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Berkeley, California R. Stohler, Founding Editor Volume 33 January 2, 1990 to October 1, 1990 TABLE OF CONTENTS Number 1 (January 2, 1990) The functional morphology of the pedal musculature of the ma- rine gastropods Busycon contrarium and Haliotis kamtschat- kana. JANICE VOLTZOW The Recent eastern Pacific species of the bivalve family Thra- ciidae. EUGENE’ Vo COAN (22s ghenane eae aus anter 20 Laubierinidae and Pisanianurinae (Ranellidae), two new deep- sea taxa of the Tonnoidea (Gastropoda: Prosobranchia). ANDERS WAREN AND PHILIPPE BOUCHET ........... 56 Reproductive systems of neritimorph archaeogastropods from the eastern Pacific, with special reference to Nerita funiculata Menke, 1851. Roy S. Houston Indirect evidence of a morphological response in the radula of Placida dendritica (Alder & Hancock, 1843) (Opisthobran- chia: Ascoglossa/Sacoglossa) to different algal prey. J. SHERMAN BLEAKNEY 111 Trail following in Littorina irrorata: the influence of visual stimuli and the possible role of tracking in orientation. RICHARD A. TANKERSLEY Number 2 (April 2, 1990) Allozyme variation in the Australian camaenid land snail Cris- tilabrum primum: a prolegomenon for a molecular phylogeny of an extraordinary radiation in an isolated habitat. DaviID S. WOODRUFF AND ALAN SOLEM 129 Comparative zoogeography of marine mollusks from northern Australia, New Guinea, and Indonesia. FRED E. WELLS 140 New records for western Pacific Morum (Gastropoda: Harpidae) with biogeographic implications. WILLIAM K. EMERSON 145 Seasonal, diurnal, and nocturnal activity patterns of three species of Caribbean intertidal keyhole limpets (Mollusca: Gastro- poda: Fissurella). CRAIG J. FRANZ 155 Anatomy of the circulatory system of the nudibranch Platydoris argo (Linné, 1767) with comparisons among Doridacea (Gastropoda: Opisthobranchia). F. J. Garcia AND J. C. GARCIiA-GOMEZ The malacological contributions of Ida Shepard Oldroyd and Tom Shaw Oldroyd. EUGENE V. COAN AND MICHAEL G. KELLOGG 174 Calcium source for protoconch formation in the Florida apple snail, Pomacea paludosa (Prosobranchia: Pilidae): more evi- dence for physiologic plasticity in the evolution of terrestrial eggs. RICHARD L. TURNER AND CATHLEEN M. McCaseE ... 185 Ca-binding glycoproteins in molluscan shells with different types of ultrastructure. TETSURO SAMATA 190 A new species of the genus Cyerce Bergh, 1871, from the Cape Verde Islands (Opisthobranchia: Ascoglossa). JESUS ORTEA AND JOSE TEMPLADO Number 3 (July 2, 1990) Photic vesicles. RICHARD M. EAKIN 209 Continuous reproduction and episodic recruitment of Lacuna vincta (Montagu, 1803) in the Gulf of Maine. EDWARD J. MANEY JR. AND JOHN P. EBERSOLE 215 Recruitment of the estuarine soft-bottom bivalve Polymesoda car- oliniana and its influence on the vertical distribution of adults. DAN C. MARELLI 222 Additional opisthobranch mollusks from Oregon, with a review of deep-water records and observations on the fauna of the south coast. JEFFREY H. R. GODDARD 230 Temporary northern range extension of the squid Loligo opales- cens in Southeast Alaska. BRucE L. WING AND ROGER W. MERCER 238 A review of the Recent eastern Pacific Acanthochitoninae (Mol- lusca: Polyplacophora: Cryptoplacidae) with the description of a new genus, Americhiton. G. THOMAS WATTERS 241 New species of Late Cretaceous Cypraeacea (Mollusca: Gastro- poda) from California and Mississippi, and a review of Cretaceous cypraeaceans of North America. LINDSEY T. GROVES il New Paleogene siliquariid and vermetid gastropods from the Pacific coast of southwestern North America. RICHARD L. SQUIRES 286 Tibiaporrhais, a new Late Cretaceous genus of Aporrhaidae re- sembling Tibia Roding. WILLIAM P. ELDER 293 Relocation of Ervilia Turton, 1822 (Bivalvia) from the Meso- desmatidae (Mesodesmatoidea) to the Semelidae (Tellino- idea). BRIAN MoRTON AND PAUL H. Scott 299 First occurrence of the Tethyan bivalve Nayadina (Exputens) in Mexico, and a review of all species of this North American subgenus. RICHARD L. SQUIRES 305 Nanomelon vossi, a new deep-water Zidoninae from off southern Brazil (Gastropoda: Volutidae). José H. LEAL AND ELIEZER DE C. RIos New records of two rare aeolid nudibranchs from the coast of California. TERRENCE M. GOSLINER Number 4 (October 1, 1990) Reproductive, morphological, and genetic evidence for two cryptic species of northeastern Pacific Nucella. A. RICHARD PALMER, ScOoTT D. GAYRON, AND DavIpD S. WOODRUFF . 325 Survey for functional kleptoplasty among West Atlantic Ascog- lossa (=Sacoglossa) (Mollusca: Opisthobranchia). KERRY B. CLARK, KATHE R. JENSEN, AND HUGH M. STIRTS NR eee aris tues cu hawne eee Gers 339 Range limits and dispersal of mollusks in the Aleutian Islands, Alaska. GEERAT J. VERMEIJ, A. RICHARD PALMER, AND DaviD R. LINDBERG 346 Burrowing times of Donax serra from the south and west coasts of South Africa. THEODORE E. DONN, JR. AND SHALEEN F. ELS 355 Chemoautotrophic sulfur bacteria as a food source for mollusks at intertidal hydrothermal vents: evidence from stable iso- topes. GEOFFREY C. TRAGER AND MICHAEL J. DENIRO 359 Survival, growth, and fecundity of the West Indian topshell, Cittartum pica (Linnaeus), in various rocky intertidal hab- itats of the Exuma Cays, Bahamas. ADOLPHE O. DEBROT 363 The diet and feeding selectivity of the chiton Stenoplax heathiana Berry, 1946 (Mollusca: Polyplacophora). BARRY FOLSOM PUTMAN ill Movement patterns of the limpet Lottia asm: (Middendorff): net- working in California rocky intertidal communities. DAVIDPRPLEINDBERG Be oi ana Ae a see waeeic acre: 375 Sexual dimorphism in Pomacea canaliculata (Gastropoda: Am- pullariidae). INESTIOR I CAZZANIGA tet aigy Acie uu See asic: 384 The use of tetracycline staining techniques to determine statolith growth ring periodicity in the tropical loliginid squids Lo- liolus noctiluca and Loligo chinensis. GEORGE DAVID JACKSON ........................ 389 The eastern Pacific species of the bivalve family Spheniopsidae. UGENE WSIG@OAND Ware yas ton, oo se ha ese aoe 394 The molluscan publications and taxa of Lorenzo Gordin Yates (1837-1909). EUGENE V. COAN AND PAUL H. Scotr ............ 402 On Cochlicopa lubrica (Miller, 1774) and Cochlicopa lubricella (Porro, 1837) (Gastropoda: Pulmonata: Cochlicopidae) in the Sierra de O Courel (Lugo, NW Spain). A. OUTEIRO, S. MATO, I. RIBALLO, AND T. RODRIGUEZ 408 Tattoo ink as a tag for nudibranchs. Kim KIEST AUTHOR INDEX BEEAKNEY, “J:0S5 00.005 oho eI ae ethene aes eee ae 111 BOUGHET A Bane Sen te ery enn ee ae ee 56 CAZZANIGA SNES Jno a tence Meer Rees 384 GEARK? Kee Bet en awe seca icine eet EO Re Rn a ea ne 339 COANSES VAS aos aoe eee 20, (127), 174, 394, 402 TEBROT As! @ roa oe lease ah een neryea) Sas era ale Pa eee 363 DENTRO SME, Je ie es ate arenes ee cee cre eee cea 359 ONIN 01g OR eecsiece he che tegen ye at otra ei a re arate 355 EVAN oR 5 VES acco aan eet ay we trope ein H caret A ae ey ta 209 EBERSOLE.) VP hese Rie eet Roe Ren eee 215 EDDER AW AIRE 25 irate se eeicee ins ee ae ee 293 1 DR ECEa io yial Ske aan cece Gee ecstasy olen arto hates Sue emer ei 355 ESMIERSON(S Witte epee cl ardent see er ee es Re a 145 ERANZ Gh ene ae inte ee eee ei ees Sek aR ener 155 GARGIAS Fe yici ta outencas oe sites hacen, casei wstc ener eer 166 GARCIA= VOKES (1956:763) dated Thracia from SOWERBY (1823), and KEEN (1969:850) followed this. However, VOKES (1967:339; 1980: 211) later changed his mind and regarded Sowerby’s use of the name as being a nomen nudum, a conclusion with which I agree. Sowerby said only that some of his fossils resemble Leach’s genus Thracia, which has an external ligament and which has been associated with Anatina by Lamarck. This is not sufficient in- formation to recognize any genus; there is no figure attributed to Thracia; and no species are included in it. The type species of Thracia has also been variously interpreted. BLAINVILLE (1824) included two species—7. corbuloidea and T. pubescens. I regard the first of these as being a nomen nudum in this work. The type species is thus fixed by monotypy as 7. pubescens. Blainville says of 7. corbuloidea only that it belongs in a group of Thracia containing ““Espéces qui n’ont qu’un cuilleron sur une valve” [Species which have a resilifer only in one valve]. This does not characterize any 7hracia, including the species that Blainville eventually made available under this name in 1827. Perhaps he was confused by some unusual specimen of Mya. This same information appears in the text of BLAINVILLE’s Man- uel (1825), and here he cites a figure. However, his plates were evidently not published until 1827, only then making the name available. Some workers have considered 7. corbuloidea to be available in 1824, perhaps believing Blainville’s description of the genus to cover this particular species, or perhaps simply not questioning the adequacy of the one-line description. Some of these authors also considered BLAINVILLE’s (1827) “‘restriction” of his concept of Thracia to T. corbuloidea to constitute a type designation. However, the modern JCZN does not allow this method of designation. (If 7. corbuloidea were to be considered available in 1824, then the type would have been fixed as 7. pubescens by the subsequent designation of ANTON, 1838:2). * The genus “‘Osteodesma Blainville, 1825,” was placed by KEEN (1969:850) as an objective synonym of Thracia, with the same type species by original designation. (The ‘““Nouvelles Additions et Corrections” section of Blainville’s Manuel is correctly dated 1827, not 1825 as is the rest of the volume.) The path to the designation of the correct type species of this genus is more complex than has been assumed, and it has never been clearly delineated. Blainville did not use the word “type” in the modern sense, and he employed it in connection with two different nom- inal species: (a) Anatina myalis Lamarck, 1818, on p. 659, and (b) Mya “declives” [misspelling of declivis] PENNANT, 1777:79, on p. 660. Blainville regarded the latter as being an older name for Mya pubescens of Pulteney and of MONTAGU (1803:40-41). Later authors have regarded Lamarck’s A. myalis as being a junior synonym of Mya pubescens; “Mya pubescens Pulteney,” of Mon- tagu, as being a synonym of Thracia phaseolina (LAMARCK, 1818: 492), or of its synonym Tellina papyracea POLI, 1791 (p. 43; pl. 15, figs. 14, 18) (non GMELIN, 1791:3231); and Mya declivis Pennant as a nomen dubium, although perhaps an earlier name for Thracia phaseolina. (I regard Pennant’s taxon as being most probably a junior synonym of Mya arenaria LINNAEUS, 1758: 670.) Thus, the type species of Osteodesma is not fixed by original designation, and two more species were also included by Blain- Page 24 No other species of 7hracia is closely similar to the type species of the genus, the large, thick-shelled eastern At- lantic 7. pubescens, in which the external ligament is partly sunken in a projecting resilifer. The right valve is some- what more inflated. The pallial sinus is shallow. I have seen one specimen 92 mm in length (SBMNH 25955). Analysis of the subgeneric names under 7hracia and the proposal of additional subgenera is a task that awaits a world-wide review of this family. I leave the following two Arctic-Boreal species in 7hracia s.]. They have a ligament without a projecting resilifer, no evident lithodesma in the adult, and a pustulose external surface, Thracia devexa Sars, 1878 (Figures 3-5) Thracia truncata [Brown] var. devexa Sars, 1878: SARS, 1878: 84-85; figs. lla, b; LAMy, 1931:296-297; SooT-RYEN, 1941:23-25, 39; pl. 12, figs. 5-10; pl. 4, fig. 4; pl. 9, fig. 5 [as Thracia devexa]; OCKELMANN, 1959:156-158; pl. 3, fig. 5; fig. 11; BERNARD, 1979:60- 61; fig. 105; LuBINSKy, 1980:48, 93, 105; pl. 11, fig. 8; map 40; BERNARD, 1983:64 [as Thracia (Ixartia)]. Thracia “sp. aff. T. truncata Brown (=T. myopsis Moller)”: MAcNEIL, 1957:106; pl. 11, figs. 9, 11, 15. [non (BROWN, 1844), non MOLLER, 1842—see synonymy under next species. | Type material and locality: Zoological Museum, Oslo, D.13738, Type 41/2, holotype, pair; length, 28.0 mm; height, 21.8 mm; convexity, 12.9 mm (Figure 3). SooT-RYEN (1941:24-25) gives reasons for believing that this specimen is Sars’ holotype, in spite of the fact that it does not match the 34 mm length given by Sars. Vadso, Varanger Fjord, northern Norway (70°5'N, 29°47’E). Description: Medium-sized (length to 40 mm; OCKEL- MANN, 1959; east Greenland; largest specimen from study area: 26.7 mm; LACM 71-482; Arctic coast of Alaska); thin; right valve slightly more inflated; approximately equilateral; posterior end somewhat produced, truncate; beaks prominent; pallial sinus shallow; surface almost smooth, with irregular growth lines, generally with sparse, well-spaced pustules, especially on posterior slope; peri- ostracum tan; pallial sinus broad, shallow. Hermaphroditic; planktonic stage either short or lacking (OCKELMANN, 1959:158). I here illustrate specimens from the Beaufort Sea (Fig- ure 4) and British Columbia (Figure 5). Distribution and habitat: From northernmost Norway south to Skjerstadfjord (67°13'N), as well as in Novaya ville in this genus: Anatina “‘trapezoidalis” [misspelling of trape- zoides| LAMARCK, 1818:464, now regarded as a synonym of Peri- ploma margaritacea (LAMARCK, 1801:137), and Anatina rupicola LAMARCK, 1818:465, a synonym of Thracia distorta (Montagu, 1803). The first subsequent designation I have located is that of DALL (1903): Anatina myalis Lamarck. Because this species is most probably a synonym of Thracia pubescens (Pulteney), Os- teodesma is indeed an objective synonym of Thracia s.s. the Veliger; Vols 335 Nom Zemlya, Spitsbergen (Svalbard) (SOOT-RYEN, 1941:39); east Greenland (from 73°16'N to about 60°N) (OCKEL- MANN, 1959); northern Canada (from 82°N south into Hudson Bay to 54°N) (LUBINSKY, 1980). On the Arctic Coast of Alaska from off Barter Island, Beaufort Sea (72°15'30"N, 143°39'36"W) (LACM 71- 397), to off ‘Pitt Point (71° 14/427 Nis 225313 0p) (SBMNH 35085); in the Bering Sea, from the Navarin Basin (60°26'36"N, 178°17'36"W) (UAM 4709) to SE of the Pribilof Islands (56°13’N, 168°20’W) (UAM 4710). Evidently, isolated populations are on the coast of British Columbia: Shidegate Inlet, Queen Charlotte Islands (53°19'30"N, 131°6’W) (LACM 60-113.1); off south- eastern Vancouver Island (49°57'N, 123°38’W) (LACM 69-126.1). The depth range for material from Arctic Alas- ka through British Columbia is from 7 to 348 m (mean, 80 m). The only bottom type recorded is mud. I have seen 15 lots from the study area. This species has been recorded in the Nuwok Formation on the Arctic coast of Alaska as “Thracia sp. aff. T. truncata Brown (=T7. myopsis Moller)” by MACNEIL (1957). This formation is now regarded as being of late Miocene or early Pliocene age (L. Marincovich, verbal communica- tion, Oct. 1988). Discussion: I am reporting herein the first records of this uncommon species from the northern Bering Sea and from British Columbia. Future studies may possibly connect these populations. Thracia myopsis Moller, 1842, ex Beck MS (Figures 6-10) Thracia myopsis Moller, 1842, ex Beck MS: MOLLER, 1842:94 [as “Beck”’]; REEVE, 1859:pl. 1, fig. 5a, b; CONRAD, 1869:54; Lamy, 1931:296-297; Soot-RYEN, 1941:22-23, 38-39; pl. 2, figs. 1-4; pl. 6, fig. 4; pl. 8, fig. 4; FitaTova, 1957:57; OCKELMANN, 1959:155-157; pl. 3, fig. 4; fig. 11; MACGINITIE, 1959: 162-163; pl. 23, fig. 9; pl. 24, fig. 4; BERNARD, 1979: 60-61; fig. 106; LuBINSKy, 1980:48-49, 94; pl. 11, fig. 11; map 41; SCARLATO, 1981:287-288; fig. 156; BERNARD, 1983:64 [as Thracia (Ixartia)]; THEROUX & WIGLEY, 1983:55, 121; fig. 107; SCARLATO, 1987:234; fig. 143. Thracia couthouy: Stimpson, 1851: STImPSON, 1851a:8; STIMPSON, 1851b:13; CONRAD, 1869: 54; DALL, 1903:1525 [as a synonym of 7. myopsis]. “Thracia truncata Brown,” auctt., non Brown, 1844 [Turton, 1822]: Sars, 1878:84-85; pl. 6, figs. 10a, b [as “forma typica’’]. [non BROWN, 1844:110; pl. 42, fig. 28, which is Anatina truncata TURTON 1822:46-47, 277; pl. 4, fig. 6, a syn- onym of Thracia (Ixartia) distorta (Montagu, 1803), as well as a junior primary homonym of Anatina truncata LAMARCK, 1818:463.] Thracia bering: Dall, 1915: Cooper, 1894:[12] [nomen nudum]; DALL, 1915:442- 443; I. OLDROYD, 1924:28; pl. 7, fig. 3; I. OLDROyYD, 1925:85; pl. 43, fig. 4; LAMy, 1931:233-234; SCARLATO, 1981:287 [as a synonym of 7. myopsis]; BERNARD, 1983: 64 [as Thracia (Crassithracia)]. Raises Tage Explanation of Figures 3 to 7 Figures 3-5. Thracia (s.1.) devexa Sars. Figure 3: Holotype; ZMO D.13738 (Type 41/2); length, 28.0 mm. Figure 4: SBMNH 35085; off Pitt Point, Beaufort Sea; 40 m; length, 24.4 mm. Figure 4a: Close-up view of posterior slope of left valve. Figure 5: LACM 69-126.1; off SE coast of Vancouver Island, British Columbia; 45 m; length, 25.9 mm. Figures 6, 7. Thracia (s.l1.) myopsis Moller. Figure 6: Lectotype (herein); ZMC no number; length, 17.7 mm. Figure 7: Lectotype (herein) of 7. bering: Dall; USNM 221555; length, 36.6 mm. Page 26 Type material and localities: 7. myopsis—Zoologisk Museum, Copenhagen (ZMC) no number, lectotype (herein); length, 17.7 mm; height, 13.2 mm; convexity, 7.0 mm (Figure 6); ZMC, paralectotypes, 18 pairs, 17 valves in same lot, plus three other lots; BM(NH) 1843.6.23.204, probable paralectotype; BM(NH) 1988041/1-3, probable paralectotypes, including the specimen figured by Reeve. Greenland; the lot from which a lectotype was selected, the largest available intact pair, has no exact locality. The type locality is here clarified as Kap Hope, Scoresby Sund (70°29'N, 22°17'W), where OCKELMANN (1959: 155) obtained the species. T. couthouyi—Lost in the Chicago fire of 1871 (DALL, 1888:132-133). Massachusetts Bay (approx. 