1VIBL/WH01 THE NAUTILUS Volume 108 1994-1995 AUTHOR INDEX Allmon, W. D 15 Lanfair, G. A 25 Dayan, N. S 49 Marshall, B. A 1, 27, 83 Dillon, R. T. Jr 49 McLean, J. H 39, 80 Fell, P. E. 42 Morris, P. J 15 Filippova, Yu. a 67 Nesis, K, N 67 Gordon, ME 55 Nikitina, I. V 67 Harasewych. M G 61 Pierce, H G 23 Helvenston, L. L 42 Smith, ME 25 HouART, R 27 Spelke, J A 42 Kabat, a. R 61 TippETT, D. L 37 Khromov, D. N 67 Tucker, J. K 37 KiLBURN, R. N 34 Yokes, HE 9 Kohl, B 9 NEW TAXA proposed IN VOLUME 108 (1994-1995) gastropoda Osteopeltidae Osteopelta praeccps Marshall, 1994, new species 3 Skeneidae Bruceiella laevigata Marshall, 1994, new species 5 Bruceiella pruinosa Marshall, 1994. new species 5 Trochidae Costomargarites McLean, 1995, new subgenus 80 Margarites (Costomargarites) haxteri McLean, 1995, new name 81 Calliostomatidae CaUiostoma (Otukaia) alcrtae Marshall, 1995, new name 117 Calliostoma (Maurea) antipodense Marshall, 1995, new species 105 CaUiostoma (Maurea) aupourianum Marshall, 1995, new species 97 Calliostoma (Maurea) eminens Marshall, 1995, new species 100 Calliostoma (Maurea) gibbsorum Marshall, 1995, new species 93 ' Calliostoma (Maurea) jamiesoni Marshall, 1995, new species 93 Calliostoma (sensu lata) hopua Marshall, 1995, new species 123 Calliostoma (serisu lata) limatulum Marshall, 1995, new species 122 Calliostoma (Maurea) maui Marshall, 1995, new species 106 Calliostoma (Maurea) penniketi Marshall, 1995, new species 115 Calliostoma (Maurea) regale Marshall, 1995, new species 96 Calyptraeidae Grandicrepidula McLean, 1995, new subgenus 80 Naticidae Bentliobulbus McLean, 1995, new genus 39 Euiimidae Pseudosabinella McLean, 1995, new genus 39 Muricidae Poirieria syrinx Marshall & I louavt, 1995, new species 30 Ocenotrophon McLean, 1995, new genus 40 Buccinidae Retimohnia McLean, 1995, new geiuis 40 Mitridae Charitodoron rosadoi Kilburn, 1995, new species 34 Turridae Pseudotaranis McLean, 1995. new genus 81 Leucosijrinx kantori McLean, 1995, new name 81 Xylodisculidae Xylodiscula osteophila Marshall, 1994. new species 7 THE NAUTILUS Volume 108, Number 1 June 17. 1994 ISSN 0028-1344 A quarterly devoted to malacology. Marine Biologicai Laboratoiy/ Woods Hole Oieanographic Institution Library JUN 3 0 1994 Woods Hots. MA 0;i543 EDITOR-IN-CHIEF Dr. M. G. Harasevvych Division of Mollusks National Museum of Natural History Sniitlisonian Institution Wasliington, DC 20560 ASSOCIATE EDITOR Dr. R. Tucker Abbott American Malacologists, Inc. P.O. Box 2255 Melbourne, FL 32902 CONSULTING EDITORS Dr. Riidiger Bieler Department of Invertebrates Field Museum of Natural History C;hicago, IL 60605 Dr. Robert T. Dillon, Jr. Department of Biology (lollege of Charleston Charleston, SC 29424 Dr. William K. Emerson Department of Living Invertebrates The American Museum of Natural History New York, NY 10024 Dr Robert Hershler Division ol Mollusks National Museum of Natural History Smithsonian Institution Washington, DC 20560 Mr. Richard 1. Johnson Department of Mollusks Museum of Comparative Zoolog\ Harvard University Cambridge, MA 02138 Dr. .\urele La Rocque Department of Geology The Ohio State University Columbus, OH 43210 Dr. James H. McLean Department of Malacology Los Angeles County Museum of Natural History 900 Exposition Boulevard Los Angeles, CA 90007 Dr. Arthur S. Merrill % Department of Mollusks Museum of Comparative Zoology Harvard University Cambridge, MA 02138 Ms. Paula M. Mikkelsen Harbor Branch Oceanographic Institution, Inc. Ft. Pierce, FL 33450 Dr. Donald R. Moore Division of Marine Geology and Geophysics Rosenstiel School of Marine and Atmospheric Science University of Miami 4600 Rickenbacker Causeway Miami, FL 33149 Dr. Gustav Paulay Marine Laboratory University of Guam Mangilao, Guam 96923 Mr. Richard E. Petit P.O. Box 30 North Myrtle Beach, SC 29582 Dr. P'dward J. Petuch Department of CJeology Florida Atlantic University Boca Raton, FL 33431 Dr David H. Stansbery Museum of Zoology The Ohio State Universit> Columbus, OH 43210 Dr. Ruth D. Turner Department of Mollusks Museum of Comparative Zoology Harvard University C:ambridge, MA 02138 Dr. Geerat J. Vermeij Department of Geology University of California at Davis Davis, CA 95616 SUBSCRIPTION INFORMATION The subscription rate per volume is US $25.00 for individuals, US $40.00 for institutions. Postage outside the United States is an atlditional US $5.00 for surface and US $12.00 for air mail. All orders should be accompanied by payment and sent to: THE NAUTILUS, P.O. Box 7279, Silver Spring, MD 20907-7279, USA. Change of address: Please inform the publisher of your new address at least 6 weeks in advance. All communications should include both old and new addresses (with zip codes) and state the effective date. THE NAUTILUS (ISSN 0028- 1344) is published quarterly by Trophon Corporation, 8911 Alton Parkway, Silver Spring, MD 20910. Second Class postage paid at Silver Spring, MD and additional mailing offices. POSTMASTER: Send address changes to: THE NAUTILUS P.O. Box 7279 Silver Spring, MD 20907-7279 T H E t^N AU T I L U S CONTENTS Volume 108, Number 1 Jiiiw 17. 1994 ISSN 0028-1344 Bruce A. IHarshall Deep-sea Gastropods from the New Zealand Region Associated with Recent Whale Bones and an Eocene Turtle . Barry kohl Harold E. Yokes On the Living Habits of Aceita buUiai (\'okes) in Cheniosv nthetic Bottom Communities, Gulf of Mexico Paul J. Morris ^ arren D. Allnion Shell Alignment for the Morphometric Analysis of High- spired Gastropods 15 Harold G. Pierce On Planorbula nebraskensis Leonard, 1948, and its position in the evolutionar\ sequence of North American planorbulids 23 Michael E. Smith Gerrv A. Lanfair Effects of Preservati\es on Wet-weight Biomass of the Asiatic Clam, Corbicula fluminca 25 Marine Blolooical Laboratory/ Wood! Hole Oteanographic IrtttitutJon Liiirary JUN 3 0 1994 Woods Hot*. MA 02543 THE NAUTILUS 108(1):1-S, 1994 Page 1 Deep-sea Gastropods from the New Zealand Region Associated with Recent Whale Bones and an Eocene Turtle Bruce A. Marshall Museum of New Zealand Te Papa Tongarevva P.O. Box 467 Wellington, New Zealand ABSTRACT Five species of gastropods are new iy recorded from deca> ing whale bone from the deep-sea floor off New Zealand: Para- cocculina cervae (Fleming, 1948). and new species of Osteo- pelta. Marshall, 1987 (Cocculiniformia), Brucciclla Waren & Bouchet, 1993 (Vetigastropoda), and Xylodiscula Marshall, 1988 (Heterobranchia). RecentK discovered material of Osteopelta mirabilis is recorded, and this or a closely similar species is recorded associated with turtle bones from the Middle Eocene of New Zealand. The new Bruceiella and Xyhidiscula species are the first records of skeneimorph gastropotis from bone. .\part from w hale bone, Paracocculina cervae is associated with sunken wood and algal holdfasts (nev\ record), so it has the most generalised habitat of any know n cocculiniform limpet. Key words: Recent, fossil, Mollusca, Gastropoda, deep-sea, w hale bone, turtle bone. New Zealand. INTRODUCTION Since discovery of the limpet Osteopelta mirabilis Mar- shall, 1987, the first record of a gastropod living in as- sociation with decaying whale bones, a second species of Osteopelta has been recorded from bone off Iceland (Waren, 1989). More recently McLean (1992) has re- corded a nev\ species of Coeeiilina Dall, 1882 (Coccu- linidae) and three species of Pywpelta McLean & Hasz- prunar, 1987 (Pyropeltidae) from whale bone from the eastern Pacific. One of these pyropeltiii limpets was de- scribed as new, w hile the other two were evidently con- specific w ith specimens originally obtained from hydro- thermal vents (McLean & Haszprunar, 1987). Dell (1987) has reviewed my tilid bivalves associated with whale re- mains. Additional ta.xa have been recorded by Smith et al. (1989), who drew attention to similarities between faunas associated with decaying whale bones, hydro- thermal vents and hydrocarbon seeps, and suggested that whale skeletons may provide stepping stones for wide dispersal of deep-sea chemosynthetic communities (see also Smith, 1992). In this paper I introduce a third species of Osteopelta. another cocculinid, and species of Bruceiella \\'aren & Bouchet, 1993 and Xylodiscula Marshall, 1988 new to the fauna of this unusual habitat Bruceiella was based on a species from a hydrothermal vent, while previously known Xylodiscula species occurred in association with sunken wood, old sea grass (Posidonia) fibres, and a hy- drothermal vent. ABBREVIATIONS .\MS .Australian Museum, S\dne\ BMNH The Natural History Museum, London L.\CM Los ,\ngeles Count>- Museum of Natural His- tor\' MNHN Museimi National d'Histoire Naturelle, Paris MNZ Museum of New Zealand, Wellington NMP Natal Museum, Pietermaritzburg OL' Geology Department, Uni\ersit\ of Otago, Dunedin USNM National Museum of Natural History, Wash- ington, DC Order ARCHAEOGASTROPODA Thiele, 1925 Suborder COCCULINIFORMIA Haszprunar, 1987 SuperfamiK COCCULINOIDEA Dall. 1882 FamiK COCCULINIDAE Dall, 1882 Genus Paracocculina Haszprunar, 1987 Paracocculina Haszprunar, 1987:321. Type species (by original designation!: Cocculina laevis Thiele, 1903: Recent, off Nias Island, Sumatra. Remarks: The genus Paracocculina was introduced by Haszprunar (1987) for species that differ from Cocculina Dall, 1882 primarily in having a prominent subpallial gland, a pedalK innervated copulator\ organ on the right side of the foot, and a hypoathroid nervous system with the osphradial ganglion situated at the far left side. Page 2 THE NAUTILUS, Vol. 108, No. 1 Figure 1. Map of New Zealand region showing localities for Osteopelta praeceps n. sp. and O. mirahilis (stars), BruceieUa laevigata n. sp. (solid circle), and BruceieUa pruinosa n. sp. (open circle), 200 and 1,000 meter contours indicated. Paracocculina eervae (Fleming, 1948) Cocculina eervae Fleming, 19-48:88, text fig. la-d; Marshall, 1986:508, figs. 2A, 3A-C, 12AB. Tecticrater eervae. Dell, 19.56:60; Powell, 1979:81, fig. 10/9 Paraeoeeulina eervae. Haszprunar, 1987:321. IVlaterial examined: (Additional to that recorded bv Marshall, 1986); 39°53.2'S, 168°01.2'E, Challenger Pla- teau, New Zealand, alive on whale bone, 908-912 m, 3 July 1989, f.v. Amaltal Explorer (3 MNZ); 44°40.5'S, 174°01.0'E, off Banks Peninsula, alive on whale skull, 844 m, 11 May 1987, f.v. Oijcing 7 (5 MNZ); 38°38.9'S, 178°38.3'E, off Gable End Foreland, New Zealand, alive on algal holdfast, 529-565 ni, 23 September 1985, f.v. Wanaka stn WK2/68/85 ( 1 MNZ); 43°02.5'S, 174°09.0'E, off Kaikoura, New Zealand, alive on algal holdfast, 848- 877 m, 24 September 1988, f.r.v. James Cuok stn J12/ 10/88 (7 MNZ); ,50°02.6'S, 169°31 'F, east of the Auckland Is, New Zealand, alive on algal holdfast, 614-620 m, 4 December 1992, f.r.v. Tangaroa stn 92011/97 (many MNZ). Distribution: North ('ape to the .Auckl; ds, Nt Zealand, 18-891 ni, on sunken wood, algal holdfasts and whale bone. Remarks: Specimens from whale bone and deep-sunken algal holdlasts and wood proved to be indistinguishable in shell and radular morphology and in e.xternal anatom\ . Superfamilv LEPETELLOIDEA Thiele, 1908 Family OSTEOPELTIDAE Marshall, 1987 Genus Osteopelta Marshall. 1987 Osteopelta Marshall, 1987:121. T\pe species (by original des- ignation): Osteopelta mirabilis Marshall, 1987; Recent, New Zealand. Osteopelta mirabilis Marshall, 19S7 Osteopelta mirabilis Marshall, 1987122, figs. lA. B-D, F-J, 2A, 2B, 3C; Haszprunar, 1988:6, figs. 1.5-24 (anatomy). Material examined: (.ALilditional to that recorded by Marshall, 1987); .39°.53.2'S, 168°01.2'E, Challenger Pla- teau, New Zealand, alive on whale bone, 908-912 m, 3 July 1989, f.v. Amaltal Explorer (18 MNZ); off Mernoo Bank, Chatham Rise, New Zealand, alive on large whale skull, ca. 900 m, October 1988, f.v. Amaltal Explorer (5 MNZ). Distribution (figure 1): Challenger Plateau and Chat- ham Rise, New Zealand, on whale bone, 800-955 m. Remarks: The Challenger Plateau specimens are in- distinguishable from type and other material from the Chatham Rise. The protoconch is retained in a specimen 5,00 mm long, and is bilateralK symmetrical and 200 /um long, with a long tapered apical fold tip on each side. Regrettably the sculpture is unknown, as the outer shell layer has been etched away. The Icelandic species Os- teopelta ceticola Waren, 1989, also from whale bone, differs in minor details of radular morphology (Waren, 1989). Osteopelta sp cf. mirabilis Marshall. 