be obtained. Mantle relatively thick and black, bearing a few widely spaced papillae (too contracted to deter- mine whether they were simple or branched ) ; sides of foot black, head and eye tentacles pale-orange in the juvenile specimen and white in the adult ( but because the black pigment had partly sloughed off the mantle and foot of both specimens due to the poor preservation, it is possible that lack of pig- mentation of the head and eye tentacles may be an artificial consequence of the state of preservation). Radula — The radula was successfully extracted from only one specimen (juvenile Rottnest Id. speci- men). The following is given therefore as a tenta- tive description of the radula of this species. The lateral and marginal teeth resemble those of other species of Zoila, but are rather wider and the medial cusps shorter; the inner basal denticle of the lateral tooth is very much smaller than any other species. The medial tooth is rather high, bears no subtending bract and no basal denticles are evi- dent. The internal dumb-bell-shaped bract of the medial tooth is high and prominent. Synonymy — 1948 Zoila rosselli Cotton, Trans. Roy. Soc. So. Australia, vol. 72(1), p. 30, pi. 1, figs. 1-6. Specimens dissected (cursory examination only) — One juvenile animal, collected by Max Shaw off Rottnest; soft-parts decomposed; radula mounted. One adult animal, collected by Max Cramer at Geraldton (shell not seen by us); soft-parts dam- aged during extraction from the shell. Records — WESTERN AUSTRALIA: W. of Dirk Hartogs Id. (craypot, 30-40 fms., V. Lombardo, WAM); Geralton harbour (diving 2 fms.. Max Cramer, see Cate, 1964); 5 mi. N.E. of Rottnest Id. (dredged 20 fms., Mariel King, “Ha- waiian-W.A. Exp.,” 1960, broken pieces, WAM); W. of Rottnest Id. (on sponge, 35 fms.. Max Shaw, WAM); be- tween North Wharf, Fremantle and Leighton Beach (beach specimens, H. Rossell, So. Aust. Mus., WAM and private collections); Mudarup Rocks, Cottesloe (beach collected, D. Wignall, WAM); Binningup Beach, 15 mi. N. of Bun- bury (beach collected, E. Nickel, see Hawaiian Shell News, 1961, vol. 10, no. 2, p. 7). [10-487] 484 Zoila B. R. Wilson and J. A. McComb Cypraeidae INDEX TO ZOILA NAMES IN VOL. 1, NO. 8 The number following the name refers to the pagination found at the top of the page. The column at the right con- tains the looseleaf pagination. anatomy, 458 [looseleaf] 10-458 bakeri Gatliff, 477 10-481 Bernaya, 466 10-466 bibliography, 468 10-468 catei Schilder, 477 10-481 consobrina McCoy, 465 10-465 contraria Iredale, 471 10-475 decipiens E. A. Smith, 478 10-482 dorsata Tate, 465 10-465 episema Iredale, 477 10-481 friendii Gray, 470 10-474 gabrieli Chapman, 465 10-465 gendinganensis Martin, 465 10-465 Gigantocypraea, 467 10-467 gigas McCoy, 465 10-465 marginata Gaskoin, 479 [looseleaf] 10-483 mulderi Tate, 465 10-465 platypyga McCoy, 465 10-465 radula, 463 10-463 roseopunctata Sowerby, 477 10-481 rosselli Cotton, 481 10-485 schilderi Dey, 465 10-465 scottii Broderip, 471 10-475 simplicior