42°20'N, 70°30'W). T. beringi—USNM 221555, lectotype (herein), pair; length, 33.6 mm; height, 24.6 mm; convexity, 11.6 mm (Figure 7). USNM 859378, paralectotypes, 2 pairs, 6 valves; SBMNH 34449, paralectotype, 2 valves. Kiska Harbor, Kiska Island, Rat Islands Group, Aleutian Is- lands, Alaska (51°58'N, 177°34’E). DALL (1915) cited a type locality of the “Commander Islands,” but no material from this locality is in the USNM collection. The lot from which a lectotype is selected bears the only number listed in the original description, and it was labeled “‘type.” Description: Medium-sized (length to 40 mm; USNM 271727; St. Paul Island, Alaska, about 5 mm larger than any reported from elsewhere); average in thickness; right valve larger, more inflated; approximately equilateral; an- teroventral margin often somewhat sinuous; posterior end slightly truncate; beaks lower and less prominent than in Thracia devexa; pallial sinus shallow; surface with growth lines and dense pustules, particularly on posterior slope and in young specimens. Specimens from populations in the southern Bering Sea and the Aleutian Islands are often more elongate and thick- er-shelled than is Thracia myopsis from the Arctic Ocean, and the surface sculpture is often worn off. This was named T. beringi by DALL (1915). However, such material in- tergrades with more typical 7. myopsis, and the densely pustulose young specimens from these populations are identical to young 7. myopsis from other areas. SCARLATO (1981) was the first to synonymize these two taxa. Hermaphroditic, with a short or absent planktonic stage (OCKELMANN, 1959:156). he Veligers Volh335 Nom I have illustrated here specimens from the Bering Sea (Figure 8) and from southeast Alaska (Figure 9), and a juvenile specimen showing a lithodesma (Figure 10). Distribution and habitat: Circum-Arctic: In the Barents, White, Kara, Laptev, East Siberian, and Chukotsk seas (FILATOVA, 1957); Novaya Zemlya, Spitsbergen (Sval- bard), Jan Mayen, and in the eastern Atlantic, south to Bergen, Norway (60°N) (SooT-RYEN, 1941); Iceland; in eastern Greenland south to Tasissaq, near Angmagssalik (about 65°N) (OCKELMANN, 1959); in the western Atlantic south to off Massachusetts (about 42°N) (THEROUX & WIGLEY, 1983); in west Greenland south to 66°N, and in eastern Canadian Arctic south into southern Hudson Bay to about 54°N (LUBINSKY, 1980)°; in the western Pacific south to Zaliv Petra Velikogo (about 43°N) (SCARLATO, 1981). Throughout the Arctic coast to Alaska, the Bering Sea, and Aleutian Islands, south to Uncluelet, Barkley Sound, Vancouver Island, British Columbia (48°55'N, 125°33’'W) (LACM 67-199.1). However, there is a single, small right valve from off Point Loma, California (about 32°33'N, 117°40'W) (USNM 208957)! In the area of study, from the intertidal zone to 183 m (mean, 44 m), with most records from sand and mud bottoms, but some recorded on gravel or among rocks. It is not uncommon; I have seen 101 lots from the study area. THEROUX & WIGLEY (1983) found this species in the western Atlantic from 95 to 114 m (mean, 105 m) on gravel substrates. Subgenus (Cetothrax) IREDALE, 1949:19 Type species: Thracia alciope ANGAS, 1872:611; pl. 42, fig. 6; =Anatina imperfecta LAMARCK, 1818:464; by original designation—western Australia. Shells medium-sized to large. The right valve is larger and more inflated. The ligament is entirely external. It differs from Thracia (Crassithracia) in its greater size and inflation and from other subgenera in its lack of conspic- uous pustules on the external shell surface. The type species of this genus is more elongate than the eastern Pacific > LUBINSKY’s (1980) map illustrating the distributions of Thra- cia myopsis and T. septentrionalis (map 41) does not match the distributions of these taxa given in the text. Evidently, the symbols for the taxa in the map’s legend were accidentally reversed. Explanation of Figures 8 to 12 Figures 8-10. Thracia (s.1.) myopsis Moller. Figure 8: USNM 859368; USGS Loc. M.6283; Bering Sea off Seward Peninsula; length, 29.2 mm. Figure 8a: Close-up of posterior slope of right valve. Figure 9: USNM 220579; Sitka, Alaska; length, 31.0 mm. Figure 10: LACM 73-23; Kachemak Bay, Cook Inlet, Alaska; 9 m; close-up of lithodesma in left valve of a juvenile specimen; scale bar, 200 um. Figures 11, 12. Thracia (Cetothrax) condoni Dall. Figure 11: USNM 214143; near Westport, Clatsop Co., Oregon; Pittsburg Bluff Formation; middle Oligocene; length, 46.7 mm (from Moore, 1976:pl. 16, fig. 1). Figure 12: LACM 57-12.7; Kasitsna Bay, Cook Inlet, Alaska; 46 m; length, 67.0 mm. E. V. Coan, 1990 Page 27 Page 28 species and has a shorter ligament and a less conspicuous periostracum. Thracia (Cetothrax) condoni Dall, 1909 (Figures 11, 12) Thracia condoni Dall, 1909: DALL, 1909:135; pl. 19, fig. 5; DALL, 1915:447; B. CLARK, 1918:137; pl. 11, fig. 12; pl. 12, fig. 2; TEGLAND, 1933: 113; pl. 6, fig. 5; WEAVER, 1943:119; pl. 25, fig. 10; pl. 29, fig. 15; DURHAM, 1944:141; pl. 13, fig. 6; ZHIDKOVA et al., 1968:137, 159, 174; pl. 7, fig. 4; pl. 43, fig. 2; HICKMAN, 1969:72-73; pl. 9, figs. 10-14; MoorE, 1976: 54; pl. 16, figs. 1, 3. Thracia (Crassithracia), n. sp.: ROTH, 1979:410-412; pl. 8, fig. 2 [thesis]. Type material & locality: 7. condoni—USNM 110460, holotype, cast of a right valve, length 60 mm; height 44 mm; convexity, 8 mm (not refigured here). Smith’s Quarry, Eugene, Lane Co., Oregon (approx. 44°3'N, 123°4'W); Eugene Formation; early to middle Oligocene; T. Condon & C. A. White. Description: Large (to 83 mm in length; LACM 60-68.1; Nuka Island, Kenai Peninsula, Alaska); thin-shelled for size; right valve decidedly more inflated; approximately equilateral; posterior end broadly truncate; beaks promi- nent; pallial sinus short, narrow; surface smooth, with only concentric growth lines; periostracum light tan, darker on posterior slope. I here figure a specimen from the Oligocene of Oregon (Figure 11) and a Recent specimen from British Columbia (Figure 12). Distribution and habitat: In the Recent fauna, this species is known from St. Paul Harbor, Kodiak Island (57°44'24’N, 152°25'42”W) (USNM 221313), through the Gulf of Alas- ka, as far north as Kasitsna Bay, Cook Inlet (59°29'N, 151°36'W) (LACM 57-12.7), south to Clover Point, Vic- toria, British Columbia (48°24'N, 123°21'W) (CAS 066617). Some of the available material was collected washed up on the shore, with depths recorded for only 5 lots—16 to 81 m (mean, 48 m). The bottom type is recorded for only 1 lot: mud. In the Recent fauna, this rare species is known from only 13 lots representing 10 stations. Discussion: This species has long been present in collec- tions from Alaska and British Columbia, generally iden- tified as Thracia challisiana Dall. Conspecific material is present in late Pliocene and Pleistocene strata of north- western California and in Pleistocene strata of south- western Oregon (ROTH, 1979, as “Thracia (Crassithracia), n. sp.”). Here I tentatively place this Recent and fossil material into 7. condoni, which has been reported from a number of formations of Oligocene age in Alaska, Wash- ington, Oregon, and California (MOORE, 1976, reviews these records). It has also been reported from deposits of Miocene age in Siberia (ZHIDKOVA et al., 1968). Future study may demonstrate whether there are suf- The Veliger, Vol. 33, No. 1 ficient morphological differences to regard Pliocene- Recent material as a separate species. Subgenus (Crassithracia) SOOT-RYEN, 1941:19 Type species: Thracia crassa BECHER, 1886:71, 82; pl. 6, figs. 1, 1a-c; by original designation; =7hracia septentrionalis crassa Becher, 1886 (herein) (Figure 13)—Jan Mayen Island, Arctic Atlantic. This subgenus contains only the following species and its subspecies, Thracia (C.) septentrionalis crassa. It is char- acterized by its smooth shells, without pustules. The shells are often thickened, and in most material the periostracum is shiny. The shells of this subgenus are smaller, thicker, and less inflated than those of Thracia (Cetothrax). Thracia (Crassithracia) septentrionalis Jeffreys, 1872 (Figures 13-18) Thracia truncata Mighels & Adams, 1842, non Anatina trun- cata Turton, 1822, a Thracia: MIGHELS & ADAMS, 1842:38; pl. 4, fig. 1; MIGHELS & ADAMS, 1843:48; CONRAD, 1869:55; CLENCH & TURNER, 1950:353-354; pl. 43, figs. 5-7. [non Anatina truncata TURTON, 1822:46, 277; pl. 4, fig. 6, a synonym of Thra- cia (Ixartia) distorta (Montagu, 1803), and a junior pri- mary homonym of A. truncata LAMARCK, 1818:463.] Thracia septentrionalis Jeffreys, 1872 [new name for Thracia truncata Mighels & Adams, 1842, non “Brown, 18277”): JEFFREYS, 1872:238; SOOT-RYEN, 1941:19-22, 38; pl. 1, figs. 9, 10; pl. 6, fig. 2a, b; pl. 8, fig. 3a—e; OCKELMANN, 1959:153-155; pl. 3, fig. 1; LUBINSKY, 1980:49, 94; map 41; THEROUX & WIGLEY, 1983:55-56, 121, 169; fig. 107; tables 306, 307. Thracia crassa Becher, 1886: BECHER, 1886:71; 82; pl. 6, figs. 1, 1a-c; SOOT-RYEN, 1941:19-22 [as a possible synonym of 7. septentrionalis]. Macoma truncaria Dall, 1916: DALL, 1916a:37 [nomen nudum]; DALL, 1916b:414; I. OLDROYD, 1925:177; COAN, 1969:281-282 [as Thracia (Crassithracia)]; BERNARD, 1983:64 [as a synonym of 7. bering: Dall]. ?Thracia seminuda Scarlato, 1981: SCARLATO, 1981:288-289; fig. 157. Type material and localities: 7. truncata Mighels & Adams (and 7. septentrionalis)—MCZ 165595, lectotype (CLENCH & TURNER, 1950), pair; length, 14.5 mm; height, 10.8 mm; convexity, 5.2 mm (Figure 14). Casco Bay, Cum- berland Co., Maine (43°45'N, 70°11'W); stomachs of haddock; 1840; 4-5 specimens. T. crassa—Naturhistorisches Museum, Vienna (NMV) (646) 61967, lectotype (herein), pair; length, 24.1 mm; height, 19.5 mm; convexity, 10.6 mm (Figure 13); NMV, paralectotypes, a smaller pair in the same lot, plus three pairs in NMV 117. Jan Mayen Island, Atlantic-Arctic (70°59'N, 8°40'W); on the beach. ° This name is not present in the first edition of BROWN, pub- lished in 1827. Instead, it first appears in the second edition, published in 1844. However, it is not a new species, merely a reassignment of Anatina truncata Turton, 1822, to Thracia. | | > \S rt aes Explanation of Figures 13 to 18 Figure 13. Thracia (Crassithracia) septentrionalis crassa Becher. Lectotype (herein) of Thracia crassa; MNV (646)61967; length, 24.1 mm. Figures 14-18. Thracia (Crassithracia) septentrionalis septentrionalis Jeffreys. Figure 14: Lectotype of 7. truncata Mighels & Adams (septentrionalis Jeffreys); MCZ 165595; length, 14.5 mm. Figure 15: Holotype of Macoma truncaria Dall; USNM 210916; length, 15.0 mm. Figure 16: Holotype of 7. seminuda Scarlato; length, 20.1 mm (from SCARLATO, 1981). Figure 17: CAS 066618; Bristol Bay, Alaska; 49 m; length, 13.0 mm. Figure 18: USNM 859369; USGS Loc. M.6077; Bering Sea, off Seward Peninsula, Alaska; 30 m; length, 28.0 mm. Page 30 Macoma truncarra—USNM 210916, holotype, broken pair; length, 15.0 mm; height, 10.5 mm; convexity, ap- proximately 4.5 mm (Figure 15). Between Cape Halkett and Garry “River” [Creek], Arctic coast of Alaska (70°38- 48'N, 152°11-27'W). T. seminuda—Presumably Zoological Institute, Lenin- grad (Figure 16). Zaliv Petra Velikogo, USSR, Sea of Japan (approx. 42°N, 132°E). Description: Small (length to 28.0 mm; USNM 859369; off Cape Woolley, Bering Sea, Alaska); oblong; relatively thick shelled; right valve more inflated; longer, sharply rounded anteriorly; posterior end truncate, produced in some; beaks produced; surface smooth, with growth lines only; periostracum light to dark tan, often shiny in Arctic populations, silky in some southern populations; pallial sinus moderately deep, extending past median line; inner ventral margin with vertical striations. Hermaphroditic, with a short or absent planktonic stage (OCKELMANN, 1959:155). I have illustrated here two specimens from the Bering Sea (Figures 17, 18). Distribution and habitat: Probably circum-Arctic: Ba- rents, White, and Kara seas (FILATOVA, 1957); Spitsbergen (Svalbard), Jan Mayen, Iceland (SooT-RYEN, 1941); in east Greenland south to Tasissaq, near Angmagssalik (about 65°N) (OCKELMANN, 1959); in the western Atlantic south to off Rhode Island (40°N) (THEROUX & WIGLEY, 1983); in west Greenland south to about 64°N; in the Canadian Arctic to northern end of Hudson Bay (about 63°N) (LUBINSKY, 1980) (see footnote 5). If Thracia semi- nuda Scarlato, 1981, is indeed a synonym, this species occurs as far south as Zaliv Petra Velikogo, USSR, in the Sea of Japan (about 43°N). In the study area, this species is known from six stations on the Arctic coast of Alaska and nine lots from the Bering Sea; south to Popoff Strait, Shumagin Islands (south of the Alaska Peninsula; about 55°15’N, 160°10’W) (USNM 859374). It is recorded from 11 to 69 m (mean, 35 m). The only bottom types recorded are sand and sandy silt. I have examined 16 lots from the study area. THEROUX & WIGLEY (1983) found this species in the western Atlantic between 23 and 74 m (mean, 54 m), chiefly on sand. The taxonomic situation here may be more complicated than can be expressed with a single name at the species level, but there is as yet insufficient material to justify recognizing more than one species. However, I here rec- ognize Thracia septentrionalis crassa as a subspecies; no material from anywhere else reaches the extreme thickness represented by its type material from Jan Mayen Island. Material from New England, the type locality of 7. sep- tentrionalis, has a silky periostracum, not shiny as in Arctic specimens. Large adult specimens from the Bering Sea (USNM 859369) (Figure 18) are also silky rather than shiny. Thracia seminuda Scarlato, 1981, seems to be of this form as well. The Veliger, Vol. 33, No. 1 A related fossil species is Thracia transversa LEA, 1845 (p. 237; pl. 34, fig. 11), described from the Yorktown Formation at Petersburg, Virginia (types, ANSP 1585), now regarded as being of Pliocene age. As pointed out by GARDNER (1943:44), 7. transversa is much smaller, none yet found being larger than 10 mm. Additionally, Lea’s species is thinner for its size and is longer posteriorly (based on examination of USNM 164645). Thracia brioni WARD & BLACKWELDER (1987:161-162; pl. 29, figs. 7-10) has been proposed for material from the late Pliocene and early Pleistocene of North Carolina that is very similar to 7. transversa. Subgenus (Homoeodesma) FISCHER, 1887:1171 Type species: Thracia conrad: COUTHOUY, 1839:153-158; pl. 4, fig. 2; by monotypy—eastern Atlantic. This subgenus is characterized by species with relatively large shells that are inflated anteriorly. They have a lig- ament that is external in the adult and does not project below the hinge margin in a resilifer. The shell surface is generally very pustulose. In addition to the two eastern Pacific species discussed below, the following six Recent taxa belong in this subgenus: T. (H.) conradi COUTHOUY, 1839—western Atlantic T. (H.) convexa Woop (1815:92; pl. 18, fig. 1)—Europe T. (H.) corbuloidea BLAINVILLE, 1827:pl. 76, fig. 7 [1825: 565|—Mediterranean T. (H.) itoi HABE, 1962:143; App., p. 40; pl. 64, fig. 17— Japan T. (71.) khakumana (YOKOYAMA, 1927:168, 177-178, 182; pl. 47, fig. 14)—Japan T. (H.) stearnsi DALL, 1886:307; DALL, 1890:275; pl. 13; fig. 2—western Atlantic. The morphology and behavior of Thracia (H.) conradi was discussed by Morse (1913, 1919) and THOMAS (1967). Thracia (Homoeodesma) trapezoides Conrad, 1849 (Figures 19, 20) Thracia trapezoides Conrad, 1849: CONRAD, 1849c:723; pl. 17, fig. 6a [no 6b present]; CAR- PENTER, 1857b:367; CARPENTER, 1864b:679 [1872:165] [as a possible synonym of 7. curta]; DALL, 1909:135; pl. 2, fig. 14; pl. 13, fig. 7; DALL, 1915:447; I. OLDROyYD, 1924:27; pl. 7, fig. 2; I. OLDROYD, 1925:84-85; pl. 43, fig. 8; GRANT & GALE, 1931:257-258, 906; pl. 13, fig. 8; Moore, 1963:84-85; pl. 26, fig. 3; pl. 31, fig. 6; BERNARD, 1983:64. ?