1987 IVlaterial examined: Isolated concretion of Waihao Greensund on right bank of South Braiiich of Waihao River, north of Waihao Dow ns (map ref. J40/45000030), closely associated with bones of fossil leatherback turtle (Dermochelys sp. — Reptilia, (-helonia. Dermochelyi- dae), coll. R. E. Fordyce, A. Grebneff, C. M. Jones and P. A. Maxwell, .^ge: Bortonian (Middle Eocene) (1 spec- imen Ol'41112). Remarks: The single specimen is enibeddetl api.'.\ (low n in a block of gritt\ niatri.x with the full\ exposetl interior surface showing clearK tlefineil muscle scars. .\ mould of the exterior where a small piece of the shell margin has liroken away shows impressions of weak concentric sculpture. The specimen is indistinguishable from the Recent specimens of Osteopelta mirahilis in shape, shell thickness, teleoconch sculpture, muscle scar outline, and size (length est. 7.50 nun, width (i,05 mm). While it is B. A. Marshall, 1994 Pages impossible to be absoluteh certain of the relationships of this limpet from the teleoconch alone, particularK the interior, the association with bone suggests that it is prob- alily an osteopeltid. The W'aihao Green sand beds w ere considered by Max- well (1992) to ha\e been deposited at 150-250 ni depth, which is substantialK shallower than the batlnmetric range of li\ing O. mirabilif; (800-955 m), Osteopella praeceps n. sp. (figures 2-6. 18, 19) Description: Shell up to 6,25 mm long, translucent white, thin and brittle, high])- arched, anterior end occup\ing 48.8-62.5% of shell length in specimens over 4 mm in length. .Aperture elliptical, anterior end more narrowly rounded than posterior; apertural plane shallowly con- cave at ends, weakly convex at sides; broadest slightK' behind midlength. .\pex tightK- rounded, anterior slope flat or weakK conca\e, posterior slope broadly convex, lateral slopes weakK con\ex. Periostracuni transparent, ver> thin, smooth. Protoconch 200 urn long, bilaterally symmetrical, api- cal fold producing a small, shallow dimple on each side, sculptured throughout with densely crowded minute punctations arranged in spiral lines, Teleoconch sculptured with fine collabral grciw tli lines, obscure radial lines, and many minute, randomly dis- tributed, shallow pits. Animal white. Foot broad and thick. Mantle edge thickened, with weak left anterolateral fold. Snout very large, tapered, longer than broad, deeply concentrically wrinkled, mouth a vertical slit in small, circular, flattened tip. No oral lappets. Cephalic tentacles slender, tapered, tips rounded, minute black eyes sunken in swellings at outer bases. A large secondary gill extending along right mantle groove to above right of head, comprising about 30 anteriorly enlarging leaflets. Seminal groove not de- tected. Two slender blunt-tipped, dorsoventrally flat- tened epipodial tentacles at posterior end. Radula (figures 18, 19) with the formula oo -I- 6 + 1 -H 6 + oo, indistinguishable from that of O. mirahilis. Type data: Holot\'pe MXZ Ml 16970 (length 5.50 mm, width 4.20 mm, height 3.40 mm) and 57 paratypes (51 MNZ; others AMS, BMNH, LACM, MNHN, NMP, USNM): 43°34.14'S, 176°18.69'E, E of Mernoo Bank, Chatham Rise, New Zealand, 372-379 m, 29 January 1992, f.r.v. Tangaroa stn 9106 168, alive on whale ver- tebra among crowded indi\iduals of a species of Idas Jeffreys, 1876 (Mytilidae). Other material examined: (Several hundred specimens MNZ): Topotypes (several hundred juveniles, MNZ M. 117279); 39°53.2'S, 168°01.2'E, Challenger Plateau, New Zealand, 908-912 m, 3 July 1989, f.v. Amaltal Ex- plorer stn 34S ' 1 29, alive on a piece of whale bone among crowded mytilids (Idas sp. ) (8 subadult specimens, MNZ M.92453). Distribution (figure 1): C^hatham Rise and Challenger Plateau, New Zealand, on whale bone, 372-912 m. Remarks: Ostcopclta pracccp.s differs from O. mirahilis in attaining smaller size and in ha\ing a narrower, taller shell w ith a longer anterior end. Their external anatomies and radulae are extremely similar. O. praeceps and O. mirahilis occurred li\ing together on the same piece of whale bone from the Challenger Plateau (M.92453 and M. 92451, respectively). Immature specimens of O. praeceps are strikingly sim- ilar to young of the cocculinid Paracocculina cervae that also occurred on the C^hallenger Plateau bone. P. cervae, however, is distinguishable by its larger adult size, lack of shell micropunctations, in having a prominent copu- latory organ behind the right cephalic lappet, and in having a snout that is short and broad instead of pro- boscis-like. Moreover, their radulae and anatomies are entirely different The highK arched shell and narrow aperture of O. praeceps enable it to live deeply (and inconspicuously) among the associated mstilids, which form densely crowded aggregations at nutrient-rich sites on the bones. By contrast, the larger and broader-shelled species O. mirahilis has been obser\ed adjacent to m\ti!id aggre- gates (Marshall 1987: fig. L\), and is probably unable to live as deeply among them. Etymology: Latin praeceps (steep). Suborder VETIGASTROPODA Salvini-Plawen, 1980 Superfamilv TROCHOIDEA Rafinesque. 1815 .^FamiK SKENEIDAE Clark, 1851 Genus Bruceiella Waren & Bouchet, 1993 Brucciclla Waren & Bniichet, 1993:26. Type species (by orig- inal designation): Bruceiella globulus Waren & Bouchet, 1993; Recent, North Fiji Basin (hydrothermal vent). Remarks: In ha\ing epipodial tentacles and papillate cephalic tentacles, the animal of Bruceiella bears a gen- eral resemblance to those of members of the family Ske- neidae, the limits of which have been tightly restricted by Waren (1992) and Waren and Bouchet (1993). The lack of a propodial penis and the distally expanded snout, however, led Waren and Bouchet (1993) to suggest that the genus probably does not belong in Skeneidae, but by implication perhaps in a family of its ow n. The radula bears a striking resemblance to those of a variety of archaeogastropods associated with hydrothermal vents and seeps, especially neomphalids and peltospirids, a sim- ilarity that Waren and Bouchet (1993) consider to be the result of convergence. The otherwise simple shell of Bruceiella species is dis- tinctive in having a protoconch sculpture of minute den- dritic threads and granules arranged in fine spiral lines, and a pronounced varix almost immediately after the protoconch-teleoconch boundary. Unlike Waren and Bouchet (1993), who interpret the varix as the termi- nation of the protoconch, I consider that the \arix is on the teleoconch. and that the actual protoconch teleo- Page 4 THE NAUTILUS, Vol. 108, No. 1 Figures 2-14. Shells of Osteopelta and Bruceiella spj) 2-6. Osliopilla pnieccps ii- sp, 2, .1. Holot\pe, length 5,50 nnn. 4. Immature specimen, Challenger Plateau, MNZ M,9245;5, length 2.20 mm. 5, 6. Protoconch of liolot) pe. 7-9. BruccicUa laevigata n, sp., holot\pe, height 1.70 mm. 9. Protoconch, with protoconch /teleoconch boundary arrowed. 10-14. Bruceiella prninosa n. sp., holotype, height 1.43 mm. 10. Detail of inner part of base. 13. Protoconch. with protoconch teleoconch bonndar\ arrowed. 14. Detail of protoconch sculpture. Scale bar 14 = 10 ^m, others = 100 fim. B. A. Marshall, 1994 Page 5 Figure§ 15-17. Shell of holntvpt- (it XijIotliMula u^teopliila u sp., width l,(iO mm 17. Protdcunch, note inroiled tip. Scale bar 100 Mm. conch boundary is a sharpK defined line at a distance behind the vari.x equi\ aleiit to about one vari.x thickness (figures 9, 13). I also disagree with Waren and Bouchet (1993) that the varix in their genus VeiUsia is part of the protoconch and instead consider that it is situated at the end of the first quarter teleoconch whorl. Species of the related genus Xyloskenea Marshall, 1988 exhibit a change in teleoconch sculpture following a sharply defined growth scar at an equivalent position (Marshall, 1988, figs: 4EJ, 5E; Waren & Bouchet, 1993: fig. 23A). The zone between the varix and the protoconch/ teleoconch boundary is analogous to protoconch II of higher gastropods, though not homologous because protoconch enlargement through marginal incrementation is unknown in archaeogastro- pods other than Neritomorpha. These growth distur- bances probably represent a period of crisis, perhaps a change in feeding mode during the earliest stages of postsettlement development Bruceiella laevigata n. sp. ^figures 7-9, 20-22) Description: Shell up to 1.70 mm higli, slightK higher than broad, thin, translucent, gloss\ , with narrow um- bilical chink, periostracum smooth. Protoconch 300 fxm w ide, delineated b\ fine groove, apical fold tip broadly roundetl, sculptured w ith minute anastomosing dendritic threatis arranged in fine spiral lines. Teleoconch of up to 2.20 strongh and rather evenly con\ ex whorls. First sixteenth w horl minuteU granulate; next sixteenth whorl occupied b\ strong, rounded \'arix, immediateK follow ed b\ fine groo\ e; elsew here smooth. .\perture subcircular, lips thin, parietal contact area nar- row. .Animal. Snout subquadrate; cephalic tentacles dorso- ventralK flattened, tapered, similar, edges ciliated; foot large, anteriorly indented. Small right and large left sub- optic tentacle, 3 small right epipodial tentacles beside operculum, 1 large left epipodial tentacle, 2 small left epipodial tentacles beside operculum. No eyes. Radula (figures 20-22) with the formula co -^ .5 -t- 1 -I- 5 -I- oo, teeth longer than broad. Central tooth stout; cutting area narrow ly angulate, w ithout secondary cusps, proininently hooded; shaft face subtriangular, suddenly narrowed at base. Lateral teeth stout, enlarging out- wards, cutting areas large, hooded, roundly angulate, coarsely serrate, terminal cusp largest; shafts outwardly bowed, strongly flanged and con\'oluted to interlock with adjacent teeth. Marginal teeth slender, outwardly nar- rowing and with smaller cutting areas and finer cusps, outermost few pairs w ith spathulate tips and fused shafts. Type data: Holotype (height 1.70 mm, diameter 1.50 mm, 2.20 teleoconch whorls) MNZ M. 116969; paratypes (5 MNZ, 1 AMS, 1 MNHN): 43°00.17'S, 174°05.5'W, NE of Chatham Islands, New Zealand, 1,242 m, 23 August 1989, f.v. Otago Buccaneer tow 31; alive amongst crowd- ed mytilids (Idaa sp. ) on a whale skull. Distribution (figure 1): Northeast of Chatham Islands, New Zealand, on whale bone, 1,242 m. Remarks: Compared w ith the ty pe species. Bruceiella laevigata is more tightly coiled with a higher spire, but otherw ise the two species are similar. .Among the host of superficially similar skeneimorph gastropods known from the New Zealand region (MNZ — many undescribed), B. laevigata is characterised by the combination of distinc- tise protoconch sculpture, the strong postlarval varix, and tlie radular morphology. Etymology: Latin laecigalus (smooth), alluding to the lack of shell sculpture after the postlar\al \arix. Bruceiella pruinosa n. sp (figures 10-14, 23) Description: Shell (holotype) 1.43 mm high, slightly higher than broad, thin, translucent, glossy , with narrow umbilical chink, periostracum smooth. Protoconch 280 nm w ide, delineated by fine groove, apical fold tip broadly rounded, sculptured with minute anastomosing dendritic threads arranged in fine-spiral lines. Teleoconch of 2.10 strongly and rather e\enly convex Page 6 THE NAUTILUS, Vol. 108, No. 1 B. A. Marshall, 1994 Page 7 whorls. First sixteenth whorl niinutel\ granulate; next sixteenth whorl occupied by strong, roundeil varix, ini- mediateK lollowed b\ fine groove. Adapical quarter of spire whorls, and inner half of base w ith minute granules and very fine axial w rinkles, stronger on base. Aperture subcircular, lips thin, parietal contact area narrow. .Animal. Similar to that of B. laevigata but w ith 1 large right and 2 large left epipodial tentacles, instead of 4 right and 3 left epipodial tentacles, Radula (figure 23) as in B laevigata. Type data: Holot>pe M\Z M.11696S (heigiit 1.43 mm, diameter 1.33 mm, 2.10 teleoconch whorls): 39°53.2'S, 168''01.2'E. Challenger Plateau, New Zealand, 908-912 m, 3 July 1989, f.v. Amaltal Explorer stn 348/129; alive amongst crowded mytilids {Idas sp.) on a piece of whale bone. Distribution (figure 1): Challenger Plateau, New Zea- land, on w hale bone, 908-912 m. Remarks: Bruceiella pniinosa differs from both B. glob- ulus and B. laevigata in ha\ing a teleoconch sculpture of minute granules and fine axial w rinkles. Etymology: Latin pruinosus (frosts), alluding to the fineK granulate surface. Subclass HETEROBRANCHIA Grav, 1840 Order HETEROSTROPHA Fischer,' 1885 Family XYLODISCULIDAE Waren, 1992 Genus Xylodiscula Marshall, 1988 Xylodiscula Marshall, 1988:988. T\pe species (by original des- ignation): Xylodiscula vitrea Marshall. 