Schilder, 465 10-465 Siphocypraea, 466 10-466 species, list of, 465 10-465 sorrentoensis Schilder, 477 10-481 thatcheri Cox, 477 10-481 thersites Gaskoin, 472 10-476 toxorhyncha Tate, 465 10-465 venusta Sowerby, 473 10-477 vercoi Schilder, 471 10-475 Zoila, 469 10-473 Published by The Department of Mollusks Academy of Natural Sciences of Philadelphia 19th and the Parkway Philadelphia, Pennsylvania 19103 [10-488] December 8, 1967 INDO-PACIFIC MOLLUSCA, vol. 1, no. 8 Index 485 INDEX TO VOLUME 1, NUMBERS 1-8 Several indices have already been issued (Strombus, Turrinae, Pinnidae and Tridacnidae). They are not repeated here, except for the listings of all generic and subgeneric names. All new names proposed in volume 1 are listed in this index and are in boldface type. A list of issues and con- tributing authors appears in vol. 1, no. 8, p. 489 (p. 00-007). In this index, the number following the name refers to the pagination of vol. 1 found at the top of the page. The col- umn at the right contains the looseleaf pagination. Abbott, R. T., 15 [looseleaf] 20-403 33 09-831 147 10-051 401 09-945 445 09-801 455 09-919 Afer, 31 20-471 Alio, 425 23-145 Altivasum, 25 20-431 anesthetic for mollusks, 394 62-082 458 10-458 Antiplanes, 339 23-021 Artopoia, 449 09-809 Atrina, 203 53-569 Austroturris, 412 23-108 Avicularium, 356 62-016 Aviculipinna, 178 53-504 Bibliographies for Cypraea (Zoila), 468 10-468 Pinnidae, 185 53-511 Strombus, 46 09-850 Terebellum, 488 09-804 Tridacnidae, 357 62-021 Turrinae, 238 22-672 Byssocardium, 356 62-016 Campylacrum, 282 22-824 Canarium, 63 09-891 455 09-919 Carinoturris, 281 22-823 Chametrachea, 379 62-067 Chimaera, 187 53-533 Chimaeroderma, 187 53-533 Cinguliturris Powell, 319 22-949 Commensal Crustacea, 226 53-632 396 62-004 Conomurex, 135 10-005 Coronia, 281 22-823 Crustacea (in Pinnidae), 226 [looseleaf] 53-632 (in Tridacnidae), 396 62-004 Cryptogemma, 279 22-791 Curvulites, 178 53-504 Cynodonta, 15 20-403 Cypraea (subgenus Zoila), 457 10-457 Cyrtopinna, 187 53-533 Digitata, 151 10-057 Dilatilabrum, 62 09-880 Dinodacna, 367 62-043 Dolomena, 89 09-927 401 09-945 Doxander, 111 09-963 Echinoturris, 403 22-947a Emerson, W. K., 397 09-881 Eopleurotoma, 282 22-824 Eoturris, 283 22-825 Epalxis, 284 22-826 Epidirella, 297 22-875 Epidirona, 299 22-879 List of species, 300 22-880 Euprotomus, 125 09-991 399 10-001 Exitopinna, 188 53-534 Feliciella, 425 23-145 Fenestrosyrinx, 425 23-145 Fusiturris, 241 22-685 Gataron, 367 62-043 Gemmula, 243 22-695 dampierana Powell, 248 22-700 diomedia Powell, 253 22-715 hombroni Hedley, 262 22-734 439 22-734a interpolata Powell, 435 22-703 microscelida Dali, 440 22-734b miocoronifera Powell, 254 22-716 pseudomonilifera Powell, 250 22-704 sibukoensis Powell, 258 22-730 tessellata Powell, 439 22-734a unilineata Powell, 437 22-716a Gibberulus, 141 10-015 Goniocardium, 356 62-016 Harpago, 169 10-083 Hemipleurotoma, 281 22-823 [91-001] 486 Index Editors Index [looseleaf] [looseleaf] Heptadactylus, 151 10-057 Lambis (continued) , 147 10-051 Hesperiturris, 