““Thracia ventricosa Conrad,” auctt., non T. ventricosa Phil- ippi, 1844: MEEK, 1864:11 [nomen nudum]. [non PHILIPPI, 1844: 17.] ?Thracia jacalitosana Arnold, 1910: ARNOLD, 1910:68-69; pl. 16, fig. 4; DALL, 1915:447. Thracia kanakoffi Hertlein & Grant, 1972: HERTLEIN & GRANT, 1972:338-339; pl. 42, figs. 11, 13-15. Type material and localities: 7. trapezoides—USNM 3604, holotype, a cast, mold, and latex impression of mold; E. V. Coan, 1990 length, 33.8 mm; height, 26.7 mm; convexity, 15.5 mm (Figure 19); paratypes, USNM 561515, 3 specimens. As- toria, Klatsop Co., Oregon (46°10’N, 123°45’W); Astoria Formation; middle Miocene; J. D. Dana; 1841. T. jacalitosana—USNM 165579, holotype, broken left valve; length, 51 mm; height, 44 mm; convexity, 8 mm [not refigured here]. USGS Loc. 4763, “on Stone Canyon and Coalinga Road,” 183 m N of Jacalitos Creek crossing, 22.5 km SW of Coalinga, either Fresno Co. or Monterey Co., California (approx. 36°2’30"N, 120°29'W); “Jacali- tos” [Etchegoin] Formation; “Upper Miocene”; R. Arnold and F. Stokes, Jr. T. kanakofi—LACM 4839, holotype, pair; length, 81.0 mm; height, 54.2 mm; convexity, 32 mm [not refigured here]. LACM 4840-4881, paratypes. LACM Loc. 291; 0.8 km S of Humphrey Railroad Station, Los Angeles Co., California (34°24'18”N, 118°26'21”W)); silt beds exposed in a gully in the center of the S half of Sec. 27, T.4N., R.15W.; Pico Formation; middle Pliocene. The paratypes are from both the Pico Formation, Los Angeles Co., and the San Diego Formation, San Diego Co. Description: Large (length to 65 mm in Recent material; CAS 066634; Departure Bay, Vancouver Island, British Columbia; and to 132 mm in Pliocene material from San Diego Co., Calif.; HERTLEIN & GRANT, 1972); trapezoi- dal; approximately equilateral; anterior end inflated, rounded, and often with a slight flexure and radial depres- sion about one-third of distance to posterior end; posterior end produced, truncate, set off by a flexure and, posterior to it, a low ridge; beaks prominent; sculpture of prominent pustules, particularly dense on posterior slope; periostra- cum dark tan, darker on posterior slope; pallial sinus mod- erate in size, barely reaching a vertical line from beaks in some specimens. A Recent specimen from British Columbia is illustrated here (Figure 20). Distribution and habitat: This eastern Pacific species is known from Wide Bay (S side of Alaska Peninsula), Alas- ka (57°22'’N, 156°11’W) (CAS 066619), throughout the Gulf of Alaska, along the coasts of British Columbia, Washington, Oregon, and California, south to off Isla Cedros, Baja California Norte (28°19’N, 115°10’W) (LACM 71-154). Specimens from populations in the shel- tered waters of Puget Sound and the islands of British Columbia attain the largest size; individuals in populations elsewhere are smaller. The species has been recorded from 11 to 199 m (mean, 71 m) on sand and mud bottoms, the latter predominating. I have examined 129 Recent lots. Thracia trapezoides has been recorded in a number of formations of middle Miocene to Pliocene age from Wash- ington, Oregon, and California. These are not detailed here. It may date from the Oligocene if 7. schencki B. Clark, 1932, proves to be a synonym (see Discussion). It is also recorded from early Pleistocene strata in south- ern California: A. CLARK (1931:opp. p. 30), Hoots (1931: 120), RoDDA (1957:2484), and WoOoODRING et al. (1946: 85). Page 31 Discussion: 7hracia kanakoffi was differentiated by its au- thors from Recent specimens—not from material from the type locality of the species in the Miocene of Oregon— because of its (1) larger size; (2) less steeply sloping pos- terodorsal margin; (3) the presence of a ridge on the pos- terodorsal margin of the right valve; and (4) its less de- veloped radial depression anterior to the ridge defining the posterior slope. This Pliocene population clearly attained a larger size than any Recent (or Miocene) material yet observed. However, some Recent material also has a less steeply sloping posterodorsal margin than other specimens, and the posterior slope of the type of 7. trapezoides and other specimens that have been illustrated from the Mio- cene of Oregon seem not to have a steeply sloping pos- terodorsal margin. The degree of production of the pos- terior end, and the extent of the radial depression setting it off, is also variable in Recent material, as is the strength of posterodorsal ridges in both valves. With this degree of variability, and with the lack of a case for differentiating Pliocene material from specimens from the Miocene of Oregon, I think it unwise to recognize a separate species from the Pliocene of southern California. It is possible that Thracia jacalitosana ARNOLD, 1910 (pp. 68-69; pl. 16, fig. 4), described from the Miocene of central California, is a poorly preserved specimen of 7. trapezoides. The collection of additional material from its type locality would be required to prove this. Thracia schencki B. CLARK, 1932 (pp. 801, 808, 845; pl. 15, figs. 2, 3, 5), ex Tegland MS, described from the upper Oligocene of Alaska (see also TEGLAND, 1933:112-113, 154; pl. 6, figs. 6-11), is similar to 7. trapezoides and is either another synonym or ancestral to it. It was differ- entiated by its author as having a more acute umbonal angle and in lacking a shallow anterior depression. Thracia kidoensis KAMADA, 1955 (pp. 11-12, 14; pl. 1, figs. 1, 2a, b), from the Oligocene of Japan, is suspiciously similar to T. schencki. Thracia (Homoeodesma) trapezoides is closely related to the western Atlantic type species of the subgenus, 7. (//.) conradi Couthouy, 1839. Recent material of that species attains a much larger size (up to 95 mm in material that I have seen), is more inflated, has a straighter posterodorsal margin, a more conspicuous escutcheon, and a light tan periostracum. THEROUX & WIGLEY (1983:55, 120, 169; fig. 106; tables 304, 305) found 7. (H.) conrad: from 34 to 126 m (mean, 70 m), predominantly in silt. Thracia (Homoeodesma) challisiana Dall, 1915 (Figure 21, 21a) Thracia challisiana Dall, 1915: DALL, 1915:443; I. OLDROYD, 1924:27-28, 209; pl. 7, fig. 1; I. OLDROYD, 1925:84; pl. 43, fig. 7; BERNARD, 1983:64 [as Thracia (Crassithracia)]. Type material and locality: USNM 272096, holotype, pair; length, 46.2 mm; height, 35.7 mm; convexity, 19.8 mm (Figures 21, 21a). San Juan Island, San Juan Co., Washington (approx. 48°30'’N, 123°W); B. M. Challis. Page 32 The Veliger, Vol. 33, No. 1 E. V. Coan, 1990 Description: Medium-sized (length to 59.5 mm; LACM 140317; Craig, Alaska); oblong; right valve more inflated; anterior end rounded; posterior end decidedly longer in adult, broadly truncate; beaks prominent; surface with fine, conspicuous pustules (Figure 21a); periostracum dark tan; pallial sinus broad, short. Distribution and habitat: Kasitsna Bay, Cook Inlet (59°21'N, 151°33'58”W) (Baxter Coll.), and Point Wood- cock, Montagu Island, Prince William Sound, Alaska (59°54'15”N, 147°48'40”W) (LACM 65-184.1), south to type locality at San Juan Island, Washington (48° 30'N, 123° W); and from off Redondo Beach, Los Angeles Co., California (33°50’N, 118°25'W) (LACM 72-204), to off Isla Guadalupe, Baja California Norte (28°52'N, 118°17’W) (SBMNH 35086). No material is yet known from between these two sets of occurrences. This species is recorded from 29 to 229 m (mean, 72 m), with the deepest records from its southern distribution. I have ex- amined 20 lots. This species was tentatively reported (as “‘cf., juv.”’) from a formation of early Pleistocene age in southern California (VALENTINE, 1961:407). Discussion: Thracia (/7.) ito1 HABE, 1962 (pp. 143; App. 40; pl. 64, fig. 17) is a closely related Japanese species, described from Onagawa Bya, Miyagi Prefecture, with similar densely pustulose sculpture (Figure 22). It is more elongate, and it may attain a larger size, the type measuring 64.5 mm in length. The ligament is proportionately shorter and extends somewhat ventrally on a resilifer. In the area of overlap with Thracia myopsis, young spec- imens of this species can be distinguished by their more elongate ligament, straighter ventral margin, more pus- tulose sculpture, and less prominent beaks. Subgenus (Jxartia) LEACH, 1852:272 Type species: Mya distorta MONTAGU, 1803:42-44; pl. 1, fig. 1; by monotypy—eastern Atlantic. [=Rupicola FLEURIAU-BELLEVUE, 1802a:348, 354; 1802b: 106-107; genus without named species (only species name present is a vernacular). Non Rupicola BRISSON, 1760, vol. 4:437. Original list and subsequent desig- nation by RECLUZ, 1846:409, 424: Anatina rupicola LA- MARCK, 1818:465; =Mya distorta Montagu, 1803.] [=Rupicilla SCHAUFUSS, 1869:18, presumably a new name for Rupicola Fleuriau-Bellevue.] Page 33 [?=Pelopia H. ADAMS, 1868:16-17, non MEIGEN, 1800:18. Type species: P. brevifrons H. ADAMS, 1868:17; pl. 4, figs. 16, 16a, by monotypy—locality unknown. ] Rupicola Fleuriau-Bellevue, 1802, was the first generic unit proposed in the Thraciidae, but it is a homonym. Pelopia H. Adams, 1868, also a junior homonym, was synonymized by KEEN (1969:850) with Jxartia. However, it was described as having a large lithodesma, suggesting that it does not belong here. Its type specimen should be reexamined, and a replacement name provided if it proves to be a useful generic unit. Species in this subgenus are nestlers, although some taxa, such as the eastern Pacific Thracia curta, may also be free-living. Species may be distorted by their nestling habitat, and even the free-living forms show significant variability in shell form. Members of this subgenus have a projecting resilifer and sculpture of conspicuous pustules. In addition to the two eastern Pacific species, the following taxa appear to belong in this subgenus: T. (.) morrisoni PETIT, 1964:157-159; figs. 1-6—south- eastern USA [synonym: 7. corbuloidea Blainville, auctt., non Blainville, 1827] T. (21) brevifrons (H. Adams, 1868)—locality unknown T. U.) cuneolus REEVE, 1859:pl. 1, fig. 2—southern Japan and the Philippine Islands T. (.) distorta (Montagu, 1803)—eastern Atlantic [synonyms (partial list): Anatina rupicola LAMARCK, 1818:465; An- atina truncata TURTON, 1822:46-47, 277; pl. 4, fig. 6 (non LAMARCK, 1818:463); Thracia brevis DESHAYES, 1846:297; pl. 81, figs. 4-6; Thracia concentrica RECLUZ, 1853:122, 129-131, ex Fleuriau-Bellevue MS] T. (1) rudis REEVE, 1859:pl. 3, fig. 21—Malacca [Strait] T. (.) semilis CouTHOUY, 1839:150-152; pl. 4, fig. 3—south- ern Caribbean [synonyms: 7. rugosa ORBIGNY, 1846: 519, ex Conrad MS; 7. distorta Montagu, auctt., non Montagu, 1803] T. (.) n. sp.2—Argentina [ANSP 103366, 343169; LACM 78-92]. Thracia Uxartia) curta Conrad, 1837 (Figures 23-27) Thracia curta Conrad, 1837: ConraD, 1837:248; pl. 19, fig. 8; CARPENTER, 1857a: 210; CARPENTER, 1857b:194, 300, 349; CARPENTER, 1864b:540, 2602, 638 [1872:26, ?88, 124]; CoNnRaD, 1869:54; DALL, 1915:442; I. OLDROYD, 1924:27, 209; Explanation of Figures 19 to 22 Figures 19, 20. Thracia (Homoeodesma) trapezoides Conrad. Figure 19: Holotype of 7. trapezoides; USNM 3604; length, 33.8 mm. Figure 20: LACM 62-120.1; Howe Sound, British Columbia; 46 m; length, 63.7 mm. Figure 21. Thracia (Homoeodesma) challisiana Dall. Holotype; USNM 272096; length, 46.2 mm. Figure 21a: Close- up of posterior slope of right valve. Figure 22. Thracia (Homoeodesma) itoi Habe. Holotype; NSMT 53347; Onagawa Bay, Miyagi Pref., Japan; length, 64.5 mm. Page 34 The Veliger, Vol. 33, No. 1 Explanation of Figures 23 to 28 Figures 23-27. Thracia (Ixartia) curta Conrad. Figure 23: Holotype; BM(NH) 1861.5.20; length, 27.1 mm. Figure 24: Holotype of Lepton clementinum; length, 0.9 mm (Carpenter’s figure, from BRANN, 1966:pl. 14, fig. 157). Figure 25: Holotype of 7. quentinensis Dall; USNM 333112; length, 46.0 mm. Figure 26: LACM 140425; Bahia San Carlos, Sonora, Mexico; length, 37.0 mm. Figure 27: LACM 71-170; Punta Thurloe, Baja California Sur; 16 m; close-up of right valve showing lithodesma and anterior lateral tooth; scale bar, 250 um. Figure 28. Thracia ([xartia) anconensis Olsson. Holotype; ANSP 218955; length, 34.0 mm. E. V. Coan, 1990 pl. 7, fig. 4; 1. OLDRoyD, 1925:83; pl. 43, fig. 6; GRANT & GALE, 1931:258-259; HERTLEIN & STRONG, 1946: 95; HERTLEIN, 1957: 63, 74; pl. 13, figs. 7, 8; KEEN, 1958:230-231; fig. 589; KEEN, 1966a:171; KEEN, 1971: 295; fig. 760; BERNARD, 1983:64 [as Thracia (Ixartia)]. ?Lepton clementinum Carpenter, 1857: CARPENTER, 1857b:248, 308 [nomen nudum]; Car- PENTER, 1857c:110-111; KEEN, 1958:107; BRANN, 1966: 39; pl. 14, fig. 157; KEEN, 1968:396; KEEN, 1971:140- 142; fig. 326 [as ?Mysella]; BERNARD, 1983:32. Thracia quentinensis Dall, 1921: DALL, 1921:21; DALL, 1925:28; pl. 11, fig. 1; GRANT & GALE, 1931:250. Type material and localities: 7. curta—BM(NH) 1861.5.20, holotype, pair; length, 27.1 mm; height, 22.8 mm; convexity, 17.3 mm (Figure 23). Conrad was mis- taken in calling this a single valve. Santa Barbara, Santa Barbara Co., California (about 34°24’N, 119°43'W); T. Nuttall; spring 1836 (GRAUSTEIN, 1967:313-315). L. clementinum—Lost (KEEN, 1968). Carpenter said he had a single valve, but his drawings, published by BRANN (1966), show both valves (Figure 24); perhaps he guessed what the hinge of the left valve would have been like? In any event, Carpenter had damaged the specimen, and it is now missing from its BM(NH) mount. The original specimen measured 0.9 mm in length, 0.6 mm in height, and 0.5 mm in convexity (a pair might have been about 1.0 mm in convexity). Mazatlan, Sinaloa, Mexico (23°12’N, 106°25'W); off Spondylus; F. Reigen. T. quentinensis—USNM 333112, holotype, left valve; length, 46.0 