1988; Recent, New South Wales. Remarks: Xylodiscula was originalK referred to Or- bitestellidae for want of a more appropriate position (Marshall, 1988). Subsequent reevaluation of Orbitestel- lidae by Ponder (1990), however, suggested that this placement is untenable, and VVaren (1992) has segre- gated Xylodiscula in a family of its ov\n. The new species described below is the first record of a xylodisculid from whale bone. Other Xylodiscula spe- cies live at 90-1,100 m in association w itli sunken wood (Marshall, 1988; Waren, 1992) and old sea grass (Posi- donia) fibres (Waren. 1992), Most recentK a Xylodiscula species has been recorded from a h\drothermal vent at 2,000 m depth in the North Fiji Basin (Waren & Bouchet, 1993), Xylodiscula osteophila n sp (figures 15-17, 24, 25) Description: Shell (holotspe) 1,60 mm wide, markedly w ider than high, spire weakly elevated, thin, translucent, colorless, umbilicate, periostracum smooth. Protoconch 270 /um wide, 1.75 whorls; tip infolded, ver\ small, finely granulate; last w horl smooth Teleoconch of 2.25 convex whorls, suture shallowly channelled. First 1.25 whorls evenly convex; last whorl w ith weakly convex side, and strongK rounded periph- ery. Base convex, evenly rounded into umbilicus, obscure spiral lines throughout. Collabral grow th lines geiUK pro- socline on spire, weakly sigmoidal on base. Umbilical diameter 17''t of shell diameter. Aperture subcircular. Outer lip thin; inner lip thickened and angled against umbilical rim, thin adapically. Parietal area broad, in- ductura extremely thin, ■Animal unknown (dried). Operculum thin, translu- cent, chitinous multispiral, Radula (figures 24, 25) with the formula 2 -I- 1 4- 0 + 1-1-2, extremeK small. Lateral teeth small, thin, subquadrate, cutting area almost straight and finely ser- rate. Marginal teeth similar, large, stout, slender, curved, \ er\ long fineK' serrate cutting area on both edges. Type data: Holot\ pe (height 1,07 mm, width 1,60 mm, 2,25 teleoconch whorls) MNZ M, 116971 off Mernoo Bank, Chatham Rise, New Zealand, ca. 900 m, October 1988, f,v, Amaltal Explorer; alive on large w hale skull amongst crowded m>tilids (Idas sp.). Distribution (figure 1): Off Mernoo Bank, Chatham Rise, New Zealand, on v\hale bone, ca. 900 m. Remarks: Xylodiscula osteophila differs from other named Xylodiscula species in the greater shell height relative to width, the thickened and angled inner lip, and the considerabK narrower umbilicus. Etymology: From the Greek osteon (bone) and philios (loving). ACKNOWLEDGEMENTS I am grateful to the numerous fisheries scientists and observers (MAF Fisheries, Wellington) who obtained the samples of whale bone, to .A. Grebneff (I'niversity of Otago) who drew my attention to the fossil osteopeltid, and to J. H. McLean (Los Angeles Count) Museum of Natural History) for constructive comments on the manuscript. Thanks also to W. St George (Institute of Geological and Nuclear Sciences, Low er Hutt) for access to the scanning electron microscope, D. Wilkinson for w ord processing, and to T, Meek for photographic print- ing' Figures 18-2,5. Radulae ex holotypes. 18. 19. Ostcopclta praeceps n, sp,, full v\idth (18) and detail of central and lateral teeth (19) 20-22. Brticciclla laevigata n, sp,, full width (20), central and lateral teeth (21) and marginal teeth (22), 23. Bruceiella pruirjo.'ia n sp , central (left), lateral and inner marginal teeth. 24, 25. Xylodiscula osteophila n, sp,, marginal teeth and the small central tooth (left center in 24), Scale bars = 10 ^m. Pages THE NAUTILUS, Vol. 108, No. 1 LITERATURE CITED Dell. R, K. 1956. The Arcluhenthal Mollusca of New Zealand. Ddiiiinion Museum Bulletin l.S:l-234. Dell, R. K. 1987. Mollusca of tlie famih Mytilidae (Bivalvia) associated with organic remains from deep w ater off New Zealand, with revisions of the genera Adipicola Dautzen- berg, 1927 and Idasola Iredale, 1915. National Museum of New Zealand Records 3:17-36. Fleming, C. A. 1948. New species and genera of marine Mollusca from the Southland fiords. Transactions of the Royal Society of New Zealand 77:72-92. Haszprunar, G. 1987. .\natomy and affinities of cocculinid limpets (Mollusca, Archaeogastropoda). Zoologica Scripta 16:305-324. Haszprunar, G. 1988. .\natom\ and relationships ot the bone- feeding limpets CoccuhncUa minutissima (Smith) and Os- tcopclta mirabilis Marshall (.\rchaeogastropoda). Journal of Molluscan Studies 54: 1-20. McLean, J, H 1992. Cocculiniform limpets (Cxicculinidae and P\ ropeltidae) living on whale bone in the deep sea off California. Journal of Molluscan Studies 58:401-414. McLean, J. H. and G. Haszprunar. 1987 P\ ropeltidae, a new familv of cocculiniform limpets from h\drothermal \ents. The \'eliger 30:196-205. Marshall. B. .\. 1986. Recent and Tertiary Cocculinidae and Pseudococculinidae (Mollusca: Gastropoda) from New Zealand and New South Wales. New Zealand Journal of Zoology 12 505-546. Marshall. B .\ 1987 Osteopeltidae (Mollusca: Gastropoda): a new family of limpets associated with whale bone in the deep-sea. Journal of Molluscan Studies 53121-127. Marshall. B. A. 1988, Skeneidae. Vitriueliidae and Orbites- tellidae (Mollusca: Gastropoda) associated with biogenic substrata from bathyal depths off New Zealand and New South Wales, Journal of Natural History 22:949-1004. Ma,\well, P, \. 1992. Eocene Mollusca from the vicinity of McCulloch's Bridge, Waihao River, South Canterbury, New- Zealand: paleontology and systematics. New Zealand Geo- logical Survey Paleontological Bulletin 65:1-280. Ponder, W. F, 1990, The anatomy and relationships of the Orbitestellidae (Gastropoda: Heteroliranchia). Journal of Molluscan Studies 56:515-532, Powell, A. W, B. 1979. New Zealand Mollusca: marine, land and freshwater shells, Collins, .Auckland. 500 p. Smith, C. R. 1992. Whale falls. Chemoss nthesis on the deep sea floor. Oceanus 35:74-78. Smith, C. R.. H. Kukert. R. A. V\ heatcroft. P .\ Jumars, and J, W. Deming. 1989. N'ent fainia on u hale remains. Na- ture 341:27-28. Waren, A, 1989, New and little know n Mollusca from Iceland. Sarsia 74:1-28. Waren, .•\, 1992, New and little known "skeneimorph" gas- tropods from the Mediterranean Sea and the adjacent At- lantic Ocean, Bollettino Malacologico 27:149-247, Waren, A. and P, Bouchet, 1993, New records, species, gen- era, and a new family of gastropods from hydrothermal vents and hydrocarbon seeps, Zoologica Scripta 22:1-90. THE NAUTILUS 108(1):9-14, 1994 Page 9 On the Living Habits of Acesta biiUisi (Yokes) in Chemosynthetic Bottom Communities, Gulf of Mexico Barry kohl Harold E. Yokes tieulijg) Department Tulane University New Orleans, LA 70118 USA ABSTRACT The supposedly pathologic paratype of Acesta huUisi (Vokes,1963) has been re-evaluated based on additional infor- mation from the type locality The specimen is not aberrant, as originally thought, but represents a normal mode of growth In uliich the shell is attached over the anterior end of che- inos\ nthetic tube worms of ihe g,enus Luim'llihrachia sp., which occur at Indrocarbon seeps in the Gulf of Mexico, It is shown thai the paratype is a final stage in the ontogen\ of A. bullisi. which lives attached to the tube worm Lamcllihrachia. Key W'urds: Acesta, Chemos> nthetic, Lanicllilirucliia. CUill of Mexico, tube worms. INTf^ODUCTION Hydrothermal vents and their oases of previously un- described species were discovered in 1977 on dives of the submersible Alvin along the Galapagos Rift in the Pacific Ocean. Giant tube worms, large clams, and mus- sels form an assemblage of invertebrates dependent on hydrogen sulfide-rich waters. These animals contain bac- terial symbionts that o.xidize the sulfide. An assemblage of tube worms, clams, and mussels was reported in the Gulf of Mexico near a saline seep at the base of the Florida Escarpment (depth 3,266 meters) by Paull et al., (1984). This was the first record of vent organisms occurring on a passive margin. The following year, Kennicutt et al. (1985) described a fauna associated with oil and gas seeps from trawls along the Louisiana continental slope. One trawl contained specimens of Acesta bullisi and an entanglement of vestimentiferan tube worms of the genus Lamellibrachia , taken near Green Canvon Block 234, (27°45'N, 9ri4'W) (Figure 1). Since the discoveries by Paull ct al and Kennicutt et al.. there have been many projects designed to document, by manned-submersibles, the occurrences of chemosyn- thetic communities in the Gulf of Mexico associated with oil and gas seeps. Shore-based research is investigating how these organisins derive their energy from oil, meth- ane, and sulfides, which are toxic to most invertebrates. In addition to unicjue chemosynthetic species, there are opportunistic non-chemosynthetic organisms that use either the carbonate substrate surrounding the vents or the chemosv nthetic forms for attachment. Acesta bullisi is one of these opportunistic organisms found attached to the tube worm Lamellibrachia sp. at many sites in the Gulf of Mexico (Carnev, 1992; MacDonald & Ambler 1992). The supposedly "pathologic " hypotype [paratype] of the species described as Lima (Acesta) bullisi by Yokes (1963, pi. 2, figs. 1-4), was given to H. E. Yokes in 1962 by H. R Bullis, Jr. It was alive when retrieved in a trawl from the R/Y Oregon Station 3741, at a depth of 548 meters in the Gulf of Mexico (29°10'N, 88°01.5'W). This location can be equated to a sampling station in Viosca Knoll Block 826 (YK 826, Figure 1), 29°11.0'N, 88°00'W, at a depth of 545 meters (L' S. Dept of Interior, 1992, vol. 1, table 1) used by the Geochemical and En- vironmental Research Group (GERG) of Texas A & M University. According to MacDonald and Ambler (1992), this location is the most easterly site within the Gulf of Mexico of chemosynthetic communities associated with hydrocarbon seep activity found to date. The chemo- synthetic fauna at the YK 826 site is represented by living tube worms {Lamellibrachia sp. and Escarpia sp) and ]X)ssibl\- lucinid clams (MacDonald & Ambler, 1992). The purpose of this paper is to document that the paratype of Acesta bullisi described by Yokes in 1963 was a form attached to tube worms from a chemosyn- thetic commimity. We hope that this paper will also stimulate an investigation of other "deformed mollusk species in collections and thereb\ identify locations of other chemosynthetic communities. MATERIALS AND METHODS Specimens of Acesta bullisi were collected by personnel from Tulane University and Louisiana State University (LSL') on cruises in the Gulf of Mexico during September 1992 and July 1993. The Johnson Sea-Link (JSL) I & II, manned-submersibles, were used for bottom sampling on a total of 24 dives. Photographs and video were taken Page 10 THE NAUTILUS, Vol. 108, No. 1 Figure 1. Location map showing Garden Banks Block 427, Green C'anvon Blocks 18.5, 234 and Viosca Knoll Block 826. of clumps of Lamellibrachia and the attached individuals of Acesia. Specimens of Acesta bullisi used for the pres- ent study were collected for research at LSI' and were photographed on shipboard by B. Kohl to document the living position of Acesta in relationship to the tube worm Lamellibrachia. A collection of invertebrates retrieved on the dives is maintained at LSU. The sites used for this paper are Green Canyon Block 185, 27°46.93'N, 91°.30.47'W, dive 3301 (1992), depth 541 meters, and Garden Banks Block 427, 27°33.61'N, 92°25.09'\V, dives 3305 (1992) and 3568 (1993), depth 611 meters, (Bgure 1). Ten specimens of A. bullisi at- tached to Lamellibrachia were retrieved by the JSL's mechanical arm by selectively collecting individual tube worms. RESULTS Biology of Tube Worms Lamellilnachia spp., classified as vestimentiferans, lack a mouth, anus and digestive system. They live in a tough, thick-walled, chitinous tube about one centimeter in di- ameter and up to two meters in length. The posterior end may either be buried in soft sediments or attached to a rocky substratum. A small red-orange plume, the obturaculum, (approx. one cm in diameter) is extended from the anterior opening and serves as a gas exchange organ. The plume can be withdrawn into the tube at will (Fisher, 1992; MacDonaid et a/., 1990). Most of the body volume consists of a trophosome, the organ containing the bacterial symbionts. The animal absorbs HjS, CO2 and O2 through its obturaculum (plume) and, through a complex chemical process, the symbiotic bacteria and enz\ nies convert CO2 to organic carbon. See Fisher (1990) for a detailed review of the chemistry involved. New research has suggested that the O2 is absorbed through the plume and H2S is absorbed through the posterior part of the tube, which is located below the sediment,' water interface. It has been observed that the living animal extends the entire length of the tube. Ab- sorption of CO2 possibly takes place along the length of the tube above the sediment/water interface through the porous chitinous wall (R. S. Carney, personal commu- nication). Relationship of A. bullisi to L.