282 22-824 radix -bryoniae Morch, 155 10-061 Heteroturris, Powell, 411 23-107 robusta Swainson, 166 10-076 sola Powell, 411 23-107 rugosum Sowerby, 172 10-086 Hippopus, 361 62-031 scorpio Murray, 164 10-074 scorpius Linne, 164 10-074 Infracoronia, 281 22-823 sebae Kiener, 156 10-062 sinuatus Perry, 166 10-076 Jung, P„ 445 09-801 sowerbyi Morch, 156 10-062 truncata Humphrey, 154 10-060 Kuroshioturris, 293 22-865 undulata Roding, 171 10-085 violacea Swainson, 167 10-077 Labiostrombus, 107 09-955 Lentigo, 117 09-973 Laevipinna, 178 53-504 Lophiotoma, 303 22-913 Laevistrombus, 47 09-855 Lophioturris Powell, 311 22-931 Lambis, 147 10-051 Lucerapex, 285 22-837 aculeatus Perry, 158 10-064 adenica Powell, 286 22-838 arthritica Roding, 173 10-087 Lucis, 449 09-809 aurantia Lamarck, 158 10-064 auranticum Sowerby, 158 10-064 McComb, Jennifer A., 457 10-457 bengalina Grateloup, 155 10-061 Micropleurotoma, 431 23-175 bryonia Gmelin, 155 10-061 Millepes, 161 10-071 camelus Gray, 153 10-059 Monodactylus, 125 09-991 cerea Roding, 153 10-059 chiragra Linne, 170 10-OS4 Okutani, T, 399 10-001 crocata Link, 157 10-063 Oostrombus, 141 10-015 crocea Reeve, 163 10-073 Optoturris, 295 22-871 davilae, 155 10-061 Oxyacrum, 283 22-825 digitata Perry, 163 10-073 Oxysma, 178 53-504 divergens Perry, 173 10-087 Palaeopinna, 177 53-503 elongata Swainson, 163 10-073 Persikima, 375 62-057 harpago Roding, 171 10-085 Pinguigemmula, 277 22-789 hermaphrodita Roding, 153 10-059 luzonica Powell, 278 22-790 indomaris Abbott, 165 10-075 okinavensis MacNeil, 277 22-789 kochi Freyer, 172 10-086 philippinensis Powell, 278 22-790 laciniata Roding, 153 10-059 thielei Finlay, 279 22-791 lambis Linne, 151 10-057 Pinna, 187 53-533 lamboides Roding, 153 10-059 Pinnarius, 187 53-533 lobata Roding, 153 10-059 Pinnidae, 175 53-501 maculata Roding, 153 10-059 Index to names, 225 53-631 millepeda Linne, 161 10-071 Key to Recent species, 180 53-506 multipes Deshayes, 168 10-078 Pinnigena, 178 53-504 nigricans Perry, 173 10-087 Pinnula, 187 53-533 nodosa Lamarck, 165 10-075 Pleuroliria, 315 22-935 novem-dactylis Deshayes, 163 10-073 Pleurotoma, 327 22-977 pilsbryi Abbott, 158 10-064 Polystira, 315 22-935 pseudo-scorpio Lamarck, 167 10-077 Powell, A. W. B., 227 22-661 purpurea Swainson, 168 10-078 403 22-947 radix Roding, 155 10-061 409 23-101 [91 - 002] December 8, 1967 INDO-PACIFIC MOLLUSCA, vol. 1, no. 8 Index 487 [looseleaf] [looseleaf] Pterocera, 151 10-057 Terebellum (continued) , 445 09-801 See species under Lambis spirale Perry, 452 09-812 Pteroceras, 151 10-057 subulatum Lamarck, 452 09-812 Pteroceres, 151 10-057 terebellum Linne, 449 09-809 Pterocerus, 151 10-057 terebra Bose, 452 09-812 Ptychosyrinx, 289 22-851 variegatum Link, 452 09-812 teschi Powell, 291 22-853 Terebrina, 449 09-809 Pyrella, 32 20-472 Trichites, 178 53-504 Pyrenella, 32 20-472 Tricornis, 53 09-871 Pyropsis, 31 20-471 397 09-881 Pyrula bengalina Grat., 155 10-061 Tridachna, 367 62-043 Tridacna, 367 62-043 Quantulopinna, 187 53-533 Tridacne, 367 62-043 