mm; height, 33.3 mm; convexity, 11.7 mm (Figure 25). Bahia San Quintin, Baja California Norte (about 30°26'N, 115°56'W); Pleistocene; C. R. Orcutt, Nov. 1888. Description: Medium-sized (length to 53.4 mm; Evans Coll.; Bahia San Carlos, Sonora, Mexico; cited in DRAPER, 1987:39; 53 mm; Skoglund Coll.; Puerto Lobos, Sonora, Mexico); shells often thickened; oval to trigonal, depending on habitat; right valve generally more inflated; rounded anteriorly; slightly to decidedly longer posteriorly; poste- rior end moderately to decidedly truncate, depending on habitat; posterior end sometimes sharply separated from central slope by a radial ridge; valves sometimes twisted to the right posteriorly; ventral margin sometimes sinuous; beaks low; surface with pustules, particularly prominent on posterior slope, and concentric growth lines; pallial sinus broad, shallow; shell sometimes greenish internally. I have here illustrated a specimen from Sonora, Mexico (Figure 26). Distribution and habitat: O’ Neal Islet, San Juan Islands, Washington (48°36'N, 123°5’W) (CAS 066632); off S end of Vancouver Island, British Columbia (48°32'N, 125°2'W) (LACM 64-130.1); Monterey, California (36°38’N, 121°56’W) (LACM 72-12; CAS 066620-066622, 066631; UCMP 239; USNM 5233, 74229, 742163), southward in California and Baja California, throughout the Gulf of California, south to Punta Quepos, Puntarenas Prov., Cos- Page 35 ta Rica (9°24'43"N, 84°9'41”W) (LACM 72-58); from the intertidal zone to 48 m (mean, 12.6 m—but some lots without depth data were probably obtained from the in- tertidal zone, so this mean may be too deep). It nestles in rock crevices and empty pholad holes, but can also be found free-living on various bottom types. This is the most com- mon thraciid in collections from the eastern Pacific; I have seen 132 lots, including the type specimens. Published records of this species from Alaska were based on specimens of other species, including Thracia myopsis, T. trapezoides, and T. challisiana. This species has been reported from strata of Pleistocene age from Long Beach (T. OLDROYD, 1914:82) and San Nicolas Island (VEDDER & NorRIS, 1963:46), in Califor- nia; Bahia San Quintin, Baja California Norte (DALL, 1921, as “7. quentinensis”); and Bahia Santa Inéz, Baja California Sur (HERTLEIN, 1957). Discussion: This species displays a variety of forms, de- pending on habitat. As with most, if not all, species of Zhracia, juvenile specimens have a proportionately large lithodesma, which only later develops a predominantly external ligament. (In the case of adult 7. curta and a number of other species, the external ligament is partly seated in a projecting re- silifer.) The lithodesma is never lost but remains tiny and hard to detect. However, young specimens can be mistaken for members of unrelated groups. For example, Carpen- ter’s tiny Lepton clementinum, described from Mazatlan, Mexico, was probably a juvenile thraciid, and not, as KEEN (1971) and BERNARD (1983) thought, a Mysella, some species of which have a lithodesma (W. CLARK, 1855:145- 146). Carpenter’s specimen most likely was a young 7. curta, the distribution of which includes Mazatlan (as CAS 066633; SBMNH 35087). I here illustrate a young spec- imen of 7. curta (Figure 27), but not one as small as the type of Lepton clementinum. It also shows an anterior lat- eral tooth, present in right valves of juveniles, that dis- appears with growth. Thracia quentinensis Dall is a typical free-living form of 7. curta. Live-collected specimens similar to material from the Pleistocene of Bahia San Quintin have been found throughout the distribution of 7. curta. The type specimen of 7. quentinensis is somewhat unusual in that it is an inflated left valve, whereas in most 7. curta the right valve is the more inflated. However, in other material from the type locality of 7. quentinensis, the right valve is more inflated (SBMNH 35105). In the Panamic province, small specimens of 7. curta may be distinguished from similar- sized specimens of 7. squamosa by the former’s more pro- jecting resilifer, denser pattern of pustules, and longer posterior end. Thracia Uxartia) anconensis Olsson, 1961 (Figure 28) Thracia anconensis Olsson, 1961: OLSSON, 1961:458-459; pl. 83, figs. 4, 4a; KEEN, 1971: 295-296; fig. 758; BERNARD, 1983:64. Page 36 Type material and locality: ANSP 218955, holotype, left valve; length, 34.0 mm; height, 22.0 mm; thickness, 6.3 mm (Figure 28). Punta Ancon, Santa Elena Peninsula, Guayas Prov., Ecuador (2°20’S, 80°53’30"W), presumably washed up on the beach; A. A. Olsson, 1958. Description: Medium-sized (to 34 mm; holotype); similar to the free-living form of 7hracia curta; anterior end round- ed; posterior end longer than that in 7° curta, only slightly truncate; ventral margin evenly curved, unlike the sinuous margin of most 7. curta. As more material comes to light, perhaps from between the most southerly known station for 7hracia curta in Costa Rica and Ecuador, the relationship between the two will have to be reexamined. Distribution and habitat: Known only from the holotype. Subgenus (Odoncineta) Costa, 1829:xiv, cxxx1; pl. 2, figs. 1-4 Type species: Tellina papyracea POLI, 1791:43; pl. 14, figs. 14-18; by monotypy; non Tellina papyracea GMELIN, 1791:3231; =Amphidesma phaseolina LAMARCK, 1818: 492’—Europe. [?=Eximiothracia IREDALE, 1924:181, 199. Type species: Thracia speciosa ANGAS, 1869:48-49; pl. 2, fig. 12; by original designation—Australia. | The name of this subgenus has been subject to many misspellings and unjustified emendations, too many to list here. Species of this subgenus are elongate and thin-shelled, with a ligament that is chiefly external in the adult. A lithodesma is evident in the adult and ranges from small to fairly conspicuous. The shell surface is covered with fine pustules. Eximiothracia differs from Odoncineta only in the presence of iridescence on the inside of the valves. There are three European species, which were reviewed by SooT-RYEN (1941) and ALLEN (1961): T. (O.) gracilis JEFFREYS, 1865:37 [synonym: 7. rectangularis SOOT-RYEN, 1941:28-29; pl. 3, figs. 11-14; pl. 7, fig. 3; pl. 10, fig. 11] T. (O.) phaseolina (Lamarck, 1818) [synonym: Tellina pa- pyracea Poli, 1791, non Gmelin, 1791; there are a num- ber of additional synonyms] T. (O.) villostuscula (MACGILLIVRAY, 1827:370-371, 410; pl. 1, figs. 10, 11) [synonym: Anatina intermedia W. CLARK, 1855:141-142]. There appears to be a new species of this subgenus in the western Atlantic, including material reported from Yucatan by DALL (1886:308) as “Thracia phaseolina Kie- ner” (USNM 64062) (see Discussion under 7. (O.) bere- nicede). There is one probable Australian member of this sub- ’ GMELIN (1791) was published prior to 14 May (HOPKINSON, 1908). It is not known when POLI (1791) was published, so it must be dated as 31 Dec. JCZN Art. 21(c)(ii)). Therefore, Tellina papyracea Poli is a junior homonym of 7. papyracea Gmelin, and we must use the next available name for this species, Am- phidesma phaseolina Lamarck, 1818. The Veliger, Vol. 33, No. 1 genus (cited above). Two Asian species appear to belong here as well: T. (O) concinna (REEVE, 1859:pl. 3, fig. 17, ex Gould MS)— Japan & Philippine Islands T. (O.) koyamar (HABE, 1981:187-188; pl. 3, fig. 4)—Japan. Thracia (Odoncineta) squamosa Carpenter, 1856 (Figures 29, 30) Thracia squamosa Carpenter, 1856: CARPENTER, 1856:229-230; CARPENTER, 1857b:287, 300, 366; REEVE, 1859:pl. 3, fig. 16; CARPENTER, 1864b: 619 [1872:105]; CONRAD, 1869:55; DALL, 1915:444; Lamy, 1931:233; KEEN, 1958:230-231; fig. 590; PAL- MER, 1963:320, 393; pl. 63, figs. 16, 17; KEEN, 1971: 295-296; fig. 761; BERNARD, 1983:64. Type material and locality: BM(NH) 1966570, holo- type, partly broken pair; length, 27.9 mm; height, 16.0 mm; convexity (of left valve), 4.1 mm (pair would have been about 8.2 mm) (Figure 29). Mazatlan, Sinaloa, Mex- ico (23°12'N, 104°20'W); C. Shipley. Description: Medium-sized (length to 36 mm; SBMNH 35088; Isla Gibraleon, Archipiélago de las Perlas, Pana- ma); thin; approximately equivalve; somewhat longer, sharply round anteriorly; truncate posteriorly; posterior slope set off by a low ridge; surface with conspicuous pustules, most prominent on posterior slope; hinge plate unbroken under umbones; lithodesma small; periostracum tan; pallial line broad, stopping just short of vertical line from beaks. Distribution and habitat: Bahia Magdalena, Baja Cali- fornia Sur (24°38'N, 112°19’W) (CAS 066623), through- out the Gulf of California, south to Isla Gibraleon, Ar- chipiélago de las Perlas, Panama (8°31’N, 79°3’W) (SBMNH 35088), and Isla Rancheria, Golfo de Chiriqui, Panama (7°37'N, 81°43'W) (Skoglund Coll.). This species has been recorded from the intertidal zone to 61 m (mean, 19 m); there are four records from sand, one from mud. I have examined 32 lots, including the type specimen. I have illustrated here a specimen from the southern Gulf of California (Figure 30). Discussion: Young specimens of 7hracia squamosa in the Panamic province can be distinguished from specimens of T. curta of similar size by their shorter posterior end, less projecting resilifer, and sparser pattern of pustules. (See next species for additional comparative comments.) Thracia (Odoncineta) bereniceae Coan, sp. nov. (Figure 31, 31a) Type material and locality: SBMNH 35089, holotype; length, 17.5 mm; height, 10.1 mm; convexity, 5.0 mm (Figure 31). SBMNH 35090, paratypes, 5 pairs and 1 valve. One of these paratypes will be placed in the USNM, CAS, and ANSP. Bahia Cholla, Sonora, Mexico (31°21'N, 113°37'W); dead on sand bars at low tide; C. Skoglund; 26 Feb. 1967. 30 31a Explanation of Figures 29 to 31 Figures 29, 30. Thracia (Odoncineta) squamosa Carpenter. Figure 29: Holotype; BM(NH) 1966570; length, 27.9 mm. Figure 30: SBMNH 35103; La Paz, Baja Calif. Sur; intertidal zone; length, 21.7 mm. Figure 31. Thracia (Odoncineta) bereniceae Coan, sp. nov. Holotype; SBMNH 35089; length, 17.5 mm. Figure 31a: Close-up of lithodesma, left valve. Page 38 The Veliger, Vol. 33, No. 1 36 37 Explanation of Figures 32 to 37 Figures 32, 33. Asthenothaerus (A.) villosior Carpenter. Figure 32: Holotype; USNM 16292; length, 9.6 mm. Figure 33: SBMNH 35104; Puerto San Carlos, Bahia Magdalena, Baja California Sur; 4 m; length, 11.8 mm. E. V. Coan, 1990 Page 39 SBMNH 35106, paratypes, 3 pairs; Skoglund Coll., paratypes, 3 pairs. Bahia Cholla, Sonora, Mexico; dead on sand bars at low tide; M. Johnson, April 1967. Description: Small (length to 25.0 mm; a paratype), thin; right valve slightly more inflated; anterior end much lon- ger, sharply rounded; posterior end narrowly truncate; surface very finely granular, not punctate; pallial sinus long and narrow, reaching past beaks; hinge with an ex- ternal ligament; hinge plate with a slot for a conspicuous lithodesma that reaches the shell wall of each valve un- derneath the umbones. Distribution and habitat: As yet known from only 7 lots, the following 5 in addition to the type lots: LACM 73-3—Bahia del Coyote, Bahia Concepcion, Baja California Norte; 9-27 m—1 valve LACM 73-122—Isla Blanca, Bahia Concepcion, Baja Cal- ifornia Norte; 11-18 m—3 valves LACM 66-30—La Paz, Baja California Sur; 37-55 m; mud—2 valves SBMNH 35091—Gulf of Tehuantepec, Oaxaca, Mexico; 9-27 m; sand—1 valve SBMNH 35092—Bahia Ballena, Gulf of Nicoya, Costa Rica; 15-21 m—1 pair. Thus, the species is known from Bahia Cholla, Sonora, Mexico (31°921'N, 113°37’W), to Bahia Ballena, Gulf of Nicoya, Costa Rica (9°44'N, 85°W); from the intertidal zone to 46 m (mean, 20 m); both sand and mud bottoms. Discussion: This species differs from Thracia (O.) squa- mosa in having a large lithodesma (Figure 31a), more elongate dimensions, a longer anterior end, a narrower posterior end that is generally more rounded in adults, smoother surface, and more elongate pallial sinus. This new species is similar to an as yet undescribed western Atlantic species known from 9 lots: ANSP 175658 and USNM 83162 from Florida; ANSP 298819, 298928, and 329526 from the Bahamas; USNM 64062 from Yu- catan Strait; and AMNH 191043 and 191075 and ANSP 249402 from the Virgin Islands. It differs from the new species in having stronger concentric sculpture and a more truncate posterior end, and in being more inflated. The species name is in recognition of the help my moth- er, Berenice Coan, has given me over the years in my work in malacology, particularly with proofreading. Asthenothaerus CARPENTER, 1864a:311 Type species: A. villosior Carpenter, 1864; by monotypy— eastern Pacific. Species of Asthenothaerus differ from Thracia in their complete lack of an external ligament. The hinge is thin, and there is a butterfly-shaped lithodesma (Figure 35). (The anatomical discussion and figure of Asthenothaerus —_— in PELSENEER (1911) was probably based on something else—see Introduction herein.) Subgenus (Asthenothaerus) s.s. Shells small; lithodesma small. In addition to the two eastern Pacific species, there is one in the western Atlantic: A. (A.) hemphillit DALL, 1886:308-309; DALL, 1902:510; pl. 31, fig. 9—Florida [synonym: A. bales: REHDER, 1943a: 189; pl. 19, figs. 13, 14]. Two Japanese species have also been placed in this genus and probably belong in this subgenus: A. sematana (YOKOYAMA, 1922:173; pl. 14, figs. 17, 18) A. isaotakii OKUTANI, 1964:83-85; text fig. 6. Asthenothaerus (A.) villosior Carpenter, 1864 (Figures 32, 33) Asthenothaerus villosior Carpenter, 1864: CARPENTER, 1864a:311 [1872:209]; CARPENTER, 1864b: 618 [1872:104]; CONRAD, 1869:56; DALL, 1915:446; I. OLDROYD, 1925:86-87; SCHENCK, 1945:516; pl. 66, figs. 11, 12; KEEN, 1958:230-231; fig. 591; PALMER, 1958: 75-76, 329; pl. 4, figs. 5-9; PALMER, 1963:320-321 [as A. “villosier”]; KEEN, 1971:296-297; fig. 762; BERNARD, 1983:64. Type material and locality: USNM 16292, holotype, broken pair; length, 9.6 mm (may have been closer to 10 mm); height, 6.4 mm; convexity, 3.8 mm (Figure 32). Cabo San Lucas, Baja California Sur (22°52'N, 109°54'W); J. Xantus. Description: Small (to 10 mm; holotype), elongate; right valve somewhat more inflated; anterior end longer, sharply rounded; posterior end produced, truncate; surface with very fine granulations and conspicuous concentric undu- lations that become more evident toward ventral margin; periostracum light tan; pallial sinus elongate, reaching well past beaks. Distribution and habitat: E side of Isla de Cedros, Baja California Norte (28°13'N, 115°9'30”W) (LACM 71-94), into and throughout the Gulf of California, south to Punta Quepos, Puntarenas Prov., Costa Rica (9°22'43’N, 84°9'41"W) (LACM 72-58), from the intertidal zone to 73 m (mean, 19 m); recorded on a variety of substrates, mostly sand and rocks, suggesting that the species lives in the sand matrix among rubble. I have examined 41 lots including the type specimen. Here I illustrate a specimen from Baja California (Fig- ure 33). Discussion: The largest specimens have been obtained from the southern part of the Gulf of California, material Figures 34, 35. Asthenothaerus (A.) diegensis (Dall). Figure 34: Lectotype (herein) of Thracia diegensis,; USNM 73604; length, 8.4 mm. Figure 35: LACM 59890; San Pedro, California; lithodesma; scale bar, 250 um. Figures 36, 37. Asthenothaerus (Skoglundia) colpoica (Dall). Figure 36: Holotype of Thracia colpoica; USNM 73639; length, 17.3 mm. Figure 37: CAS 066627; Guaymas, Sonora, Mexico; closed pair; length, 23.0 mm; open pair, showing lithodesma; length, 23.7 mm. Page 40 from the outer coast of Baja California and from Costa Rica being smaller. Specimens from Costa Rica have heavi- er concentric ribs than material from elsewhere. KEEN (1958) synonymized Asthenothaerus diegensis with this species, but it is distinct, being more