\mellibrachl\ sp. Specimens of A. bullisi attach to tall individuals of La- mellibrachia approximately one-half meter above the sea bottom, near the anterior end of the tube. Specimens collected on the 1992 and 1993 dives show that A. biillisi frequently occurs attached to the "bushlike ' congrega- tions of the tube worm Lamellibrachia sp This was also reported and illustrated by MacDonaid £>/ al. (1989) from the Green Canyon Block 184-185 area. Two attachment positions of A. bullisi were observed on our dives. The first is an attachment by a b\ssus to the side of the tube worm near the upper portion of the tube (figure 3). These individuals do not have the notch or opening and are about half the size of the adult shown in figures 2 and 4. The second attachment position can be seen in the specimens in figure 2, which completely envelop the anterior region of Lamellibrachia. A detail of one of the specimens from figure 2 is shown in figure 4. These specimens are attached by a byssus to the exterior of the tube worm very near the anterior end. The shape of the growth lines in the valves of the adult A. bullisi (figure 9) implies that the opening in the shell developed late in ontogeny. DISCUSSION The specimen described by H. E. Yokes was thought to have been damaged at a time when the animal was half grown (Yokes, 1963; 78). Observations of the living an- Figure§ 2— t. Living specimens of Acesta Imllisi 2. Live specimens of A. bullisi attached to the anterior ends of two Lamcllilyrachia sp . (Jarden Banks Block 427, dive .'5.305, depth (ill nicters. 3. A live specimen of .A, Imllisi from Green C:anyon Block 185, dive 3.301, deplli 541 meters This specimen is attached by a byssus 32 mm i>el()w (lie anterior enil of the tube worm and lacks the notch in the anierodorsal margin as seen in ligiires 4 and 9. ll is about 50 mm in height which is half tlie size of the specimen on left in figure 2. 4. Detail of the live specimen on the right in ligure 2 The specimen is approximatcK , 1 10 mm in height and surrounds the anterior portion of l.unwUihrarbia sp Note the position of the tuiie worm at center of photograph with the plume of the tube worm protruding Irom the anterior end (arrow) and its relationship to the notch When the plume is extended, during life position, it is totally contained within the mantle cavity of A. bullisi The i)yssus holds the A. Imllisi in a position to envelope the lube worm Specimen from Garden Banks Block 427, dive 3305, depth (il 1 meters. Scale in upper Icll Is in centimeters B. Kohl and H. E. Yokes, 1994 Page 11 5 cm *4 %. N .vt. // p ^^^^^^^^^^^m ^^^1 ^^^^^^^ ^^^^^H ~ Page 12 THE NAUTILUS, Vol. 108, No. 1 Figures 5-9. Specimen of /\. Imllisi eolleclecl alive Iroiii llie anterior of a lube worm. Green Canyon Block 185, dive 3301, 541 meters. Magnifieation (X 1). This specimen is comparable in size to the one figured 1)\ N'okes (1963, pi 2, figs, 1-4). Note the B. Kohl and H. E. Yokes, 1994 Page 13 imals retrieved and photographed (figures 2-4) (hiring recent dives of the JSL (Sept 1992 and JuK 1993) have shown that the specimen figured h\ \ dkes was actualK an ecophenotypic variant that grows attached to the tube worm LamcUihrachia sp^ The inlienchng of the antero- dorsal margin of A. btillisi and the de\elopment of an open gape is the result of its attachment to the anterior end of LamcUihrachia sp. Tliis has been reported from specimens collected at Viosca Knoll Block 826 (Mac- Donald & Ambler, 1992: 46) Yokes (1963: 78) suggested that "the specimen, uhich was alive when taken, had suftered serious damage to the shell and mantle in the region of the lunule at a time when the animal was approximately half grown." With- out the know leilge that the specimen was originally at- tached to a tube worm, this was a logical interpretation. We now know that inbending of the anterodorsal margin and open gape is a normal development for specimens growing on the anterior end of LamcUihrachia (Mac- donaldc? (;/,,1989). Carney (1992) reports that specimens of A. hiilh.si with thin shells are either attached to car- bonate rocks or swimming freely near the vent com- munities v\hereas the forms attached to LamcUihrachia have a distinctk thicker shell. We suggest that smaller specimens of A. buUisi ma\ move from an attacfied position on the side of the tube worm after achieving apprcximately one-half adult size, and then to an anterior location during the later stages of life where the gape is developed as it closes over the tube worm. Members of the genus Acesta can resorb their byssus filaments. Once unattached they can swim to another position, near the anterior end of the tube worm, and re-attach. Since LamcUihrachia has neither mouth, gut nor anus and is completely dependent on sulfur-reducing bacteria for its food, it is difficult to un- derstand the ecological relationship between A. huUisi and the tube worm. Could there be a symbiotic or com- mensal association between the two organisms? Acesta huUisi does not have symbiotic bacteria in its mantle and appears to be a normal filter feeder. Addi- tional research on the biology of A. hitUisi. being con- ducted by C. R Fisher at Pennsylvania State Univ. (R. S. Carney, personal communication), may answer some of these cjuestions. The description of the "pathologic" specimen of A. huUisi by Yokes (1963), collected at Yiosca Knoll Block 826, was the first record of a mollusk from an oil seep- chemosynthetic community in the Gulf of Mexico. Care- ful study of the faunal collections dredged at YK 826 in 1962 by the R/Y Oregon would probably reveal the existence of tube worms. Subseciuent dives by GERG in 1991 have documented the occurrence of LamcUihrachia at YK 826 (MacDonald & Ambler, 1992). Based on our present knowledge of the occurrence of A. hiiUisi in chemosynthetic communities, there is no doubt that the paratype of A. huUisi described l)y Yokes from YK 826 was a form attached to the anterior end of a specimen of LamcUihrachia. ACKNOWLEDGMENTS The authors thank Harry H. Roberts, director of the Coastal Studies Institute at Louisiana State University and the chief scientist on the cruises of the R/Y Edwin Link for Septemlter 1992 and the H/ Y Seward Johnson for July 1993 dives. Also, thanks go to Robert Avent of Minerals Management Service, Department of Interior for providing background material on the chemosyn- thetic faunas and to James M Rrf)oks and Ian R Mac- Donald of GERG for providing information on Acesta huUisi occurrences in the Gulf of Mexico. We also thank Robert S. Carney for the loan of a specimen of Acesta huUisi from the LSU collection for photograph} . Emily H. Yokes, of Tulane LIniversity, pre- pared the plates and reviewed and edited versions of the manuscript. Marco Taviani, Institute di Geologia Marina, Bologna, ItaK, who was on the 1992 cruise, assisted with the shipboard photograph) The authors thank two anon- \ mous reviewers for their suggestions to improve the paper. The dives were supported by the Louisiana Sea Grant Program, New Orleans Office of Minerals Management Service, National Oceanic and Atmospheric AgencN and the National L^ndersea Research Center at the University of North Carolina at Wilmington. LITERATURE CITED Carney, R .S 1992 Heterotrophic Fauna Associated with CheiiiDsyiitiietic Communities. In: MacDonald, I. (ed.). Technical Report of Northern Gulf of Mexico Chemosyn- thetic Ecosystems Study: Literature Review and Data Syn- thesis, vol. 2, V. S Department of Interior, Minerals Man- agement Service, Gulf of Mexico OC'S Region, New Oriean.s, LA, Chapter 9, p. 1-30 Fisher, C. R. 1990 Chemoautotrophic and Methanotropliic Symbioses in Marine Invertebrates- Reviews in .Aijuatic Sciences. 2 (3-4): 399-436. l-'isher, C:, R. 1992. Physiology. Biocl)emistr>, and Ecology of Vent and Seep Invertebrates with Chemoautotrophic or Methanotrophic Symbionts. In: MacDonald, I. (ed). Technical Report of Northern Gulf of Mexico Chemosyn- thetic Ecosystems Study Literature Review and Data Syn- thesis, vol. 2, U S Deparlmenl of Interior, Minerals Man- agement Service, Gulf of Mexico OC^S Region, New Orleans, LA, Chapter 7, p. 1-17. Kennicutt. M C. II, J. M. Brooks, R R Bidigare, R. R. Fay, T L. Wade and T. J. McDonald. 1985. Vent-type taxa in a hydrocarbon seep region on the Louisiana slope. Na- ture (London) 317(603.5): 351-353. position ol die notch, (F'igs. 5,6,9) vviiitli allows A. inillisi to close over the tube worm. This is also evident in Yokes (1963, pi. 2, figs. 2-3). Specimen on loan troni LSLI. Page 14 THE NAUTILUS, Vol. 108, No. 1 MacDonald, I. R and J .Ambler. 1992. Regional distribution of C^heniosynthetic Fauna and Community Ecology hi: MacDonald, I. (ed.). Technical Report of Northern Gulf of Mexico Chemosynthetic Ecosystems Study: Literature Review and Data Synthesis, vol.2, U S Department of Interior, Minerals Management Service, Gulf of Me.xico (X;S Region, New Orleans, LA, Chapter 8, p. 1-53 MacDonald, L R., G. S. Borland, J. S. Baker, J. M. Brooks, M. C. Kennicutt and R. R. Bidigare. 1989 Gulf of Mexico chemosv nthetic communities II; Spatial distribution of seep organisms and hydrocarbons at Rush Hill Marine Biology KM: 235-247, MacDonald, I R , N L. Guinasso, Jr , J. F. Reilly, J M. Brooks, W R Callender, and S. G. Gabrielle. 1990. Gulf of Mexico Hydrocarbon Seep Communities: VI. Patterns in Community Structure and Habitat, Geo-Marine Letters, 10: 244-252 Paull, C K , B Hecker, R. Commeau, R.P Freeman-Lynda, C Neumann, W.P. Corso, S. Golubic. J. E. Hook, E. Sikes, and J Curray 1984. Biological communities at the Flor- ida Escarpment resemble hydrothernial vent taxa. Science 226: 965-967. U. S Department of Interior, Minerals Management Service. 1992. Northern Gulf of Mexico Chemosynthetic Ecosys- tems Study: Literature Reyiew and Data Synthesis. Vol. 1, Executive Summary, 32 pp.; Vol. 2, Technical Report, 238 pp.; Vol. 3, Appendix, 243 pp.; Edited by Ian Mac- Donald, Geochemical and Environmental Research Group, Texas A &M University L' S Dept. of Interior, Minerals Management Service, Gulf ot Mexico OCS Regional Of- fice, New Orleans, LA, Vokes, H, E 1963 Studies on Tertiary and Recent Giant Limidae, Tulane Studies in Geologyl(2): 75-92, 2 pis. THE NAUTILUS 108(l):15-22, 1994 Page 15 Shell Alignment for the Morphometric Analysis of High-spired Gastropods. Paul J. Morris' Smithsonian Tropical Research Institute Apartatdo 2072 Balboa, Republica de Panama Warren D. Allmon Palenntological Research Institution 1259 Trumansburg Rd. Ithaca, NY 14S50 USA ABSTRACT .