Tridacnidae, 347 62-007 Rectiplanes, 339 23-021 Index to names, 395 62-003 Rectisulcus, 339 23-021 Key to species, 359 62-023 Rosewater, J„ 175 53-501 Tridacnodites, 367 62-043 347 62-003 Trypanotoma, 281 22-823 Tudicla, 31 20-471 Sawkinsia, 356 62-016 angulata Angas, 32 20-472 Scolymus, 15 20-403 capitatus Perry, 32 20-472 Servatrina, 211 53-597 carinata Roding, 32 20-472 Sinistrella, 281 22-823 costata Angas, 32 20-472 Spirillus, 32 20-472 houreqi Collignon, 32 20-472 Stegoconcha, 178 53-504 krenkeli Cox, 31 20-471 Streptopinna, 221 53-627 rames Cuvillier, 32 20-472 Strombus, 33 09-831 rostratus Schliiter, 32 20-472 Index to names, 145 09-651 rusticulus Basterot, 32 20-472 daviesi Dey, 401 09-945 spirillus Linne, 32 20-472 iredalei Abbott, 133 09-999 thebaica Cuvillier, 32 20-472 klineorum Abbott, 70 09-898 turbinata Angas, 32 20-472 listeri T. Gray, 399 10-001 Tudiclana, 31 20-471 ochroglottis Abbott, 74 09-902 Tudicula, 27 20-443 oldi Emerson, 397 09-881 armigera A. Adams, 27 20-443 orrae Abbott, 66 09-894 inermis Angas, 30 20-446 quilonensis Dey, 402 09-946 rasilistoma Abbott, 29 20-445 rugosa Sowerby, 456 09-920 spinosa H. & A. Adams, 29 20-445 wilsoni Abbott, 455 09-919 zanzibarica Abbott, 31 20-471 Subitopinna, 188 53-534 Turridrupa, 413 23-117 Sulcatipinna, 177 53-503 Key to species, 414 23-118 consobrina Powell, 418 23-126 Taranis, 425 23-145 diffusa Powell, 422 23-132 Key to species, 426 23-146 rnaoria Powell, 424 23-134 percarinata Powell, 427 ticaonica Powell, 427 23-151 23-151 prestoni Powell, 423 23-133 Terebellum, 445 09-801 weaveri Powell, 423 23-133 album Link, 452 09-812 Turrinae, 227 22-661 lineata Roding, 452 09-812 409 23-101 lineatum Perry, 452 09-812 Index to names, 341 22-651 nebulosum Roding, 452 09-812 Tunis, 327 22-977 punctatum Chemnitz, 452 09-812 intricata Powell, 332 22-982 punctulorum Roding, 452 09-812 Unedogemmula, 269 22-761 [91 - 003] 488 Index Editors Index [looseleaf] Vasidae, 15 20-403 Vasum, 15 20-403 Key to species, 16 20-404 armatum Broderip, 18 20-406 aurantiacus Verco, 25 20-431 ceramicum Linne, 19 20-407 cornigera Lamarck, 17 20-405 crosseanum Souverbie, 23 20-411 flindersi Verco, 25 20-431 imperialis Reeve, 20 20-408 nigra Perry, 17 20-405 rhinoceros Gmelin, 21 20-409 spinosa G. Fischer, 19 20-407 triangularis E. A. Smith, 23 20-411 truncatum Sowerby, 23 20-411 tubiferum Anton, 20 20-408 Vasum ( continued ), 15 [looseleaf] 20-403 turbinellus Linne, 17 20-405 variolaris Lamarck, 17 20-405 Veruturris, 404 22-948a Viridoturris Powell, 320 22-950 Volutella, 15 20-403 Wilson, Barry R., 457 10-457 Xenuroturris, 321 22-961 Key to subgenera, 403 22-947a castanella Powell, 442 22-984a cerithiformis Powell, 442 22-964a gemmuloides Powell, 443 22-967 kingae Powell, 325 22-965 Zoila, 457 10-457 Index to trivial names, 484 10-488 Key to species, 469 10-473 Published by The Department of Mollusks Academy of Natural Sciences of Philadelphia 19th and the Parkway Philadelphia, Pennsylvania 19103 [91 - 004] December 8, 1967 INDO-PACIFIC MOLLUSCA, vol. 1, no. 8 