inflated and less produced posteriorly. Juvenile specimens of this species are difficult to distin- guish from those of Thracia (Ixartia) curta. Asthenothaerus (A.) diegensis (Dall, 1915) (Figures 34, 35) Thracia diegensis Dall, 1915: DALL, 1915:443; I. OLDROYD, 1925:85; KEEN, 1958: 231 [as a synonym of A. villosior]; KEEN, 1971:297 [as a synonym of A. villosior]; BERNARD, 1983:64 [as a syn- onym of A. villosior]. Asthenothaerus villosior Carpenter, auctt., non Carpenter, 1864: WILLIAMSON, 1905:121. [non CARPENTER, 1864a:311.] Type material and locality: USNM 73604, lectotype (herein); length, 8.4 mm; height, 6.5 mm; convexity, 3.8 mm (Figure 34). USNM 859379, paralectotypes, 4 pairs, 15 valves (most probably forming pairs), plus a few frag- ments. San Diego Bay, San Diego Co., California (32°40'N, 117°10'W); 2-9 m; sandy mud. Description: Small (length to 11.0 mm; Bahia Todos San- tos, Baja California Norte; LACM 64301), oval; right valve more inflated; anterior end longer, inflated, rounded; posterior end slightly produced, truncate; periostracum tan, most evident near ventral margin; surface with very fine granules, most evident near ventral margin; pallial sinus just reaching ventral line from beaks. Distribution and habitat: San Pedro, Los Angeles Co., California (33°44'42"N, 118°11'24”"W) (CAS 066624), to Bahia Magdalena, Baja California Sur (24°38'N, 112°9'W) (USNM 217825; CAS 066625); Bahia de Los Angeles, Baja California Norte (28°55'N, 113°31'W) (CAS 066626; Skoglund Coll.); Soladita Cove, Guaymas, Sonora (27°54'N, 110°58’'W) (LACM 68-27; juveniles, probably this species). There is a single pair labeled “50 m off Newport, Oregon” (about 45°N) (LACM 140426), a locality I doubt because of the lack of any specimens of this shallow-water species from between Oregon and southern California; this may be the result of a transcription error for Newport, Cali- fornia, where the species has been obtained (CAS 018071). Recorded from the intertidal zone to 119 m (mean, 23 m) on mud and sand bottoms. Not uncommon; I have ex- amined 58 lots, including the types. Discussion: This species is closest to the western Atlantic Asthenothaerus (A.) hemphillii Dall, which attains a larger size and is less flattened. (Skoglundia) Coan, subgen. nov. Type species: Thracia colpoica Dall, 1915—eastern Pacific. Extremely thin and easily damaged, and probably as a result extremely rare in collections. It is oval in outline, The Veliger, Vol. 33, No. 1 and the right valve is decidedly more inflated than the left. The ligament is internal, supported by a large, butterfly- shaped lithodesma that abuts each valve under the um- bones. This subgenus differs from Asthenothaerus (Astheno- thaerus) in being much larger and in having a still more conspicuous lithodesma. It differs from Bushia in having very thin shells without concentric sculpture and in pos- sessing a butterfly-shaped lithodesma, not a bar-shaped one. A similar lithodesma is present in “TVhracia” rushi PILSBRY (1897:292-293; pl. 7, fig. 30) from Uruguay and Argentina. This species, which has a small segment of external ligament, has a thicker, more evenly oval shell. It was placed in Asthenothaerus by CARCELLES (1947:3- 4). The new subgenus is named for Carol C. Skoglund of Phoenix, Arizona, who generously contributed material for this and other studies. Asthenothaerus (Skoglundia) colpoica (Dall, 1915) (Figures 36, 37) Thracia colpoica Dall, 1915: DALL, 1915:443-444; KEEN, 1958:230-231; fig. 588; OLSSON, 1961:458, 556; pl. 83, figs. 7, 7a; KEEN, 1971: 295-296; fig. 759; BERNARD, 1983:64. Type material and locality: USNM 73639, holotype, pair; length, 17.3 mm; height, 14.5 mm; convexity, 8.0 mm (Figure 36). “Gulf of California,” here clarified as being Guaymas, Sonora, Mexico (27°55'N, 110°53’W), where the species has been collected (CAS 066627). Description: Small (length to 23.7 mm; CAS 066627; Guaymas, Sonora), very thin-shelled, rounded; right valve larger, more inflated; anterior end much longer, rounded; posterior end truncate; posterior slope set off by a low ridge; escutcheon present, more evident in right valve; sur- face with conspicuous growth lines; no pustules evident; periostracum thin, light tan on posterior slope; pallial sinus broad, short; internal ligament with a large, butterfly- shaped lithodesma. I have illustrated two complete specimens from Guay- mas, one showing the lithodesma (Figure 37). Distribution and habitat: Guaymas, Sonora (27°55'N, 110°53’W) (CAS 066627), and La Paz, Baja California Sur (24°10'N, 110°19’W) (Skoglund Coll.), south to Tum- bez, Tumbez Prov., Peru (3°40’S, 80°23'W) (PRI 25945), on intertidal mudflats. This species is known from only 6 lots, including the type specimen. Bushia DALL, 1886:309-311 Type species: Asthenothaerus (Bushia) elegans Dall, 1886; by monotypy—western Atlantic [see also DALL, 1889:440; pl. 39, fig. 1]. E. V. Coan, 1990 This genus has a small segment of external ligament and a large internal one with a conspicuous, bar-shaped lithodesma seated on thickened cups beneath the beaks (Figure 41). In Bushia s.s., these cups are on the shell wall. In B. (Pseudocyathodonta), they are on a shelf between the shell wall and the hinge plate. Subgenus (Busha) s.s. The four known species of Bushia (Bushia) have con- centric sculpture, prominent in three of them, subdued in the fourth. All occur offshore. DALL (1886) described the anatomy of B. elegans. The type species occurs in the west- ern Atlantic, the other three in the eastern Pacific. Bushia (B.) panamensis (Dall, 1890) (Figure 38) Asthenothaerus (Bushia) (elegans var.?) panamensis Dall, 1890: DALL, 1890:275; DALL, 1915:446 [Bushia as a full ge- nus]; KEEN, 1958:231; fig. 592; KEEN, 1971:296-297; fig. 763; BERNARD, 1983:64. Type material and locality: USNM 87583, holotype, a right valve; length, 13.9 mm; height, 11.2 mm; convexity, 3.6 mm (pair would have been about 7.2 mm) (Figure 38). SW of Isla San José, Archipiélago de las Perlas, Gulf of Panama (7°56'N, 79°41'30”W); 94 m; mud; USFC Sta. 2805; 30 Mar. 1888. Description: Small (length, 13.9 mm), oval, inflated, ap- proximately equilateral; anterior end sharply rounded; posterior end slightly truncate; surface with strong con- centric sculpture; pallial line reaching just short of midline. Distribution and habitat: Known only from the type spec- imen. Discussion: In describing the type species of Bushia as well as this species, Dall regarded Bushia as being a subgenus of Asthenothaerus, but his headings cite the species as if Bushia were a full genus. Because his intent is clear, pa- rentheses must be placed around Dall’s name when Bushia is used as a full genus. Of the three eastern Pacific and one western Atlantic species of Bushia, this one differs in being more inflated, proportionately higher for its length, and in having more central beaks. Bushia (B.) galapagana (Dall, 1915) (Figure 39) Cyathodonta galapagana Dall, 1915 DALL, 1915:446. Type material and locality: USNM 195029, holotype, a left valve; length, 25.9 mm; height, 16.9 mm; convexity, 5.0 mm (Figure 39). Off Isla Gardner, Galapagos Islands, Ecuador (1921'S; 89°40'15”W); 73 m; sand; USCF Sta. 2813; 7 Apr. 1888. Page 41 Description: Medium-sized (length to 32.8 mm; SBMNH 35093; Isla del Coco, Costa Rica), elongate; anterior end longer, sharply rounded; posterior end truncate; externally with conspicuous concentric sculpture (somewhat subdued on central ventral margin of holotype, but not in material from Isla del Coco); pallial sinus just reaching vertical line from beaks. Distribution and habitat: Known from just four stations: three in Bahia Chatham, Isla del Coco, Costa Rica (5°33'N, 87°2'30”W) (LACM 38-39; SBMNH 35093, 35094), and the type specimen from the Galapagos Islands (1°21’S, 89°40'15"W) (USNM 195029); 57-83 m (mean, 71 m). Discussion: This species has been overlooked by many workers and is not mentioned by KEEN (1958, 1971), OLSSON (1961), or BERNARD (1983). Dall, who had described the genus Bushia and both of its then-known species, did not recognize that this species was another Bushia. Instead, he placed it in Cyathodonta, which has a very different ligament, with a resilifer on the hinge plate and no deeply seated lithodesma. Of the eastern Pacific species of Bushia, this is closest to the type species, B. elegans from the western Atlantic, differing in being much larger and in having heavier, more widely spaced concentric ribs. Busha (B.) phillips Coan, sp. nov. (Figures 40, 41) Type material and locality: SBMNH 35095, holotype, right valve; length, 23.0 mm; height, 16.4 mm; convexity, 4.5 mm (Figure 40). SBMNH 35096, paratypes, one smaller right valve and two still smaller pairs. The lith- odesma of one of these pairs is illustrated here (Figure 41). N end Isla Smith, Gulf of California, Baja California Norte (29°6’N, 113°31'W); 183 m; C. & P. Skoglund; Nov. 1981. Skoglund Coll., paratypes, 2 left valves. W end of Isla Smith, Baja California Norte (29°4'N, 113°31’W); 114- 152 m; C. & P. Skoglund; Apr. 1988. Description: Small (to 23 mm; holotype), thin-shelled; right valve more inflated; anterior end longer, sharply rounded; posterior end truncate; sculpture of very subdued concentric ribs; pallial sinus broad, short; lithodesma bar- shaped, seated in thickened cups on shell wall under um- bones. Differentiation: Bushia phillips differs from the other species in the subgenus in its subdued concentric ribs and its thin shell; each of the other species has a heavier shell and more conspicuous concentric sculpture. It is less elon- gate than B. galapagana, but more so than B. panamensis. It is larger than any specimens thus far obtained of either B. panamensis or B. elegans. Distribution and habitat: Known only from four stations on the western side of the Gulf of California, from Isla Page 42 The Veliger; Vola335 Nosal 38a ; 39a E. V. Coan, 1990 Smith (29°6'N, 113°31'W) (type lots), to off Isla Danzante (25°48'N, 111°16’W) (SBMNH 35097; Skoglund Coll.); 38-183 m (mean, 104 m); no bottom types recorded. In addition to the type lots, this species is represented in collections by the following material: SBMNH 35097—off Isla Danzante, 61 m—2 valves Skoglund Coll.—off Isla Danzante, 31-46 m—4 valves. Discussion: This species is named for David W. Phillips of Davis, California, editor-in-chief of The Veliger. Bushia (Pseudocyathodonta) Coan, subgen. nov. Type species: B. (P.) draperi Coan, sp. nov.—eastern Pa- cific. The shells of this new subgenus are shaped like those of Cyathodonta and are of similar thickness, but they have a hinge similar to that in Bushia. There is a somewhat projecting resilifer for the external portion of the ligament, but it is much smaller than that in Cyathodonta and lacks a thickened calcareous pad. A subumbonal cup on a shelf, well below the hinge plate, undoubtedly holds a lithodes- ma, which is lacking in the only known specimen. In Bushia s.s. the cup is on the shell wall. There is heavy concentric sculpture, but it is not oblique, as in Cyathodonta); it is more undulating than that in Bushia s.s. This genus is known only from the type species. Bushia (Pseudocyathodonta) draperi Coan, sp. nov. (Figure 42, 42a) Type material and locality: SBMNH 35098, holotype, pair; length, 28.5 mm; height, 22.4 mm; convexity, 12.2 mm [broken] (Figure 42). In the Gulf of California, off Isla Danzante, Baja California Sur (25°48'N, 111°16’W); 61 m; C. & P. Skoglund; either Oct. 1979 or Oct. 1983. Description: Small (to 28.5 mm in length; holotype), thin; right valve larger, more inflated; approximately equilat- eral; anterior end sharply rounded; posterior end set off by a low ridge, truncate; escutcheon present, most evident in left valve; surface with conspicuous concentric sculpture, which becomes finer toward posterior end; posterior end with fine pustules; hinge plate narrow, with a subumbonal _— Page 43 cup on a shelf beneath the beaks that presumably holds a lithodesma. Distribution and habitat: Known only from the holotype. Discussion: This species is named for Bertram C. Draper of Los Angeles, California, who has helped with the pho- tographic work for many papers by various authors. Lampeia MacGinitie, 1959 Type species: Thracia (Lampeia) adamsi MacGinitie, 1959, by original designation—Arctic coast of Alaska. A narrow segment of external ligament is present along the dorsal margin, but the main ligament is internal, at- tached to an oblique structure on the shell wall under the beaks. This structure is free along its anteroventral margin, where it is supported by a series of pillars. The internal ligament is supported by a strong, curved lithodesma. The outside of the shell is covered by a heavy brown perios- tracum. The hinge of this genus is closest to Asthenothaerus, but its shell is much heavier than that of this genus; it has a thick, dark periostracum; and there is a simple, curved lithodesma. The buttressed subumbonal structure is like nothing else. This genus is represented only by the follow- ing species. Lampeia adams: (MacGinitie, 1959) (Figures 43, 44) Thracia (Lampeia) adams: MacGinitie, 1959: MAcGINITIE, 1959:163-164; pl. 18, fig. 9; pl. 21, figs. 7, 8; pl. 24, fig. 8. Type material and locality: USNM 610301, holotype, pair; length, 22.8 mm; height, 18.3 mm; convexity, 9.7 mm (Figure 43). 4 km off Point Barrow, Arctic coast of Alaska (about 71°31'N, 156°23'W); 33.5 m, mud-gravel- stone bottom; G. E. MacGinitie, 15 Sept. 1948. Description: Small (to 29.7 mm; UAM 4473; NW Bering Sea); shells average in thickness; right valve larger, more inflated; anterior end slightly longer, rounded; posterior end truncate, with an escutcheon that is evident in both valves; lunule present in left valve; surface of adults with Explanation of Figures 38 to 42 Figure 38. Bushia (B.) panamensis (Dall). Holotype of Asthenothaerus (B.) panamensis,; USNM 87583; length, 13.9 mm. Figure 38a: Close-up of subumbonal cup for lithodesma. Figure 39. Bushia (B.) galapagana (Dall). Holotype of Cyathodonta galapagana; USNM 195029; length, 25.9 mm. Figure 39a: Close-up of subumbonal cup for lithodesma. Figures 40, 41. Bushia (B.) phillipst Coan, sp. nov. Figure 40: Holotype; SBMNH 35095; length, 23.0 mm. Figure 41: Paratype; SBMNH 35096; close-up showing lithodesma under beaks. Figure 42. Bushia (Pseudocyathodonta) draperi Coan, subgen. et sp. nov. Holotype; SBMNH 35098; length, 28.5 mm. Figure 42a: Close-up of hinge of right valve showing resilifer and subumbonal cup for lithodesma. Page 44 The Veliger, Vol. 33, No. 1 Explanation of Figures 43 to 47 Figures 43, 44. Lampeia adamsi (MacGinitie). Figure 43: Holotype of Thracia (L.) adamsi; USNM 610301; length, 22.8 mm. Figure 43a: Close-up of right valve showing subumbonal slot for lithodesma. Figure 44: CAS 066628; off Point Barrow, Alaska; 39 m; length, 11.4 mm; close-up of a broken pair showing lithodesma. E. V. Coan, 1990 Page 45 heavy, dark periostracum and concentric growth lines; pal- lial sinus short, broad. Distribution and habitat: On the Arctic coast of Alaska, from off Point Barrow (71°34’N, 156°22'W) (CAS 066628), westward into the NW Bering Sea off Mys Chaplino, Chukotskiy Poluostrov (64°18'30"N, 171°8'W) (UAM 4473); 10-41 m (mean, 28 m). Sediment type is recorded only for the type specimen: mud-gravel-stone bottom. I have examined 8 lots, including the type. Cyathodonta CONRAD, 1849a:155-156 Type species: C. undulata Conrad, 1849a; by monotypy— eastern Pacific. Shells with conspicuous, oblique, undulating sculpture; hinge with an external ligament and a projecting resilifer; resilifer thickened with a calcareous pad; a small, curved lithodesma present on anterior surface of resilium. There are at least four other species in addition to four in the eastern