\ significant problem often impedes the morphometric anaKsis of high-spired gastropods. It is usualK' not possible to obtain large samples of complete specimens. Without the ontogenetic landmarks pro\ ided b\ earl\ u horls it is difficult to align in- complete specuiiens to compare homologous w horls. We discuss several methods that allow alignment of morphometric data taken from incomplete specimens of high-spired gastropods. We evaluated these methods b\ treating a set of complete specimens as incomplete. Two methods produce reliable align- ments. One of these uses an ontogenetic function of the whorl expansion rate to estimate missing whorls, the other aligns whorls of similar heights. Both produce reasonably precise and accu- rate alignments, but under different assumptions. Once a set of specimens has been aligned, it can be subjected to conven- tional statistical anaKsis using accepted coinentions to deal w ith missing data Key words: Morphometries. Gastropoda. Missing Data. Tur- ritellidae. INTRODUCTION Morphometric anal\ ses of very high-spired gastropods (such as members of the families Cerithiidae, Lo.xone- matidae, Murchinsoniidae, Nerineidae, Turritellidae, and Terebridae) are difficult because complete specimens are usualK unavailable or rare. This is especiaJK true for fossil taxa. When the juvenile whorls ha\ e been broken off of most specimens it is difficult or impossible to specif \ homologous points (i.e.. similar whorl number) among specimens. Therefore, construction of data matrices for morphometric or statistical anaKses of a large number of specimens of such incomplete gastropods is usualK problematic. This difficultv may account for the paucitv of morphometric studies of high spired gastropods in the literature (e.g.. .^llmon, 1994). ' Current Address: Paleontological Research Institution. Trumansburg Rd . Ithaca. \y"i4S50 USA 259 A variety of methods can be used to align the whorls of incomplete specimens w ith each other \\'e have im- plemented three algorithms for use on sets of traditional point-to-point measurements. We assessed the accuracy of these algorithms by their ability to predict true whorl number in a data set of complete specimens that we treated as incomplete. In this data set, we truncated complete specimens at known whorl numbers. Two al- gorithms produced good alignments of shells of turritel- lids in this test data set. Both of these align most speci- mens to within one whorl of their true position. An algorithm that uses iterative comparisons of whorl heights produces slightly more precise alignments. .\n algorithm that uses an ontogenetic function of the whorl expansion rate to infer numbers of missing whorls is slightly more accurate. There are several tradeoffs between these two algorithms. SHELL ALIGNMENT In most gastropods it is possible to identify a homologous point for aligning whorls. Either the end of the proto- conch or a varix produced in the adult shell can be used as landmarks from which whorls can be counted. It is therefore easy to align whorls that are of the same dis- tance from this landmark in different specimens. Such aligned specimens are easiK compared using traditional point-to-point measures, Raupian parameters, or moving frame parameters. In most high-spired shells, however, whorls differ only in size and shape, and lack discrete landmarks. When specimens have been broken and are incomplete, there are no clear means for inferring how- many whorls are missing. Incomplete specimens, which are particularly common as fossils, are therefore difficult to align, homologize, and anaK ze morphometricalK . This problem is depicted in Figure 1 . There are three basic ways of aligning incomplete high spired shells. These are: alignment by homologv of or- nament, inference of missing whorls from whorl expan- sion rate (W of Raup, 1966), or alignment by compar- isons of whorl heights among specimens. (One can also Page 16 THE NAUTILUS, Vol, 108, No. 1 l-i-J..V_ ' \1 Figure 1. The problem of v\horl alignnieiU in two incomplete specimens of high spired gastropods. In tlie alisence of the protoconch, juvenile w horls, or other homologous points, it is necessar) to find some means of aligning w horls in order to conduct mnrphometric anaKses The two principal \va>s of making alignments are by matching whorls of similar size, or by inferring the number of whorls missing from the small end of each specimen. use other measures that increase through ontogeny, such as \\ horl width, but for the remainder of this paper we refer exclusiveK to whorl height). We ha\e considered five variations of these methods, alignment by: 1) com- parison of homologous features of ornament, 2) inter- polation of missing whorls using the wliorl expansion rate from earliest preserved whorls of each specimen, 3) in- terpolation of missing whorls using some function of W calculated from a composite average specimen, 4) com- parison of whorl heights between each pair of specimens, and 5) iterative correction and comparison of whorl heights with a composite a\erage specimen .\ method that uses iterati\'e comparisons of whorl heights appears to be the most generalK useful. None of these, however, are entireK perfect \\a\s to infer how man> whorls are missing in an incomplete specimen. There is no ideal way to align indi\ idual specimens that lack earh whorls and homologous end points. Each of these approaches has limitations and makes compromises. .■\ first approach is alignment using homologous fea- tures of spiral ornamentation, such as the points of bi- furcation or origination of keels. This approach (because it involves homologous points) w ould seem most reliable. It is limited, howe\er, by an inability to extend homol- ogies of ornamentation be\ond small groups of related species, and b\ the need for man\ homologous points spread throughout ontogeny. As a result, it is useful only in a few specific cases. It does not lend itself to imple- mentation as a general algorithm that could appK , for example, to both turritellids and loxonematids. It also must avoid the risk of making arbitrary assignment of homologous points in contiiuiousK growing ornament. .\ second approach is alignment using w horl expansion rates to inter the ninnber ol missing w horls between the protoconch and the first [^reserved whorl in each speci- men. This could be done 1)> first calculating the whorl expansion rate from the lirsl preserved whf)rls iil each specimen. This parameter can then be used to calculate how main w horls are missing between the first preser\ed w horl and some smaller first whorl (e.g.. the protoconch, or the smallest preser\ed whorl in the data set). That is, iterativeK reducing the height of the smallest preserved whorl of a specimen b\ its whorl expansion rate. The number of steps required to reduce this height value to below the protoconch size provides an estimate for the number of missing whorls. The advantage of this method is that the estimated number of missing whorls in each specimen is independent of the other specimens in the sample. However, both measurement errors in the small- est whorls and ontogenetic changes in the w horl expan- sion rate affect these alignments. These errors can be reduced by using some ontogenetic function of the whorl expansion rate rather than an un- changing W \ alue. This must perforce be calculated from other specimens in the sample. It is not possible to infer from a single specimen how W might ha\e \ aried in its missing whorls. Ideally, this ontogenetic huiction oi W w ould be continuousK \ ariable, reflecting the real nature of ontogenetic changes in w horl expansion rate. In man\' of our samples, however, few specimens preserve the smallest whorls in the data set. It is therefore necessary to increase the sample size, and decrease the resolution of ontogenetic changes in W. We feel that three whorl expansion rates, produced by a di\ision of the data set (the composite specimen) into three parts (smallest 'i, second 'i, largest 'A), pro\ide the best compromise be- tween resolution of W and sample size. Thus, a whorl expansion rate is calculated for each pair of whorls in each specimen, and an average W is calculated from the whorls in each part of the specimen. These three W \alues are then used to infer how many w horls are miss- ing from each specimen. For example, consider a specimen in w hich the small- est preserved whorl falls within in the second U of the range of whorl sizes in a sample. The number of missing w horls in this specimen are calculated b\ iterati\el\' re- ducing the height of its first preserved w horl. While this height value remains within the second 'j, the average W V alue from the second ' j is used to calculate the height of the next smaller w horl. When this value falls into the range of heights of the smallest '4, the average W value from the smallest U is likewise used to fill in smaller wliorls. In each specimen, the number of steps required to reduce the height of the smallest preserved whorl to less than the height of the smallest w horl in the data set (or a protoconch height) are an estimate ot the nutnber of missing w horls. .\ third possible approach is the aligmnent of w horls of similar height. This is the most intuitive approach, and the one we find ourselves naturalK taking when pliv sicallv lining up specimens next to each other. There are two obv ions vv av s to carry out these aligmnents; each starts bv sorting the specimens in order of increasing size of first whorl. Specimens mav' then be aligned bv pairwise comparisons or bv constructing a composite average specimen and itcratlvcly locating and moving poorly aligned specimens, .\liginnent by height, however, offers p. J. Morris and W. D. Allmon, 1994 Page 17 several computational problems, Pairwise c can affect alignments in several ways. Such \ arialiilit\ can produce a broad spread of alignments, or it can produce non- intuitive systematic missalignments. It is essential that 4 11 4 11 4 True Whorl humber Figures 2 — I. .^ccuracN and precision of tiiree methods ot \\ horl alignment discussed in this paper. Whorl heights uere mea- sured in ten specimens of Tiirritella gonostoma and used to construct a data set of 50 truncated specimens containing true whorl numbers and whorl height. Open circles are aligned whorl number, filled circles are plus and minus one standard deviation. Lines represent correct whorl alignment and error of plus and minus one whorl. 2. Alignment from Ju\enile VV 3. .Alignment using three W \alues, 4. .Ahgnment b\ Height. .Alignment from Juvenile \V is neither accurate nor precise. The other twci metlicKls are suhstantialK laetter at correctK aligning specimens in this test. the user examine aligned samples to understand how this variabilitx may have affected the alignments. Rotation error is the result of taking a set of measurements from one plane in a continuousK growing spiral shell. In our test data set it is the principle source of reduced precision for the tw o best alignment methods. Effects of this source of error can be reduced In either using three dimensional coordinates and mo\ ing frame parameters y\ckerly, 1989; 1990; Johnston et ai. 1991), or b\ aligning specimens before measurement (rotating each specimen until a par- ticular whorl width (or height) lies in the plane of mea- surement). The relationships among size, age, and homologx' pro- vide a subtle set of philosophical problems. Age, growth rate, size, and shape are complexly interrelated in gas- tropods. Environmental factors, such as the availability of food, readily influence the growth rates of gastropods. In time averaged samples it is entireK possible that w horls of the same size (and the same true whorl number) in different individuals represent different points in their ontogenies. In most paleontological studies, how ever, the primary goal is simply to assess how shape and size vary within and between samples Patterns in the data can then constrain various taxonomic, evolutionary or pale- oecologic hypotheses. The two most common questions for w hich detailed morphometric studies are undertaken are species discrimination, and trends in variability tlirough time [i.e.. tempo and mode of evolution). Lines of ev idence beyond morphologv', such as taphonomv', are needed to assess the role ol ecophenotypic factors in increasing variance or creating clusters in the data. Page 18 THE NAUTILUS, Vol. 108, No. 1 Table 1. Comparison of advantages and pitfalls of three methods of whorl alignment. These methods are; Alignment b> Height using iterative comparison with a composite average specimen, Alignment using three W values computed from composite average specimen, and .Alignment using VV calculated from smallest three whorls of each specimen. Alignment by Height is the most suitable except for cases where 1) many specimens are being aligned and computing time is a concern, or 2) many specimens consist of just two or three whorls. If the data are three dimensional coordinates suitable for moving frame analysis we expect that a modification of the Height algorithm will probabK produce the best alignments .'Alignments of a test data set (complete specimens treated as incomplete) suggest .