List of issues 489 ISSUES AND CHANGES Looseleaf subscribers should put this sheet in the front of their binder just before the Introduction [looseleaf p. 00-051]. Note that some numbers con- tain additions and changes to previously published monographs. Remove staples from all issues and place the new sheets where they belong by follow- ing the looseleaf numbering at the bottom of the page. Throw away any blank, brown paper sheets. Replaced pages may be discarded or saved in the back of the binder behind the guide tab, “Replaced pages.” Notice that the index pages are numbered so that they will immediately follow their appro- priate tab. Volume 1 has now been completed. An index has been published. Subscribers may now have their copies bound or may keep them in our heavy-duty post binder. List of Guide Tabs Guide tabs are automatically sent to all those who receive the journal looseleaf. Those receiving the folded pamphlet style for future binding may, if they wish, order tabs at the rate of 10 cents ( U.S. ) each. The present set of ten costs $1.00. List of Issues 00-005 Guide to Contents 00-100 Phasianellidae 04-000 Strombidae 09-650 Cassidae 12-400 Vasidae 20-400 Turridae 22-500 Pinnidae 53-500 Tridacnidae 62-000 Replaced Pages 95-000 Date of First page of Volume 1 Subject Author Publication no. pp. looseleaf no. 1, pp. 1- 14 Introduction Editors Mar. 31, 1959 14 00-003 15- 32 Vasidae R. T. Abbott 18 20-403 2 33-146 Strombus R. T. Abbott Nov. 23, 1960 114 09-831 3, 147-174 Lambis R. T. Abbott Sept. 28, 1961 38 10-051 4, 175-226 Pinnidae J. Rosewater 52 53-501 5, 227-346 Turrinae A. W. B. Powell Mar. 31, 1964 120 22-661 6, 347-396 Tridacnidae J. Rosewater Apr. 30, 1965 50 62-003 397-398 Strombus oldi W. K. Emerson 2 09-881 399-400 Strombus listeri T. Okutani 2 10-001 401-402 Miocene Indian Strombus R. T. Abbott 2 09-945 403-406 T urrinae ( replacement ) A. W. B. Powell 4 22-947 407-408 Issues and Changes Editors “ 2 00-005 7, 409-444 Turrinae (concluded) A. W. B. Powell May 15, 1967 36 23-101 445-454 Terebellum Jung and Abbott “ 10 09-801 455-456 Strombus wilsoni R. T. Abbott 2 09-919 8, 457-484 Cypraea (Zoila) Wilson and McComb Dec. 8, 1967 28 10-457 485-488 Index to vol. 1 Editors 4 91-001 489-490 Issues and Changes Editors 2 00-007 [00 - 007] 490 Editors List of issues Although primarily a scientific journal, INDO-PACIFIC MOLLUSCA is attrac- tively designed and richly illustrated in full color to serve the serious amateur interested in marine shells. Subscription The subscription rate is 7^ U.S. per page. Numbers are issued at irregular intervals, as research is completed. It is hoped that a yearly average of a hun- dred pages will be maintained. A list of back numbers, their prices and an order form may be obtained from the pub- lisher. The journal is not available on a library exchange basis, and discounts to magazine agencies are not made. The blue and gold-embossed, expandi- ble post binder shown here will hold 600 pages. 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