Pacific: C. cruziana DALL, 1915:446—western Atlantic C. granulosa (A. ADAMS & REEVE, 1850:82, 87; pl. 23, fig. 16)— Japan C. plicata (DESHAYES, 1832:1039-1040)—west Africa C. rugosa (LAMARCK, 1818:464)—western Atlantic [syn- onyms: Thracia magnifica JONAS, 1850:170; pl. 4, fig. 7; T. semirugosa REEVE, 1859:pl. 2 (nomen nudum); T. plicata (Deshayes), auctt., non Deshayes, 1832; 7. (C.) dalli MANSFIELD, 1929:7, 10; pl. 4, figs. 1, 2; T. dissimilis Guppy, 1875:52; C. rectangulata MaAcsoTAY, 1968:87- 88, 410; pl. 4, figs. 1, 2—see Discussion under C. un- dulata). Cyathodonta undulata Conrad, 1849 (Figures 45-47) Cyathodonta undulata Conrad, 1849: CONRAD, 1849a:155-156; CONRAD, 1849b:230; CaAR- PENTER, 1864b:633 [1872:119]; CONRAD, 1869:53; DALL, 1915:444; GRANT & GALE, 1931:259 [in part; not figs. ]; Lamy, 1931:285-286; HERTLEIN & STRONG, 1946:96; HERTLEIN & STRONG, 1955:181; pl. 3, figs. 1, 2; KEEN, 1958:232-233; fig. 595; OLSSON, 1961:459; KEEN, 1971: 297, 299; fig. 766; BERNARD, 1983:64. Cyathodonta granulosa (Adams & Reeve), auctt., non Adams & Reeve, 1850: GOULD, 1853:407 [species’ author not given; nomen nu- dum]; CARPENTER, 1857b:231 [as “C. granulosa Gould’; nomen nudum]. [non Thracia granulosa A. ADAMS & REEVE, 1850:82, 87; pl. 23, fig. 16.] Thracia plicata Deshayes, auctt., non Deshayes, 1832: CARPENTER, 1857b:231, 297, 352; REEVE, 1859:pl. 2, fig. 7b, c [not 7a] [according to LAMy, 1931:285]; CAR- PENTER, 1864b:541, 564, 619 [1872:27, 50, 105]; CONRAD, 1869:53; STEARNS, 1894:157; Lamy, 1909:253. [non DESHAYES, 1832:1039.] Thracia magnifica Jonas, auctt., non Jonas, 1850: MABILLE, 1895:76. [non JONAS, 1850:170; pl. 4, fig. 7.] Cyathodonta lucasana Dall, 1915: DALL, 1915:445; HERTLEIN & STRONG, 1946:96 [in part; not figs.]; KEEN, 1958:232 [in part; not figs.]; KEEN, 1971:297 [in part; not figs.]; BERNARD, 1983:64. [but not C. lucasana Dall, auctt., =C. dubiosa—which see. ] Cyathodonta dubiosa Dall, auctt., non Dall, 1915: DuruHaAM, 1950:70, 161; pl. 16, figs. 2, 7. [non DALL, 1915:445.] Cyathodonta undulata peruviana Olsson, 1961: OLSSON, 1961:459, 556; pl. 83, fig. 2-2b; KEEN, 1971: 297 [as a synonym of C. undulata]; BERNARD, 1983:64 [as a synonym of C. undulata]. Type material and localities: C. undulata—ANSP 55406, lectotype (herein), pair; length, 48.5 mm; height, 35.0 mm; convexity, 15.3 mm (Figure 45). ANSP 372699, para- lectotype, a smaller pair. “Coasts of Lower California and Peru”; restricted to the east coast of Baja California by HERTLEIN & STRONG (1946:96); here further clarified as being La Paz, Baja California Sur (24°12'N, 110°22'W), where the species has been taken (for example, LACM 60-7). C. lucasana—USNM 15910b, holotype, left valve; length 7.8 mm; height, 5.3 mm; convexity, 1.3 mm (Figure 46). Cabo San Lucas, Baja California Sur (22°52'N, 109°54'W); J. Xantus. C. undulata peruviana—ANSP 218953, holotype, right valve; length, 49.2 mm; height, 39.0 mm; convexity, 10.9 mm (Figure 47). Puerto Pizarro [Tumbez], Tumbez Prov., Peru (3°29'S, 80°23'W); A. A. Olsson, 1958. Description: Medium-sized (length to 50.2 mm; SBMNH 35099; Puertecitos, Baja California Norte), oval; right valve much more inflated; anterior end longer, rounded; poste- rior end set off, truncate; surface with oblique concentric undulations, strongest on anterior end, and pustules that generally form a radial pattern (pattern most evident on central part of valves); posterior end more densely pus- tulose; periostracum tan, evident only on posterior slope; pallial sinus moderate in length, reaching almost to vertical line from beaks. Distribution: Bahia Magdalena, Baja California Sur (24°38'N, 112°9'W) (LACM 140427), throughout the Gulf of California, south to Punta Organos, Peru (4°8’S, 81°7’W) (CAS 066629); Isla San Cristobal, Galapagos Islands (LACM 38-188). Most lots are beach material; live-col- lected material has been obtained from the intertidal zone to 64 m (mean, 14m). The bottom type most often recorded is sand. I have examined 109 lots, including the type ma- terial. This species has been reported in Pliocene strata of the Imperial Formation in southern California (HANNA, 1926: 466; POWELL, 1988:16) and of Isla Carmen, Baja Cali- Figures 45-47. Cyathodonta undulata Conrad. Figure 45: Lectotype (herein); ANSP 55406; length, 48.5 mm. Figure 46: Holotype of C. lucasana Dall; USNM 15910b; length, 7.8 mm. Figure 47: Holotype of C. undulata peruviana Olsson; ANSP 218953; length, 49.2 mm. Page 46 fornia Sur (EMERSON & HERTLEIN, 1964:342, 349). It has also been recorded in formations of Pleistocene age at Bahia Magdalena, Baja California Sur (JORDAN, 1936:112, 123), and on the Burica Peninsula, Panama (OLSSON, 1942:162). HOFFSTETTER (1952:45) reported it as a “subfossil” on the Santa Elena Peninsula, Ecuador. Records in Pleisto- cene formations in southern California need to be reex- amined (see Discussion under Cyathodonta pedroana). Discussion: There is a similar C'yathodonta in the western Atlantic, and Iam unable to differentiate C. undulata from some specimens of this rare species. The earliest name for the Atlantic species appears to be Anatina rugosa LAMARCK, 1818 (p. 464), described from Santo Domingo [Hispan- iola]. The holotype, a right valve measuring 41.7 mm in length, is in the Muséum d’Histoire Naturelle, Geneva (No. 1082/36) (Figure 48). I have seen only a few spec- imens of this species, and these not simultaneously, and it is possible that more than one taxon is involved. Large, intact specimens were discussed and figured by J. GIBSON-SMITH & W. GIBSON-SMITH (1983:181; figs. 11- 13). Caribbean material attains a larger size (73 mm) than C. undulata, and these authors maintain that it is more produced anteriorly than is C. undulata. However, the shape of the anterior end is variable in C. undulata, and this may prove to be the case with the Caribbean species. Cyathodonta lucasana is based on a broken, juvenile Cy- athodonta. Although no other available material is this small, it seems to match C. undulata most closely. The name C’. lucasana has been misapplied to specimens of C. dubwosa Dall. Cyathodonta undulata peruviana falls within the range of variability of this species, and oval specimens matching its type have also been obtained in the Gulf of California. Cyathodonta dubiosa Dall, 1915 (Figure 49) Cyathodonta dubiosa Dall, 1915: DALL, 1915:445; I. OLDROYD, 1925:86; pl. 9, fig. 5; HERTLEIN & STRONG, 1946:96; KEEN, 1958:232-233; fig. 593; KEEN, 1971:296-297; fig. 764; BERNARD, 1983: 64. Cyathodonta lucasana Dall, auctt., non Dall, 1915: HERTLEIN & STRONG, 1946:96, 120; pl. 1, figs. 4, 9; KEEN, 1958:232-233; fig. 594; KEEN, 1971:296-297; fig. 765. [non DALL, 1915:445.] [non C. dubiosa Dall, auctt., =C. undulata or C. pedroana— see under these species. ] Type material and locality: C. dubiosa—USNM 96450, holotype, right valve; length, 38.1 mm; height, 27.8 mm; convexity, 8.0 mm (Figure 49). Off La Paz, Baja Cali- fornia Sur (24°18'N, 110°22’W); 48 m; sand; USCF Sta. 2823, 30 Apr. 1888. Description: Medium-sized (length to 40.2 mm; Skoglund Coll.; off Tetas de Cabra, Sonora, Mexico), oval to elon- gate-oval; right valve somewhat more inflated; equilateral, The Veliger, Vol. 33, No. 1 or longer either posteriorly or anteriorly, these variations present within a single lot; concentric undulations gener- ally lower, more numerous and less oblique than those in Cyathodonta undulata; punctations denser than in the pre- ceding species, generally arranged in concentric rows; peri- ostracum tan; pallial sinus very shallow. Distribution and habitat: In Mexico, from Isla Smith, Baja California Norte (29°3'N, 113°30'W) (SBMNH 35100; Skoglund Coll.), and Punta San Antonio, Sonora (27°57'N, 111°7'W) (SBMNH 35101), to Salina Cruz (16°9'N, 95°12'W) (Skoglund Coll.) and Puerto Huatulco (15°44'30"N, 96°8’W) (CAS 066630), Oaxaca; 13 to 183 m (mean, 96 m). The only bottom type recorded, this on but one lot, is sand. I have examined just 11 lots, including the type. Records of this species from California (DALL, 1915; I. OLDROYD, 1925) are based on misidentifications of Cy- athodonta pedroana. This species has been recorded from Pleistocene strata at Bahia Magdalena, Baja California Sur (JORDAN, 1936: 112) and on the Burica Peninsula, Panama (OLSSON, 1942: 162). Discussion: In northern Mexico, this species occurs with Cyathodonta undulata, from which it can be separated by its more inflated left valve; denser punctations, which are arrayed in a concentric pattern; its finer, less oblique con- centric undulations; its shallow pallial sinus; and its off- shore habitat. This rare species seems more closely related to a number of fossil taxa than does Cyathodonta undulata. These in- clude: C. gatunensis (TOULA, 1909:757-758; fig. 15), from Miocene formations in Central America, which appears to have heavier, less oblique ribs (WOODRING, 1982:722; pl. 121, fig. 7); “Cyathodonta?” dolicha WOODRING, 1982 (pp. 721-722; pl. 91, fig. 22; ?pl. 121, fig. 8), from the middle Miocene Gatun Formation of Panama; and C. tristant (OLSSON, 1922:383; pl. 20, fig. 3) from the middle Miocene of Costa Rica. These three taxa may represent the same species. Also related may be C. guadalupensis DALL, 1903 (p. 1527; pl. 53, fig. 6) and C. spencer: DALL, 1903 (pp. 1527-1528; pl. 53, fig. 8), from a Miocene formation on Guadeloupe (according to WOODRING (1982: 722), these two names probably refer to the same species); and C. reedsi Maury, 1920 (pp. 25-26; pl. 5, fig. 2), from a Miocene formation in Puerto Rico. In the Recent fauna, Cyathodonta dubiosa is closest to the western Atlantic C. cruziana Dall, 1915, which differs in being still more densely pustulose. Cyathodonta pedroana Dall, 1915 (Figure 50) Cyathodonta pedroana Dall, 1915: DALL, 1915:445; I. OLDROYD, 1925:86; pl. 54, figs. 1- 3; BERNARD, 1983:64 [as a synonym of C. dubiosa Dall]. Cyathodonta dubiosa Dall, auctt., non Dall, 1915: DALL, 1915:445 [in part; not type specimen]. 50 Explanation of Figures 48 to 51 Figure 48. Cyathodonta rugosa (Lamarck). Holotype of Anatina rugosa; MHNG 1082/36; length, 41.7 mm. Figure 49. Cyathodonta dubiosa Dall. Holotype; USNM 96450; length, 38.1 mm. Figure 50. Cyathodonta pedroana Dall. Lectotype (herein); USNM 207527; length, 26.0 mm. Figure 51. Cyathodonta tumbeziana Dall. Holotype; ANSP 218952; length, 37.6 mm. Page 48 Cyathodona undulata Conrad, auctt., non Conrad, 1849 GRANT & GALE, 1931:259, 906; pl. 13, fig. 6a, b [in part]. [non CONRAD, 1849a:155-156.] Type material and locality: USNM 207527, lectotype (herein), pair; length, 26.0 mm; height, 19.8 mm; convex- ity, 10.8 mm (Figure 50). USNM 859377, paralectotypes, 5 pairs; SBMNH 34284, paralectotype, 1 pair. San Pedro Harbor, Los Angeles Co., California (33°43'N, 118°15'W), mud; Eschnaur. Description: Medium-sized (length to 38 mm; LACM 16956; Newport Bay, Orange Co., California), thin; right valve more inflated; left valve less flattened than that in Cyathodonta undulata; anterior end longer, rounded; pos- terior end truncate; concentric undulations, on average, intermediate between those of C. undulata and C. dubiosa (lower, more numerous, and less oblique than those of C. undulata; more prominent and oblique than those in C. dubiosa); pustules arrayed in a concentric pattern; peri- ostracum dark brown; pallial sinus shallow, but deeper than that in C. dubiosa. Distribution and habitat: Monterey Bay, Monterey Co., California (36°37'N, 121°52'30”W) (LACM 60-22; UCMP 2395), to Bahia Magdalena, Baja California Sur (24°58'15"N, 115°53’W) (USNM 212572), from 9 to 114 m (mean, 36 m). A wide variety of bottom types are re- corded, including shale, rocks, sand, and mud. I have ex- amined 58 lots, including the type specimens. Records of living Cyathodonta undulata and C. dubiosa from southern California are undoubtedly based on this species. Material from Pliocene and Pleistocene strata in south- ern California and northern Baja California must be reex- amined in light of the differentiating characters discussed here. Most of the following records may have been based on this species, though it is possible that Cyathodonta un- dulata also occurred here in the late Pleistocene: Pliocene—DURHAM & YERKES (1964:27), as C. cf. C. un- dulata. Early Pleistocene—T. OLDROYD (1925:4), as C. pedroana; A. CLARK (1931:opp. p. 30), as C. cf. pedroana; and DELONG (1941:opp. p. 244), as Thracia undulata. Late Pleistocene —WILLETT (1937:387), as Thracia (C.) un- dulata; and KANAKOFF & EMERSON (1959:22), as C. undulata. Undifferentiated Pleistocene—ORCUTT (1921:19), as C. du- biosa. Discussion: DALL (1915:446) described Cyathodonta cru- ziana from “Santa Cruz Island” in the West Indies, in- dicating that it is “analogous” to C. pedroana, but it seems closer to C. dubiosa (see under same). Cyathodonta tumbeziana Olsson, 1961 (Figure 51) Cyathodonta tumbeziana Olsson, 1961: OLSSON, 1961:460, 556; pl. 83, fig. 1, 1a [on pl. expl. ithe Veliger; Volk33, Nom as “C. tumbezensis’’; first revision herein]; KEEN, 1971: 297 [as a possible synonym of C. undulata]; BERNARD, 1983:64 [as a synonym of C. undulata). Type material and locality: ANSP 218952, holotype, right valve; length, 37.6 mm; height, 32.2 mm; convexity, 7.9 mm (pair would have been about 14 mm) (Figure 51). Tumbez, Tumbez Prov., Peru (3°29'S, 80°23’W); A. A. Olsson, 1958. Description: Medium-sized (length to 37.6 mm; holo- type), oval; right valve decidedly more inflated; approxi- mately equilateral; anterior end rounded; posterior end truncate, with a very narrow posterior slope; ventral mar- gin produced posteroventrally; concentric undulations ob- scure, overlain by fine beaded threads; posterior slope with conspicuous pustules, less dense than in other species; peri- ostracum dark tan; pallial sinus shallow. Distribution and habitat: Golfo de Tehuantepec off Puer- to Madero, Chiapas, Mexico (14°42'-52'N, 92°32’-42'W) (SBMNH 35102), to Mancora, Tumbez Prov., Peru (4°6’S, 81°4'W) (OLSSON, 1961; specimen not examined, but as- sumed to be correctly identified), 13-26 m (mean, 19 m). This species is known from only 7 lots, of which I have examined 5, including the type specimen. Discussion: This is the most distinctive eastern Pacific species of Cyathodonta, and it can be distinguished by its oval outline, produced posteroventral margin, and its nar- row posterior slope. EXCLUDED TAXA (1) Tyleria fragilis H. ADAMS & A. ADAMS, 1856 (p. 368; pl. 97, fig. 3, 3a; new genus and species), was tentatively placed in the Thraciidae by KEEN (1958:232-233; fig. 596). Later, it was discovered that the type specimen was actually a Sphenia (KEEN, 1971:263; see also BERNARD, 1983:58). (2) Thracia carnea Morch, 1860 (p. 180), proved to be a Tellina (KEEN, 1966b:13, 14; fig. 14a, b). ACKNOWLEDGMENTS There is no way that a monograph of this sort could be completed without the cooperation and advice of a great many individuals, including those connected with mu- seums as well as independent collectors. I deeply appreciate the help of the following persons: Academy of Natural Sciences of Philadelphia—Arthur Bo- gan and Mary A. Garback; Australian Museum—Ian Loch and Winston Ponder; British Museum (Natural Histo- ry)—Solene Morris; California Academy of Sciences— Terrence Gosliner, Michael G. Kellogg, Elizabeth Kools, and Robert Van Syoc; Muséum d’Histoire Naturelle, Ge- neva—Yves Finet; Museum of Comparative Zoology, Harvard University—Kenneth J. Boss, Alan R. Kabat, and Silvard Kool; National Museum of New Zealand— E. V. Coan, 1990 Page 49 Bruce A. Marshall; National Science Museum, Tokyo— Akihiko Matsukuma; Natural History Museum of Los Angeles County—Clifton Coney, George Kennedy, James H. McLean, LouElla Saul, and Gale Sphon; Naturhis- torisches Museum, Vienna—Erhard Wawra; New Zea- land Geological Survey—Phil A. Maxwell; Paleontology Museum, University of California, Berkeley—David R. Lindberg; Santa Barbara Museum of Natural History— Paul Scott; Seattle Aquarium—Roland Anderson; U.S. Geological Survey, Menlo Park—Louie Marincovich and Charles L. Powell, II]; U.S. National Museum of Natural History—Frederick J. Collier, Diane Tyler, and Tom Waller; University of Alaska Museum—Nora R. Foster; University of Colorado Museum—Shi-Kuei Wu; Univer- sity of Hong Kong—Brian Morton; Tromse Museum— Wim Vader; Zoologisk Museum, Copenhagen—Tom Schigette; Zoologisk Museum, Oslo—Karin Andersen. I appreciated the opportunity to examine material from the private collections of Carol C. Skoglund of Phoenix, Arizona; the late Frank R. Bernard of Nanaimo, British Columbia; and Rae Baxter of Homer, Alaska. I appre- ciated the advice of Riccardo Giannuzzi-Savelli of Paler- mo, Italy; Kevin Lamprell of Kallangur, Queensland, Aus- tralia; Richard E. Petit of North Myrtle Beach, South Carolina; and Barry Roth of San Francisco, California. The photographs of specimens were made by Bertram C. Draper; the SEM photographs were prepared with the assistance of Terrence Gosliner and Mary Ann Tenorio. The plates were prepared with the assistance of Sharon Williams. Useful comments on the manuscript were made by Paul Scott and Berenice Coan. LITERATURE CITED All works cited in the text, including sources of taxonomic units, are listed. Numbers of volumes, bulletins, mono- graphs, memoirs, professional papers, and special papers are in bold face; series numbers, in parentheses, precede volume numbers; issue numbers, in parentheses, follow volume numbers; supplementary information, such as sec- ondary methods of listing volumes, part numbers, and parenthetical statements, is given in brackets. Plates are listed, but not text figures, maps, charts, or tables. Exact dates of publication are given when possible. ADAMS, A. & L. A. REEVE. 