\lignment by Height as the best overall technique. .Alignment b\ Height HISPIRE .3 Three W values HISPIRE 2 Juvenile W HISPIRE 1 Specimens aligned using: .Advantages; Speed Alignment Disadvantages: Speed .Alignment Systematic Missalligment; Dependence on sample composition; Recommended for; Most applications Many specimens Few whorls Principle sources of alignment error; Measurement error Ontogenetic del W Population del W Few whorls Composite Composit Fast Precise Accurate Slow Some None Yes Yes + + + + + + + + + + + + + + + + + + + Self Fast Inaccurate Much No + + + + + + Test data set, Tiirritella gonostoma Average of SDs; Relative precision of alignment Small whorls Middle whorls Large whorls 0.93 high high moderate 1.12 high moderate moderate 1.73 low low low ASSUMPTIONS A fundamental assumption follows from the previous paragraph: The questions under consideration can be tested with information about size and shape without reference to age (see discussion in Janie.s, 1988), Beyond this, each alignment method makes its own set of as- sumptions about the specimens that are to be aligned. Both the accuracy of alignment and the assumptions behind the methods affect their suitability for particular analytic problems. There are three principal assumptions made by the method of alignment from juvenile W val- ues. It assumes that there is little measurement error in the early whorls. In addition, it assumes that the w horl expansion rate is constant throughout ontogeny. Whorl number is expected to correlate with the height of the first preserved whorl. The method of alignment using three W values allows for ontogenetic variation in the whorl expansion rate. However, it is still sensitive to errors in measurements of the earliest preserved whorl. In addition, this method takes on the assumption that there is little population variability in the w horl expan- sion rate. .Alignment by height assumes that whorls of similar heights are homologous. It does not directly as- sume that there is little population variabilit> in the whorl expansion rate, but if this exists, it may produce non-intuitive alignments. TESTS OF ALIGNMENTS We ha\e tested the accuracy of these three methods of w horl alignment b> evaluating their ability to correctly align whorls in a sample of complete specimens that we treated as incomplete. We used ten complete specimens of Tiirritella gonostoma Valenciennes, 1832, a Recent turritellid that undergoes relatively large ontogenetic changes in w horl expansion rate. We photographed these specimens, enlarged the images, and digitized points at the suture of each whorl using a Numonics digitizing tablet. The heights of successive v\ horls were calculated from these digitized points. We then truncated the data for each specimen at five different whorls spanning the region of greatest change in whorl profile. This procedure provided us w ith a data set of .50 incomplete specimens p. J. Morris and W. D. Allmon, 1994 Page 19 s E .•QO o True Whorl Number Alligned Whorl No. Figures 5, 6. Assessment of accuracy ul w liw I uligimient b> iterative comparisons of whorl height with composite average specimen. 5. Comparison of true and ahgned whorl number. Open circles are aligned whorl number, filled circles are plus and minus one standard deviation. Lines represent error of plus and minus one whorl from correct whorl alignment, 6. Whorl heights and aligned whorl number Lines connect the whorls of individual specimens Note that the accuracy of this method is strongly affected by the alignment of the two smallest specimens in the data set. Elimination of the two smallest aligned whorls will result in substantial uTipro\ement in accurac> . of various numbers of whorls, for wliicli whorl height and true whorl number w ere known in each whorl. This data set was entered into three conversion programs (HISPIRE, HISPIRE2, HISPIRE3, see Appendix 2) that implement the three whorl alignment algorithms. Figures 2-4 illustrate the different alignments of this data set b\ the three programs. These figures are graphs of true whorl number against aligned whorl number. The mean alignments for these 50 specimens (plus and minus one standard de\iation) are compared with a cor- rect alignment (plus and minus one whorl). Clearly align- ment by ju\enile VV performs least well in this test. On careful examination of the results, both the ontogenetic changes in whorl expansion rate and measurement error contribute to its failure. The other two alignment methods produce substan- tially better results. Here, alignment using three W values produces more accurate results than alignment by height, .alignment using three W values places the mean align- ment for each whorl slightly closer to the true whorl value than alignment by height. However, when the standard deviations in alignment in each v\ horl (exclud- ing the first and last whorls in each case) are averaged, alignment b\ height is more precise (Table 1). It appears that alignment by height is slightK' more precise (tighter clustering) and that alignment using three W values is slightK- more accurate (closer to true). In order to understand the quality of alignments, it is important that we consider the reasons for these differ- ences. The sources of these errors are revealed in an examination of plots of true against aligned whorl num- ber, whorl height against true whorl number (e.g.. Fig- ures 4,5), and the original data. Alignment using three W values is subject to misalignments produced by mea- surement errors in the early whorls. Such measurement errors, along with slight rotation errors and variation in ^^', tend to produce misalignments in steps of one whorl. L'sing this method, some specimens are well aligned, others suffer quantum misalignment. In contrast, align- ment by height produces misalignments that appear to be the result of rotation error. All specimens are tightly aligned with each other (Figure 6) and their spread is some combination of true variation and alignment error. Careful examination of the graph reveals that the lower accuracy of this method is also the result of rotation error. The two specimens that have the smallest whorls in the data set happen to be aligned to the left relative to the others. The alignment of these two specimens is the cause of the reduced accuracy of alignment of the rest of the specimens. A remecK suggests itself at once. Removal of early whorls represented by only a few specimens should make alignment by height the most accurate and precise of the three methods. To test this contention further, we qualitatively eval- uated these programs. We examined the alignments they produce in fifteen samples of four species of Paleocene and Eocene turritellids from the U.S. Gulf coastal plain (Turritella mortoni Conrad, 1830, T. carinata I.Lea, 1S33, T. perdita Conrad, 1S65, and T. praecincta Con- rad, 1864). .almost all the specimens in these samples are incomplete. It is therefore not possible to compare the alignments to true values. Here we assessed the precision of alignments b\' graphing aligned w horl number against w horl height. In all the samples we examined, alignment b\ height consistentK' produced tight clusters. The results shown in Figures 7,8 are typical. In contrast, alignment using three W values often placed one or more specimens outside of a cluster of aligned specimens. This again reveals a weakness in this method of alignment. Small errors (in the measurement of the first preserved whorl, or in fit of the specimen to the three W values) can easily be magnified to misalignments of one or more whorls. Page 20 THE NAUTILUS, Vol. 108, No. 1 Alhgned WhorJ No. o All igned Whorl No. Figures 7. 8. Whorl alignments in 7. eight specimens of Tiir- rilella rnortoni from the Paleocene Aquia Fm. at Piscatavva> C:reek Prince Georges Co., MD (Appendix 1) and 8. twelve specimens of Tiirritella carinata from the Eocene Gosport Sand at Clayfjorne Bluff, Monroe Clo.. Alabama, .\ligned u liorl num- ber is plotted against whorl height. Lines connect the whorls of individual specimens. These alignments are t> pical for the samples of fossil turritellids that we examined. CHECKING ALIGNMENTS Given the assumptions of these algorithms and the va- garies of real data, the alignment programs cannot be treated as black boxes that produce infallible results. It is essential that the user e\aluate the accurac)' of the aligruTients. The programs themseUes pro\ide some warnings, but graphs of aligned \\li pe O ash. and is, therefore, only slightly younger than the KX-145 locality. Mammals were di- verse, and of a strongly boreal character (Voorhies & Corner, 1991 ). Analysis of arboreal pollen confirmed that a boreal, almost taiga-like, environment existed here (Bolick, 1991), The molluscan fauna from this locality is quite similar to that of Leonards t\ pe locality. Exami- nation of the planorbuliils from NO- 104 locality showed that they had the unique placement of the labial lamella characteristic of P. nebraskensis (Figure 4). This sug- gested that the P. nebraskensis v\as not an unique ab- erration of the KX-145 locality, and nearby Lyons Coun- ty, Iowa only, but a widespread taxa. This also prompted reevaluation of several older (late Pliocene, Blancan, ca. 2 MA) faunas from Nebraska that included planorbulids, especially those reported by Pierce (1990), Careful re- evaluation, including break-back of specimens, showed that the planorbids of late Pliocene Localities KX-109, KX-143 and CD- 1 04, had been incorrectly identified as P. armigera. These specimens were clearK' P. nebras- kensis (Figure 5), and demonstrated a significant range, both spatially and temporally, for this species. Fossil planorbulids from strata older than Quaternary are not common, possiblv due to the unfortunate past habit of lumping man> disparate planispiral taxa under a form genus "Planorbis" Ta\lor (1960:36) lists P. ar- migera as a component ot the Sand Draw fauna (Blan- can), although I have yet to collect a planorbulid from that locality for comparison. Taylor (1966) refers, with question, several Idaho and Wyoming planorbulids, also of Blancan age, to Planorbiila eampestris (Dawson, 1875). Hannibal (1912:157-158) described P. mojavensis from the Miocene Barstow Formation of California, The la- mellae of P. mojavensis are unique, with only three lamellae on the outer lip (Taylor, 1954:74). Pierce (1993) described Planorbula powelli from the late Oli- gocene (Arikareean) Cabbage Patch beds of southwest- ern Montana. The lamellae of Planorbiila powelli (Fig- ure 6) are also of the pattern characteristic of modern P. wheatleyi. i.e., with a vertically oriented upper palatal lamella When compared in series, an evolutionary se- quence, beginning with Oligocene Planorbiila powelli. and extending through P. nelmiskensis to P wheatleyi is strongly suggested. This also suggests that P armigera may be a late offshoot of the older P. wheatleyi lineage. The slight inclination of the upper palatal lamella noted with the otherwise typical P. nebraskensis from the Nuckols County locality may represent the beginning ot Page 24 THE NAUTILUS, Vol. 108, No. 1 c^ sp cr> c^ r — > d < 5 ■^up u D ^ 0 H ^^==3lp ^==^ ^^:r^ ^ ^ (f \ Q (J 1 1 2 3 4 5 6 Figures 1-6. Labial lamellae of planorbulid snails, greatK en- larged. sp=suprapalatal; up=iipper palatal; lp=lo\ver palatal; b=basal. 