1848-1850. Mollusca. Pp. x + 87; 24 pls. In: A. Adams (ed.), “The zoology of the voyage of H.M.S. Samarang, under the command of Captain Sir Ed- ward Belcher, .. . during the years 1843-1846.” Reeve, Ben- ham, & Reeve: London. 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Atlas Molliuskov Verkhnego Miotsena i Pliotsena Sakhalina. Akad. Nauk SSSR, Sibirskoe Otdele- nie, Sakhalinskit Kompleksnyi, Nauchno-Issledovatel’skii Institute. 179 pp.; 50 pls. THE VELIGER © CMS, Inc., 1990 The Veliger 33(1):56-102 (January 2, 1990) Laubierinidae and Pisanianurinae (Ranellidae), Two New Deep-Sea Taxa of the Tonnoidea (Gastropoda: Prosobranchia) by ANDERS WAREN Naturhistoriska Riksmuseet, Box 50007, S-10405 Stockholm, Sweden AND PHILIPPE BOUCHET Muséum National d’Histoire Naturelle, 55 Rue de Buffon, F-75005 Paris, France Abstract. The classification of the Tonnoidea is discussed based on new information about deep-sea species. Representative radulae, opercula, and larval shells are described and figured. The conclusions agree mainly with earlier classification, with the following exceptions: Oocorythinae is moved from Tonnidae to Cassidae and its value as a subfamily is questioned. The gross anatomies of two Recent deep-water species of Pisanianura Rovereto, 1899, are described, and a new ranellid subfamily, Pi- sanianurinae, is described for P:sanianura Rovereto, 1899, formerly classified in the Buccinidae. The genera Laminilabrum Kuroda & Habe, 1961, presently in the Trichotropidae, Kaiparanura Laws, 1944, and Nawenia Ladd, 1977, presently in the Buccinidae, are considered synonyms of Prsanianura, which is known in the fossil record since the Oligocene. A new family, Laubierinidae, is erected for Laubierina gen. nov. and Akibumia Kuroda & Habe, 1958 (formerly Trichotropidae) with three Recent deep-water species. Laubierina peregrinator gen. et sp. nov. is described from deep water in the tropical Atlantic and Indian oceans. Two large (5 mm) planktonic larvae belonging to the Laubierinidae are described and one of them is remarkable for being a sexually mature male at the time of settlement. All dissected adults are females and it is speculated that Lawbierina is a protandrous hermaphrodite with neotenic males. The gross anatomies of L. peregrinator sp. nov., A. orientalis (Schepman, 1909), and A. schepmani (Habe, 1962) are described. Akibumia reticulata Habe, 1962, is referred to the Epitoniidae and Conradia minuta Golikov & Starobogatov, 1986 (described in Fossaridae) is considered a larva of Neptunellinae. Thalassocyon bonus Barnard, 1960, and 7. tui Dell, 1967, are synonymized; their anatomies are briefly described and compared with that of Ficus and it is concluded that Thalassocyon has been correctly referred to the Ficidae. Attention is drawn to the fact that the morphology of the Ficidae conforms poorly with other Tonnoidea. The value and use of larval shells as taxonomical criteria are discussed, and it is concluded that they are useful criteria, as long as clear distinction is made between “primary” (z.e., planktotrophic) and “secondary” (7.e., non-planktotrophic) types of larval shells and only “‘primary” ones are compared. Indo-Pacific area. During these expeditions, material of deep-water species of the Tonnoidea has been obtained. During the last two decades, several French expeditions Such specimens are rare and each expedition has usually have been exploring the bathyal and abyssal parts of the brought back only one or two samples. Among them were INTRODUCTION A. Warén & P. Bouchet, 1990 some specimens that could not be classified in the normally accepted suprageneric taxa. During the same time, much has been added to the knowledge of the tonnoids, mainly through the work of Dr. Alan Beu, Geological Survey of New Zealand (see Literature Cited), working on the recent and fossil ton- noids. In order to give a more complete picture of the superfamily, we have felt that it would be valuable to review also the deep-sea groups, which otherwise, through their deviate morphology, might easily be overlooked. We have also studied the consequences that the new taxa may have on existing classification by comparing them with presently accepted groups, and we have supplemented existing information with new data about larval shell and radular morphology of the known groups. We will start by giving a key to the families of Tonnoidea partly based on BEU’s (1981, 1988b) characters of the families, supplemented by our own observations. Then we continue with a review of the families of the Tonnoidea, supplemented by new information and illustrations of fea- tures used in the discussions of the new taxa. We finish with a discussion about the larval shell of the Tonnoidea and its value as a taxonomic criterion. Author names and dates not found in the text are given in the Appendix. MATERIAL Deep-water Tonnoidea are rare and the material studied here has been accumulated over a period of more than 20 years. The following expeditions contributed material that was essential to our understanding of the deep-water Ton- noidea. ““Benthedi,” 1977, on R/V Surozt, in the northern part of the Mozambique Channel, directed by B. Tho- massin, collected the first lot of Laubierinidae, which puzzled us for several years. “Walvis,” 1981, on R/V Jean Charcot, off southwestern Africa, directed by M. Sibuet, collected additional wet material of Laubierinidae. “Bio- cal,” 1985, on R/V Jean Charcot, off southern New Cal- edonia, directed by C. Levi, brought back living represen- tatives of the genus Pisanianura. Other expeditions yielded additional information and are cited when appropriate in the text. In addition, described and undescribed material from AMS, ANSP, MOM, NMNZ, SAM, SMNH, USNM, ZMA, and ZMC was examined. When necessary, type material has been examined, and is referred to in the text or accompanying figures. The author and date for the taxa discussed are given at appropriate places in the text, except for the species listed in the Appendix. ABBREVIATIONS USED AMS, Australian Museum, Sydney; ANSP, Academy of Natural Sciences, Philadelphia, MNHN, Museum Na- tional d'Histoire Naturelle, Paris; MOM, Musée Océan- Page 57 ographique, Monaco; NMNZ, National Museum of New Zealand, Wellington; NZGS, New Zealand Geological Survey, Lower Hutt; NZOI, New Zealand Oceanographic Institute; SAM, South African Museum, Cape Town; SMNH, Naturhistoriska Riksmuseet, Stockholm; USNM, U.S. National Museum of Natural History, Washington, D.C.; ZMA, Zodlogisch Museum, Amsterdam; ZMC, Zoologisk Museum, Kgbenhayn. SYSTEMATICS Superfamily TONNOIDEA Suter, 1913 The superfamily Tonnoidea has been defined by THIELE (1929) and Boss (1982). The only change in the criteria of this taxon that our investigations have necessitated is that the osphradium, which has been supposed to be bi- pectinate in all species, is monopectinate in the Laubi- erinidae. If future work shows that the Ficidae does not belong to the Tonnoidea (cf. p. 83), it should also be added that all Tonnoidea have large, complex salivary glands (small and simple in Ficidae). Morphology of the Tonnoid Veliger Veliger larvae belonging to species of Tonnoidea have been known since the mid-19th century oceanographic expeditions (FORBES, 1852; H. & A. ADAMS, 1853-1858), and these large larvae were then frequently described as distinct holoplanktonic species. The following generic names were thus based on tonnoid larvae: Macgilliwraya Forbes, 1852, larva of Jonna Britinnich, 1772; Calcarella Souleyet, 1850, probably larva of Distorsio Roding, 1798; Gemella H. & A. Adams, 1858, a cassid; Talisman de Folin, 1884, a bursid larva; and Dissentoma Pilsbry, 1945 (see PILSBRY, 1949) and Ethella H. & A. Adams, 1858, neptunelline larvae. Empty larval shells in sediments were (and still occasionally are) given specific names before being rec- ognized as larval shells. CAZENAVETTE (1853) briefly and FISCHER (1863) in more detail described the young of Zonna and identified it with Macgillivraya. Several authors (e.g., KESTEVEN, 1901, 1902; CLENCH & TURNER, 1957) then figured the larval shells of tonnoids, based on young benthic stages with preserved apical whorls. PELSENEER (1906) briefly described two ranellid larvae from the Bay of Biscay (as Coralliophila sp. A and B). SIMROTH (1911) described Cal- carella in some detail, but did not see the connection with the adult stage. Finally, SCHELTEMA (1971, 1972) and LAURSEN (1981) on a larger scale identified and figured planktonic larvae by comparison with benthic stages. SCHELTEMA (1966) introduced the term “‘teleplanic’’ to characterize the long-lived veligers of Neptunellinae and demonstrated that trans-Atlantic dispersal could take place during their planktonic life. PECHENIK e/ al. (1984) have shown that planktonic larvae of Cymatium parthenopeum (von Salis, 1793) do not grow during their dispersal across the North Atlantic via the North Atlantic drift, but remain Page 58 competent for metamorphosis during periods extending over 300 days. This growth stasis and ability to delay metamorphosis explains the more or less circumtropical distribution for many species of Tonnoidea. (It does not explain, however, why many of them are absent in the tropical eastern Pacific, and most species despite having teleplanic larvae are not cosmopolitan.) We have examined numerous species of Tonnoidea in order to find out to what extent and with what precision larval shells can be identified, because these frequently are encountered in samples of plankton and benthos, and also in order to find out if the larval shells give any clues about the systematic relations of the species. We have then sup- plemented comparison of larval shells from planktonic or benthic catches and young specimens with examination of opercula and radulae, which contain important informa- tion. Under each family we figure representative species, with operculum and radula. A sculpture of thin axial and spiral cords that meet at right angles and are clearly set off from the smooth surface is present on protoconch II of at least some species in most families of Tonnoidea (Figures 79, 90, 92-96, 102-109, 110-113). This type of larval shell is here considered to indicate planktotrophic development. We are not convinced that certain other larval shells with reticulate sculpture built up by axial and spiral ribs, but lacking the smooth surface (e.g., BEU, 1988b:fig. 1B), indicate planktotrophic development. Key to the Recent Families and Subfamilies of Tonnoidea A. Osphradium monopectinate, shell about as broad as high and without well-defined siphonal canal Od ce ee ee AN AOE G Laubierinidae A. Osphradium bipectinate, distinct siphonal canal PRESEN tae cen ict ner aah ura tea ee eae ae B B. Operculum absent or reduced in adult ......... C B. @pereulwm wellidevelopedi acme eee eer D C. Siphonal notch present, larval shell corneous; if calcified, disjunctlyicolled™ aman tenee. Tonnidae C. Siphonal canal formed by the drawn out aperture; larval shell calcified, not disjunctly coiled .. Ficidae D. Aperture with a well-developed posterior siphonal canal, jaws reduced; incremental scars aligned along both sides of shell, separated by 180° rei rr err ren il oso Micha s Oo Bursidae The Veliger, Vol. 33, No. 1 D. Aperture without distinct posterior canal, jaws large Re eee hes en 3.6.58 i030. '0.0 2 E E. Shell globular or triangular with short siphonal Canally Ase 3iie ae loonie ee oe ee Cassidae E. Shell fusiform or bucciniform, often with long ca- 502 lees EDA ee ere Moma EcAD WieiZain'a.a lal eo" 609 0 F F. Parietal shield strongly developed; proboscis very long, strongly coiled when retracted; posterior edge of central radular tooth evenly curved a oar ener cnt Ade occa 0 oc Personidae F. Parietal shield not strongly developed; proboscis straight when retracted; posterior edge of central radular tooth straight ........... G (Ranellidae) G. Shell with regularly appearing varices ......... H G. Shell without varices anasaeee sore Pisanianurinae H. Varices separated about 240° ........ Neptunellinae H. Varices separated about 180° ........... Ranellinae Family RANELLIDAE Gray, 1854 (Figures 25-40, 52-55, 67-70, 85-95, 97-104, 123-127, 142-145, 153, 154) Beu (1985) and BEU & CERNOHORSKY (1986) have pointed out the nomenclatorial reasons for using the name Ra- nellidae instead of the well-established and almost uni- versally used name Cymatiidae, and Personinae instead of Distorsioninae. However, they have continued using Cy- matiinae Iredale, 1913, as the name of the subfamily, referring to ICZN Article 40b. Because the Commission evidently refuses to consider invalidating the name Ra- nellidae, we can see no possibility to avoid using the name Neptunellinae introduced by GRay (1854) at the same occasion as the Ranellidae. It is based on the generic name Neptunella Gray, 1854 (defined by citation of the name “| Murex] cutaceum” (Linné, 1758)) and is an objective junior synonym of Cabestana Roding, 1798. This is a re- grettable consequence of the strict application of the Code. BEu (1988b) convincingly separated the Personidae as a distinct family, present already in the Upper Cretaceous, and distinguished two subfamilies of Ranellidae: Neptunellinae (=Cymatiinae), with a central radular tooth that is distinctly wider than high and is equipped with several lateral denticles; a periostracum with dense axial blades and varices separated by % of a whorl. Ranellinae, with a central radular tooth equipped with a few lateral denticles and about as high as broad; a peri- Explanation of Figures 1 to 8 Radulae of Ficidae and Bursidae. Figures 1 and 2. Thalassocyon bonus (New Zealand, NMNZ). Sc ale li ves 25 and 10 um. Figures 3 and 4. Ficus sp. (Seychelles, MNHN). Scale lines 100 and 25 pm. Figures 5 and 6. Bursa sp. (Gilbert Id., SMNH). Scale lines 100 and 25 um. Figures 7 and 8. Bufonaria marginata (Canaries, SMNH). Scale lines 100 and 25 um. A. Warén & P. Bouchet, 1990 The Veliger, Vol. 33, No. 1 Page 60 A. Waren & P. Bouchet, 1990 ostracum without dense axial blades and varices aligned up the sides of the spire, 7.e., separated by 180°. We will here add one additional subfamily, based on the genus Pisanianura Rovereto, 1899, previously classified in the Buccinidae. However, first we will give some back- ground information about the protoconch of the Ranelli- dae. The Ranellid Larval Shell There is considerable variation in the morphology and development of the larval shell of this family. It is therefore outlined for the different subfamilies, and for species with planktotrophic development. PILKINGTON (1974) described the early young of Sassza (Cymatona) kampyla (Watson, 1885) (Neptunellinae), which belongs to the oldest genus of Tonnoidea, dating back to the Late Cretaceous (Turonian, BEU 1988b). The planktonic stage of this species was mentioned by DELL (1956). In several papers Beu has later figured and de- scribed larval shells of many species of Sassia, which differ from other species of Neptunellinae in being calcareous, globular and often equipped with cancellate sculpture. The morphology and biology of larvae belonging to Cy- matium and Charonia (Neptunellinae) have been described by SCHELTEMA (1971), LAURSEN (1981), RICHTER (1984), and PECHENIK et al. (1984). Characteristic for these is that they are only partly, or not at all, calcified, and have a fairly tall spire, usually considerably higher than the ap- erture (Figures 97-104). The larval operculum has a char- acteristic shape, very slender, with a semicircular external and depressed bell-shaped inner margin (Figure 70). As in the Tonnidae the larval shell is, after settlement, filled by calcareous deposits, so that after destruction of the peri- ostracum only a disjunctly coiled, internal mould remains. This internal mould shows the original sculpture also in species that resorb primarily deposited calcium carbonate (Figures 103, 104). In species of Sassca (Neptunellinae)—e.g., S. remensa (Iredale, 1936), S. parkinsonia (Perry, 1811), and Sassia raulint (Cossmann & Peyrot, 1923) (Figure 90)—there is no interspace between the whorls of the protoconch II, even after treatment with bleach to remove remains of the periostracum. This can only be interpreted in one way, viz. that already the planktonic larva forms a calcareous shell, covered only externally by a periostracum. Explanation Radulae of Tonnidae and Cassidae. Figures 9 and 10. Jonna, larva (off SE South Africa, SMNH). Scale lines 50 um. Figure 11. Ewdolium crosseanum (the Azores, MNHN). Scale line 50 um. Page 61 We consider the sculpture of species of Cymatium, on the inside of the periostracum of the apical whorl (Figure 104) and on the remaining internal mould (Figure 103), of great interest and indicating that these species originate from species with a reticulate larval sculpture of the same type as in, e.g., Sassia. The osphradium of the larva is usually bipectinate in species of Cymatium, but in one species we found a mono- pectinate osphradium (shell: see Figure 98). The larvae of the Ranellinae are less well known, but PILKINGTON (1974, 1976) described the hatching young of Argobuccinum pustulosum (Lightfoot, 1786), Fusitriton ma- gellanicus, and KESTEVEN (1901, 1902), IREDALE (1936), and BEu (1978a, b, 1988b) figured several protoconchs from juvenile specimens. To expand on this, we have ex- amined some late larvae and early post-larvae. A specimen of Ranella australasia taken in surface plankton off south- eastern Australia has a larval shell that shows only traces of an internal calcareous coating, which certainly is of no supporting function (Figure 89). The shell is perfectly smooth and unexpectedly solid, considering that it consists only of periostracum. Immediately after settlement the young starts to form an internal calcareous coating, and later a shell. When the post-larval growth continues, the periostracum starts to disintegrate and finally a perfectly smooth, calcareous, internal mould of the larval shell re- mains (as in Figures 86, 87), following the same pattern as in Cymatium and Charonia. We have examined young benthic stages of Fusitriton magellanicus (Figure 85), Ra- nella olearia (Figure 87), and Argobuccinum pustulosum tumidum (Figure 86) that have larval shells conforming with this developmental model. Usually one can see a distinct interspace between the whorls, where once there was periostracum (Figure 87). The larval shell of the Pisanianurinae is characteristic in its large size (2.5-3.5 mm high), usually brown color, and globular shape without a distinct siphonal canal and especially in the perfectly reticulated sculpture with large smooth squares between raised axial and spiral ribs that meet at close to right angles (Figures 90, 96). It resembles that of the Laubierinidae (Figures 105, 106), but they have a still larger and proportionally broader larval shell (5-6 mm high) with an expanded aperture and a more regular spiral arrangement of the minute tubercles on pro- toconch I (cf. Figures 108 and 142). Most competent ranellid larvae have a radula similar of Figures 9 to 16 Figure 12. Jonna sp. (Madagascar, SMNH). Scale line 50 um. Figures 13 and 14. 7onna allium (Sumatra, SMNH). Scale lines 250 um. Figures 15 and 16. Oocorys sulcata (off S Portugal, MNHN). (For detail of marginal teeth, see Figure 24.) Scale lines 50 um. The Veliger, Vol 335.Noa Ae \\ \\ Ny); Jw OO FP gg ADK LAAN papi aMule 0/9 ks TT \\ {TP 7/4 _ WO RY = HAN? HTPC VU fm AAW! NIN / é CGA) ; PRS A. Warén & P. Bouchet, 1990 to adults of the species (Figures 30, 34, 36-38, 41, 42), but PELSENEER (1906:pl. 10, figs. 1-4) reported a species missing the radula and we noticed the same in a larva with a similar shell. Subfamily PISANIANURINAE Waren & Bouchet, subfam. nov. Diagnosis: Tonnoidea with a medium-sized Buccinum- like shell lacking varices, with poorly developed siphonal canal. Columella curved, contributing to giving the aper- ture a more rounded appearance. Operculum with cor- roded apical nucleus. Proboscis very short, broad, and mus- cular. Osphradium bipectinate with left row of leaflets less developed than inner ones. Pisanianura Rovereto, 1899, has always been regarded as a buccinid genus, although FISCHER (1883) and LOCARD (1897) evidently had some doubts about it and compared P. grimaldu with Oocorys. We have had access to a few specimens preserved with soft parts and the anatomy (pre- sented below) does not conform with a position in the Neogastropoda. The salivary glands have the typical struc- ture that occurs only in Tonnoidea, and there is no devia- tion in other anatomical features from what is known in that superfamily. Therefore, we conclude that Pisanianura should be transferred to the ‘Tonnoidea. The shell morphology agrees well with the Ranellidae, except that it lacks varices (z.e., it does not have periodical growth) and that the columella has a peculiar, curved shape. The anatomy is less useful for assigning Pisanianura to family because anatomical features, as far as they are known, are quite uniform throughout the superfamily and the differences so subtle and incompletely known that a detailed analysis can hardly be undertaken presently. It is, however, obvious that the families are based on anatomical or shell morphological specializations, and be- cause we are not suggesting anything better, we do not want to disturb this order. Therefore, we can exclude a position in the following families: 1. Ficidae. Salivary glands simple; proboscis pleurem- bolic, long and slender; buccal mass small and without large retractors. 2. Tonnidae. Foot large, operculum missing in adult; Page 63 distal part of proboscis suckerlike; jaw with a strong hook; shell globular and inflated. 3. Bursidae. Jaws reduced or absent. Varices present. Posterior canal strongly developed. 4. Personidae. Proboscis very long and slender, coiled when retracted. Varices present. 5. Cassidae. Central radular tooth broad and low. Shell globular with short canal. 6. Laubierinidae. Osphradium monopectinate. Shell with poorly developed siphonal canal. The Ranellidae on the other hand consists of species with obviously less specialized or modified morphology and we Can seé no reason against a position therein. (This may give reason to suspect that the classification is based on grades rather than clades, with the Ranellidae to some extent constituting a less modified stock. This may be true, but this classification is practical and in the absence of detailed anatomical information that gives a better reso- lution, we prefer to keep it.) The assignment of Pisanianura to subfamily offers more problems. In the Neptunellinae the proboscis is retracted by numerous small muscles attached to the sheath (HousBRIcK & FRETTER, 1969), whereas in Pisanianura there are two major muscles attached to the buccal mass and additional muscles are inconspicuous. Another differ- ence from the Neptunellinae is the relative size of the buccal mass, which in the Neptunellinae is small in re- lation to the proboscis sheath, whereas in Pisanianura it is large (for drawings of this see, e.g.: PANCERI, 1869; HAL- LER, 1893; SIMROTH, 1896-1907; AMADRUT, 1898; HousrRick & FRETTER, 1969). It is difficult to compare anatomically Pisanianura with the Ranellinae because few species of that group have been described anatomically (HALLER, 1893), but it seems that the species of Ranellinae closely resemble the Neptunel- linae. Also the shell morphology contradicts a position in the Ranellinae. Beu (pers. comm.) considers this a uniform, monophyletic taxon, characterized by the neat alignment of the varices along the sides, separated by 180°. We, therefore, supported by Beu (pers. comm.), consider Pisanianura a monophyletic group distinct from the Nep- tunellinae and Ranellinae, and erect a new subfamily for Pisanianura, the Pisanianurinae, in the Ranellidae. Explanation of Figures 17 to 24 Radulae of Cassidae. Figures 17, 18, and 22. Galeodea echinophora (Mediterranean, SMNH). Scale lines 100, 50, and 50 um. Figures 19 and 20. Semicassis granulatum (Mediterranean, SMNH). Scale lines 500 and 100 um. Figure 21. Semicassis saburon (West Africa, MNHN). Scale line 100 um. Figure 23. Cypraecassis testiculus, larva from plankton (Dana sta. 1286). Scale line 10 um. Figure 24. Oocorys sulcata, marginal teeth (off S Portugal, MNHN). (For a figure of the whole radula, see Figure 15.) Scale line 100 um. Page 64 It is also possible that the Pisanianurinae should be considered a distinct family in the Tonnoidea, because it (1) lacks varices, which evidently is a primitive character in the group and (2) has the left row of leaflets in the osphradium less developed (equal size in all other tonnoids except the Laubierinidae ). This view is supported by Beu (pers. comm.). The subfamily Anochetinae Cossmann, 1901, was erect- ed with Pisanianura as “type genus” (in the Buccinidae), but it is not based on a generic name and therefore has no nomenclatural status. Pisanianura Rovereto, 1899 Anura BELLARDI, 1873:201 (not Anura Hodgson, 1841). Type species: Murex inflatus Brocchi, 1814, subsequent des- ignation COSSMANN (1901:178). Pisanianura ROVERETO, 1899:104, new name for Anura Bel- lardi, 1873 (preoccupied several times). New synonyms: Kaiparanura Laws, 1944:309. Type species by original des- ignation: Phos spiralis Marshall, 1918, Miocene, New Zealand (Figures 142-145). Laminilabrum Kuroda & Habe in HaBE, 1961, app.:13. Type species by original designation: L. breviaxe Kuroda & Habe, 1961, Recent, deep water, Japan. Nawenia LADD, 1977:51. Type species by original desig- nation: N. bartholomewi Ladd, 1977, Pliocene, west Pa- cific. Remarks: Laminilabrum was described in the Trichotro- pidae, Kazparanura in the Cominellidae (=Buccinidae), and Nawenza in the Buccinidae. These positions have not been questioned since. The soft parts of Pisanianura and Laminilabrum (described after the species, below) do not indicate differences other than of specific rank. The shells of all four genera have in common a multispiral protoconch with a large-meshed reticulate sculpture, a teleoconch without varices, and a stongly curved columella. They differ mainly in the degree of axial sculpture on the te- leoconch, with Laminilabrum having no axial sculpture, Nawenia axial sculpture on the first two whorls, and Pi- sanianura and Kaiparanura on all the whorls. We regard these differences as of specific rank, and consequently treat the generic names as synonyms. The type species of Pisanianura is a Pliocene fossil from northern Italy. Brocchi’s type material has been figured The Veliger, Vol. 33, No. 1 by Rosst RONCHETTI (1955:201) and PINNA & SPEZIA (1978:pl. 34, fig. 3) and we have examined Pliocene ma- terial from Pradalbino, Bologna, Italy, kindly sent on loan by Mr. Della Bella (Figures 93, 125). The larval shell is multispiral with cancellate sculpture (Figure 93) and the teleoconch differs from the recent P. grimaldi mainly in the presence of a peripheral keel. Beside the type species, BELLARDI (1873) included Fusus borsoni (Gené in Bellardi & Michelotti, 1840) in Anura and described A. ovata, A. striata, A. craverii, A. pusilla, and A. sublaevis from the Italian Miocene and Pliocene. We have not examined these species, some of which recently have been figured by FERRERO Mortara e¢ al. (1981:pl. 7). Judging from the figures it seems certain that the Mio- cene A. craverii belongs to Pisanianura; the status of the other species is more doubtful. The genus is furthermore represented in the European Tertiary deposits by Pisanianura aturensis Peyrot, 1927, and P. benoisti Peyrot, 1927, from the Oligocene and Mio- cene of southwestern France respectively. (P. degrangei Peyrot, 1927, is probably a young cerithid related to Gour- mya Fischer, 1884 [Cerithidae]; P. Lozouet, pers. comm.) The type species of Kaiparanura is a Lower Miocene fossil from New Zealand; the teleoconch has a sculpture of coarse axial ribs. LAws (1944) introduced this new genus because he had been misled by COSSMANN’s (1901) de- scription of the protoconch of Pisanianura inflata, which erroneously had been described as sculptured with weak spiral threads and with a smooth and depressed nucleus. This synonym was pointed out to us by Dr. A. Beu. Finally, the type species of Nawenza is a Pliocene fossil from Fiji, similar to Pisanianura breviaxe but differing in having stronger axial sculpture on the first two teleoconch whorls. Pisanianura is thus known from at least two Paleogene and two Neogene North Atlantic species, two Neogene South Pacific species, and two Recent (sub)tropical deep- water species. The taxonomy and distribution of the Recent species only are here treated in detail. Pisanianura grimaldu (Dautzenberg, 1889) (Figures 25; 26,55, 68; 94595. 12655127) Hindsia grimaldii DAUTZENBERG, 1889:33, pl. 2, fig. 4. Anura clathrata DAUTZENBERG & FISCHER, 1906:25, pl. 3, figs. 6-8. (New synonym.) Explanation of Figures 25 to 32 Radulae of Ranellidae. Figures 25 and 26. Pisanianura grimaldiu (New Caledonia, MNHN). Scale lines 50 um. Figure 27. Pisanianura breviaxe (New Caledonia, MNHN). Scale line 50 yn Figure 28. Sassia kampyla, very young (New Zealand, NUNZ 9196). Scale line 25 um. Figure 29. Sassia parkinsonia (New South Wales, AMS C 50074). Scale line 50 um. Figure 30. Ranella australasia, planktonic larva (New South Wales, AMS C 147218). Scale line 25 um. Figures 31 and 32. Ranella olearia (the Azores, MNHN). Scale lines 200 and 100 um. A. Waren & P. Bouchet, 1990 \ ma. ei