1. Plariorbula armigera (Sa\ 1821). 2. P. wheatleyi (Lea 1858). 3. P. riebraskensis Leonard 1948, hypotype. 4. P. nelwaskcnsis. Localit\ NO- 104, 5. P. ni'braskrnsis. Locality KX-109, Blancan 6. Planorlnila pouclli, Oligoeene (Arika- reean) Cabbage Patch beds. F"igiires 1 and 2 alter Baker 194.5, Plate 76. Figures 7, 8. .-Xpertural lamellae of P ncliKiskcnsis, aiiproxi- niately 20X, both from lot PPLN 1027, Locality K\-145 (Leon- ard's type locality). 7. Lamellae « ithin aperture ot small spec- imen. 8. .Aperture of small specimen broken back to radial of apertnral lamellae. a divergence that led to the lamellar pattern of modern P. armigera. Planorbula vulcanata Leonard 1948, a fossil of mid-Pleistocene age (ca. (5OOKA) from southwestern Kansas and adjacent Oklahoma, appears to be the oldest confirmed planorbulid with typical P. armigera denti- tion. This suggested lineage results, however, in a dis- tributional inconsistency. Oligocene Planorlnila powelli were toinid to have existed in a climate similar to that preferred by modern P . wheatleyi, which is found only in the humid mesothermal climatic zone of the south- eastern United States. Planorbula riebraskensis appar- ently preferred a cooler, humid, microthermal climate. Planorbula armigera, on the other hand, is a very tol- erant species, widely distributed from Louisiana to the Northwest Territories of Canada, currently occupying both mesothermal and microthermal climatic zones. LITERATURE CITED Baker, F. C. 1945. The molluscan famiK Planorbidae: Col- lation, revision and additions by H. J. \'an Cleve. L'ni- versily of Illinois Press, Urbana, 5.30 p. Bolick, M R, 1991. Pollen and spore anaKsis of sediment from the .\lbert .^hrens Locality, a medial Pleistocene vertebrate paleontolog\ site (abst). Proceedings of the Ne- braska .Academy of Sciences 1991:55. Dawson, G. M. 1875. Land and fresh-water Mollusca, col- lected during the summers of 1873-74, in the vicinit\ of the forty-ninth parallel- Lake of the Woods to the Rock\' Mountains, p. 347-350. (In) North .American Boundary Commission, 1872-1876, Report on the geolog\ and re- sources of the region in the vicinity of the forty-ninth parallel, from the Lake of the Woods to the Rocky Moun- tains, with lists of plants and animals collected, and notes on the fossils. B. Westerman & Co., New York. Hannibal, H. 1912. A synopsis of the Recent and Tertiary freshwater Mollusca of the Californian Province. Pro- ceedings of the Malacological Societs 10:112-211, Lea, I, 1858, Descriptions of a new Helix and two new plan- orbes. Proceedings of the .^cademv of Natural Sciences of Philadelphia 10:41, Leonard, A, B. 1948. Five new Yarmouthian planorbid snails. The Nautilus 62:41-47, Leonard. .\, B, 1950, .\ Yarmouthian molluscan fauna in the midcontinent region of the I'nited States. Uni\ersit\ of Kansas Paleontological Contributions, Mollusca. .\rticle 3. 48 p. Pierce. H. G, 1990, Two unusual gastropods from late Plio- cene lakes in northeast Nebraska, The Nautilus 104(2):53- 56, Pierce, H, G. 1993. The nonmarine mollusks of the late Oli- gocene-early Miocene Cabbage Patch fauna of western Montana. III. Aquatic mollusks and conclusions. Journal of Paleontology 67(6):980-993. Say, T. 1821 Descriptions of univalve shells of the United States. Journal of the .\cademy of Natural Sciences of Philadelphia 2(1): 149-179. Taylor, D. W 1954. Nonmarine mollusks from Barstow For- mation of southern California. U. S. Geological Survey Professional Paper 254-C:67-80. Ta\lor, D. W. 1960. Late Cenozoic molluscan faunas from the High Plains. U. S. Geological Survey Professional Paper 337. 94 p Ta\lor, 1), W 1966, Summarx of North American Blancan nonmarine mollusks, Malacologia 4(1): 1-172, \oorhies. M, R, and R G, Corner 1991. Paleoclimatic im- plications of a new medial Pleistocene vertebrate site in south-central Nebraska; Irvingtonian mammals from the Albert Ahrens Locality (abst). Proceedings of the Nebraska ,\ca(leni\ of Sciences 1991:63. THE NAUTILUS 108(l):25-26, 1994 Page 25 Effects of Preservatives on Wet-weight Biomass of the Asiatic Clam, Corbicula fluminea Michael E. Smith Gerry A. Lanfair Department of Biology Valdosta State University Valdosta, GA 31698 USA The Asiatic clam, Corbicula fluminea (Miiller, 1774) was first collected in North America from Vancouver Island, Canada in 1924 and in the United States from near the mouth of the Columbia River during 1938 (Counts, 1986; McMahon, 1982). Since its introduction, C fluminea has spread to the east coast of the United States (Counts, 1986) and into South America (Ituarte, 1981) becoming an important component of freshwater ecosystems. The Asiatic clam is often found in high densities creating problems such as biofouling of industrial water intakes and power plant cooling systems (Isom, 1986). Corbicula fluminea is also recognized as an important biological monitor in freshwater ecosystems (Doherty, 1990). When sampling C. fluminea, individuals are usually preserved in the field for later laboratory analy sis. Pres- ervation can cause changes in wet- weight biomass (Don- ald & Paterson, 1977; Herke, 1973; Howmiller, 1972; Landahl & Nagell, 1978; Mills et ai. 1982; Stanford, 1973; Wiederholm & Eriksson, 1977; Williams & Robins, 1982) that affect estimates of population parameters such as standing crop and production. Thus, the purpose of this study was to determine the effects of two common preservatives, 70"^' ethanol and 10"^ neutral buffered for- malin, on wet-weight biomass of the Asiatic clam, Cor- bicula fluminea. Specimens of Corbicula fluminea were collected on 6 February 1993 from the confluence of Shellstone Creek and the Ocmulgee River, Blecklev Countv, Georgia (32°25'35"N, 83°28'50"W). Induiduals were removed from their shells and live wet-weights were determined prior to preservation in 70'^- ethanol (n = 18) or 10*^' neu- tral buffered formalin (n=18). Formalin and 10'"c ethanol solutions were prepared b\ diluting 37 ''c formaldehyde solution and 95% ethanol with distilled water; the 10% formalin solution included 4 g/l sodium phosphate mono- basic and 6.5 g/l sodium phosphate dibasic to produce neutral buffering. All specimens were blotted (Donald & Paterson, 1977; Wiederholm & Eriksson, 1977), weighed to the nearest 0.1 mg and placed individually into 100 ml of preservative at room temperature (25 ± 1°C). Preserved specimens were soaked in distilled water for 30 minutes (Howmiller, 1972; Mills et ai, 1982), weighed, and returned to their individual containers. Comparison of live wet-weights between the two groups was by the use of a t-test. Means, 95% confidence intervals and linear regressions were calculated from arcsine trans- formed data. Homogeneity of regression slopes was test- ed to determine if there was a difference in weight loss over time between the formalin and ethanol groups (So- kal & Rohlf, 1981). Live wet-weights were not significantly different (P>0.05) between the formalin and ethanol groups. Mean weight loss proceeded more rapidly in the ethanol pre- served specimens and after 163 days, was 41.9%- (40.7- 43.1%; 95% C.I.) and 21.6% (20.3-22.9%; 95% C.I.) for the ethanol and formalin groups, respectively (Figure 1). Stabilization of mean weight loss occurred at appro.xi- mately day 77 for the ethanol group with only a 2.9% weight loss over the next two samplings (= 86 days). The formalin group stabilized after day 163; wet- weight bio- mass was measured at day 385 for the formalin group only and mean weight loss was 22.1%, a loss of 0.5%' since da\' 163. Regression equations were y=0.0434-f0.604x, r-=0.961 for the formalin group and y=0.0265 + 0.0690x, r-=0.985 for the ethanol group (Figure 1). Equations are only descriptive for the preservatives and temperature range used in this study. The slopes of the two lines were significantly different (P<0.001) indicating that rate of weight loss was different between the two groups. Our study demonstrates that the two most commonly used preservatives for freshwater benthic organisms, 70% ethanol and 10% neutral buffered formalin, significantly decrease wet-weight biomass of Corbicula fluminea . Use of preserved specimen biomass as an accurate determi- nation of live biomass, therefore is inappropriate. Wet- weight losses for C. fluminea fall within the range ob- served in other studies for freshwater invertebrates (Donald & Paterson, 1977; Howmiller, 1972; Landahl & Nagell, 1978; Stanford, 1973; Wiederholm & Eriksson, 1977). Howmiller (1972) found, as we did, that formalin preservation resulted in less wet-weight loss compared to 70% ethanol. We recommend that the regression equations be used to convert preserved wet-weight biomass to live wet- weight biomass if 10% neutral buffered formalin or 70% ethanol is used in the temperature range of 25 ± 1°C. Page 26 THE NAUTILUS, Vol. 108, No. 1 3 O 100 125 TIME (DAYS) 175 30 IJJ o 25 DC and history of the invasion of the United States by Corbicula fluniinea (Bi- valvia: Corbiculidae). American Malacological Bulletin Special Edition No. 2:7-39. Doherty, F. G. 1990 The .\siatic clam, Corbicula spp., as a biological monitor in freshwater environments. Environ- mental Monitoring and .\ssessment 15:143-181. Donald, G L. and C G. Paterson. 1977 Effect of preservation on wet weight biomass of chironomid larsae Hydrobiol- ogia 53:75-80. Herke, W. H. 1973. Spin-drying of preserved fishes and ma- croinvertebrates. Transactions of the American Fisheries Society 102:643-645, Howmiller, R. P. 1972. Effects of preservatives on weights of some common macrobenthic invertebrates Transactions of the American Fisheries Societv 101:743-746. Isom, B. G. 1986. Historical review of Asiatic clam {Corbi- cula) invasion and biofouling of waters and industries in the Americas. American Malacological Bulletin Special Edition No. 2:1-5. Ituarte, C. F. 1981. Primera noticia acerca de la introduccion de pelecipodos Asiaticos en el area Rio Platense. Neotro- pica 27:79-82. Landahl, C.-C. and B. Nagell 1978 Influence of the season and of preservation methods on wet- and dr\ weights of larvae of Chirononius plumosus L. Internationale Revue der gesamten Hydrobiologie 63:405-410 McMahon, R. F. 1982. The occurrence and spread of the introduced Asiatic freshwater clam, Corbicula fluminea (Miiller) in North America: 1924-1984. The Nautilus 96: 134-141. Mills, E. L., K. Pittman, and B. Munroe. 1982. Effect of preservation on the weight of marine benthic inverte- brates. Canadian Journal of Fisheries and .\quatic Sciences 39:221-224, Sokal, R. R. and F, J, Rohlf , 1981. Biometrv 2nd edition. \V. H. Freeman and Co., New York Stanford, J. \ 1973. A centrifuge method for determining live weights of aquatic insect larvae, w ith a note on weight loss in preservative Ecolog\ 54:449-451, Wiederholm, T. and L. Eriksson. 1977 Effects of alcohol- preservation on the weight of some benthic in\ ertebrates. Zoon 5:29-31 Williams, R. and D B. Robins. 1982. Effects of preservation on wet weight, dry weight, nitrogen and carbon contents of Calanus helgolandicus (Crustacea: Copepoda). Marine Biology 71:271-281. 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Box 7279, Silver Spring, MD 20907-7279, USA. Chdujic of (iddress: Please inform the publisher of your new address at least 6 weeks in advance. All comnumications should include both old and new addresses (with zip codes) and state the effective date. THE NAUTILUS (ISSN 0028-1344) is publish(>d (piarterlv by Trophou (Corporation, 8911 Alton Parkway, Silver Spring, MD 20910. Second (Class postage paid at SiKer Spring, MD and additional mailing offices. POSTMASTER: Send address changes to: THE NAUTILUS PO. Box 7279 Silver Spring. MD 20907-7279 T H E t7N AUT I L U S CONTENTS Volume 108, Number 2 March 7, 1995 ISSN 0028-1344 Bruce A. Marshall Roland Houarl A Review of the New Zealand Recent Species of Poirieria Jousseaume, 1880 (Mollusca: Gastropoda; Muricidae) with Description of a New Species 27 A New Species of Charitodoron (Gastropoda: Mitridae) from Mozambique, with Notes on C. veneris (Barnard, 1964) 34 Taxonomic Notes on Kenyonia Brazier and Conopletira Hinds (Gastropoda: Conoidea) 37 Four New Genera for Northeastern Pacific Prosobranch Gastropods 39 Population Structure, Growth and Fecundity of Melampus bidcntatus (Sa\ ) from Two Regions of a Tidal Marsh Complex in Connecticut 42 48 R. >. Kilburn Donn L. Tippett John K. Tucker James H. McLean Jessica A. Spelke Paul E. Fell Lucille L. Helvenston News and Notices THE NAUTILUS 108(2):27-33, 1995 Page 27 A Review of the New Zealand Recent Species of Poirieria Jousseaume, 1880 (Mollusca: Gastropoda: Muricidae) with Description of a New Species Bruce A. Marshall Museum of New Zealand Te Papa Tongarewa P O. Box 467 Wellington, New Zealand Roland Houart Research Associate Institut Royal des Sciences Naturelles de Belgique Vautierstraat 29 B-1040 Brussels, Belgium ABSTRACT Poirieria syrinx n. sp. is described and compared with P. ze- landica (Quoy & Gaimard, 1833) and P. hopua Dell, 1956. Poirieria syrinx is recorded from Early Pleistocene beds at Palliser Bay, and living at 482-786 m off the east coast of the North Island. It is locally sympatric with P. zelandica, which is widely distributed off New Zealand at 0-540 m depth Chal- lenger Expedition specimens of P. zelandica reputedly from Tongatabu and the Kermadec Islands are considered to have been translocated from Queen Charlotte Sound, New Zealand. Poirieria hopua occurs at 490-1006 m off the east coast of the South Island. Some groups with similar species are commented upon, including Pagodula Monterosato, 1884 and Enixotro- phon Iredale, 1929 The type species of Enixotruphon (Tro- phon carduelis Watson, 1883) is newly recorded from New Zealand, and its radula is illustrated for the first time. Key words: Mollusca, Gastropoda, Muricidae, Poirieria. new species. INTRODUCTION The present contribution was initiated when we recog- nized an undescribed species of Poirieria from the con- tinental slope of the North Island of New Zealand. The new species differs markedly from the well known spe- cies P. zelandica (Quoy & Gaimard, 1833) and the rare .P. kopua Dell, 1956 in having tubular shell spines. ACRONYMS AMS The Australian Museum, Sydney BMNH The Natural History Museum, London MNHN Museum National d'Histoire Naturelle, Paris MNZ Museum of New Zealand, Wellington NZGS Institute of Geological and Nuclear Sciences, Lower Hutt NZOI National Institute of Water and Atmospheric Research, Wellington SYSTEMATICS Order Neogastropoda Thiele, 1929 Superfamily Muricoidea Rafinesque, 1815 Family Muricidae Rafinesque, 1815 Subfamily Muricinae Rafinesque, 1815 Genus Poirieria Jousseaume, 1880 Poirieria Jousseaume, 1880335. Type species (by original des- ignation): Murex zelandicus Quoy & Gaimard, 1833; Plio- cene-Recent, New Zealand. Remarks: The radula of Poirieria species is morpholog- ically very similar to those of the Recent European mur- icine Trophon echinatus (Kiener, 1840) (Bouchet & Waren, 1985, fig. 333) and a number of bathyal Indo- West Pacific species (mostly undescrit)ed — AMS, MNHN, MNZ), among them Trophon carduelis Watson, 1883 (Watson, 1886, pi. 10, fig. 7), the type species of Enix- otrophon Iredale, 1929 (type locality New South Wales: here newly recorded from off the North Island and the west coast of the South Island, New Zealand, living at 676-1217 m— MNZ, NZOI) (figures 1-4). Some of these have even longer shoulder spines than most Poirieria species, though all differ in having the shoulder spines set closer to the rim of each varix, and radular teeth that are larger in shells of comparable size. Neither Poirieria species nor Trophon echinatus or T. carduelis appear to be closely related to Trophon Montfort, 1810 (type spe- cies Buccinum geversianum Pallas, 1774; Recent, south- ern South America) or to Boreotrophon Fischer, 1884 (type species Murex clathratus Linnaeus, 1758; Recent, northeastern Atlantic), both of which differ in details of central radular tooth morphology (Harasewych, 1984; personal observation). Trophon echinatus is closely sim- ilar to Murex vaginatus De Cristofori and Jan, 1832 Page 28 THE NAUTILUS, Vol. 108, No. 2 Figures 1-4. Radulae of New Zealand Poirieria and Pagodula species. 1. Poirieria zelandica (Quoy & Gaimard, 1833), subadult, off Matakoa Point, Hicks Bay, 99-102 m, MNZ M. 60498, shell height 30 0 mm 2. Poirieria syrinx new species, holotype (adult). 3. Poirieria kopua Dell, 1956, adult paratype, Chatham Rise, .530 m, MNZ M. 10506. shell height 1-4.7 mm 4. Pagodula carduelis (Watson, 1883), adult, off Gisborne, 913-750 m, NZOI sta. E719, shell height 50.5 mm. Scale bars 1-3 = 100 Mm, 4 = 1.0 mm. (Pliocene, Italy), which is generally interpreted as the type species of Pagodula Monterosato, 1884 (Bouchet & Waren, 1985:126). It seems plausible, therefore, that Pa- godula is the most appropriate genus group name for T. carduelis, and thus by implication that Enixotrophon is a junior synonym. The type species of Paziella Jousseaume, 1880 (Murex pazi Crosse, 1869; Recent, Caribbean), and especially Actinotrophon Dall, 1902 (Trophon actinophorus Dall, 1889; Recent, Caribbean) (see Bayer, 1971, figs. .30, .35D) are so similar to New Zealand Recent Poirieria species in shell and radular morphology that separation on these characters alone seems untenable. Harasewych (1984) and Kool (1993a, b), however, have shown that shell characters (unlike anatomy) in Muricidae are subject to convergence, while Kool (1987) concluded that radular characters arc largely unrelated to food-type and are thus a useful clue to phylogenetic relationships. We fol- low Yokes (1970, 1992) in treating Paziella and Actin- otrophon as subgenera of Poirieria. Incidentally, although Paziella was introduced one line before Poirieria (Jousseaume, 1880:335), Poirieria has consistently been treated as the senior synonym (e.g. Yokes, 1970, 1992) following Cossmann (1903), who was the first reviser according to ICZN Article 24. Poirieria zelandica (Quoy & Gaimard, 1833) (Figures 1, 5, 7, 10-15) Murex zelandicus Quoy & Gaimard, 18.33: 529, pi. .36, fig. 5-7. Poirieria zelandica Maxwell, 1971 767 (earlier synonymy); Powell, 1979: 170, pi. .35, fig 2; Beu & Maxwell. 1990: 358, pi. 48K. Type material: Holotype MNHN: Cook Strait. Other material examined: About 100 Late Pliocene and Pleistocene specimens (MNZ, NZGS), and about 1500 Recent specimens in 200 lots (MNZ). Distribution: (figure 5) Early Pliocene (Opoitian) to Re- cent. Three Kings Islands southward to Stewart Island, Challenger Plateau and Chatham Islands, New Zealand, living at 0-540 ni. Remarks: This well known species is common through- out most of its range at 20-150 m depth. Small (imma- ture) living specimens occur rarely at considerably great- er depths, particularly in the Bay of Plenty (deepest record 490-540 m). One of us (B.A.M.) has taken lightly abraded short-spined specimens on numerous occasions alive from sand at low tide at the eastern end of Ohope B. A. Marshall and R. Houart, 1995 Page 29 Figure 5. Map of New Zealand showing distribution of Poir- ieria zelandica (Quoy & Gaimard, 1833). 200 and 1000 meter contours indicated. Figure 6. Map of New Zealand showing distribution of Poir- ieria syrinx new species (stars) and Poirieria koptia Dell, 1956 (solid circles). 200 and 1000 meter contours indicated. Beach, but these had probably been transported inshore by storms. There is considerable variation in the length of the spines (figures 10-14), which tend to be longest in specimens from muddy substrata, and shortest in spec- imens from coarser substrata. Grading southward through intermediate forms, specimens from off the southern South Island and Stewart Island (figure 11) attain about half the size of the largest northern specimens, and are more stoutly built, often with a few low rounded den- ticles within the outer lip. In their stout build the Recent southern form approaches Late Pliocene (Mangapanian) and Early Pleistocene (Nukumaruan) forms of P. zelan- dica (figure 15), though the fossils are consistently thick- er, have more strongly dentate outer lips, and attain the size of Recent northern specimens. It should be appre- ciated that all of the fossils are from the southern half of the North Island, and since shells of equivalent age are unknown from further north (due to lack of expo- sures), it is unknown whether northern Late Pliocene- Early Pleistocene shells were more lightly built. Maxwell (1971) concluded that P. zelandica evolved gradually from P. primigena Finlay, 1930, a species in which the shoulder is set lower on the whorls and that ranges from Late Oligocene (Duntroonian to Early Pliocene (Opoi- tian). Watson's (1886:157) records of P. zelandica from Challenger stations 171 (north of Raoul Island, Ker- madec Islands, 1097 m) and 172 (off Tongatabu, 18 m,) are highly anomalous since, apart from the Challenger Plateau record, the species has never been obtained from beyond the New Zealand continental shelf. These spec- imens (BMNH 1887.2.9.546-7, 1887.2.9.548) are per- fectly accordant with the specimens from Challenger station 167A (Queen Charlotte Sound, New Zealand, 18 n, BMNH 1887.2.9.544-5) and numerous additional specimens from the vicinity (MNZ). Although it seems clear that the specimens from stations 171 and 172 were introduced through entanglement in the nets or ropes, it is significant that they were not detected at stations 168-170, presumably through oversight or the use of a different trawl. According to the Narrative of the Cruise (Tizard et ai, 1885) and the Summary of Results (Mur- ray, 1895), stations 168-171 were sampled by trawl and station 172 by dredge. According to Tizard et al. (1885: 1012) station 167A was by dredge alone, but in fact there were also two trawlings at this station (Murray, 1895: Page 30 THE NAUTILUS, Vol. 108, No. 2 593), which would account for the presence of contam- inants in both dredge and trawl stations. Poirieria syrinx Marshall and Houart, new species (figures 2, 6, 8, 16, 17) Poirieria zelandica - Dell, 1962:76 (not Quoy & Gaimard, 1833). Poirieria kopua - Dell, 1963:212 (not Dell, 1956); Powell, 1979: 170 (text in part). Description: Shell of medium size for the genus, length up to 45 mm, broadly fusiform, spire slightly higher than aperture, suture well impressed, glossy, white or salmon pink. Protoconch 1.20 mm wide, taller than broad, of l'/4-iy4 convex whorls, smooth, glossy, terminal varix prominent. Teleoconch of up to 5Vi convex whorls, suture well impressed. Varices thin, sharp-edged, prominently spinose, numbering 5 on 1st whorl, 5 on 2nd, 5 or 6 on 3rd, 5 or 6 on 4th, and 4 or 5 on 5th whorl. Spines slender, sharp, gently curved, bases set behind varical rim, shoul- der spine strongly posterior; leading edge open to varical rim at first, becoming closed after 3rd whorl so that proximal part of each spine is fully tubular. Shoulder spine row the most prominent, situated medially on ear- liest whorls, then supramedially. Secondary spines ab- apical, 3, relative sizes variable, about equidistant, com- mencing after 3rd whorl, bases centered well in front of that of shoulder spine, adapical spine row exposed on spire, set about midway between shoulder spine row and suture or abapically, abapical spines of previous whorl in front of aperture. Aperture roundly ovate. Inner lip thin, smooth, fully contacting or free over abapical half. Figures 7-9. Outer lip profiles of New Zealand Poirieria species. 7. Poirieria zelandica (Quoy & Gaimard, 1833), Or- chard Bay, Marlborough Sounds, 29 m, MNZ M. 45110, shell height 28.0 mm. 8. Poirieria syrinx new species, holotype, MNZ M. 117782, shell height 26.0 mm. 9. Poirieria kopua Dell, 1956, paratype, Chatham Rise, 530 m, MNZ M. 10506, shell height 14.7 mm. Scale bars = 4 0 mm. Figures 10-18. Shells of New Zealand Poirieria species. 10- 15. Poirieria zelandica (Quoy & Gaimard, 1833) 10. Holotype, MNHN, shell height 52.0 mm. 1 1. Port Pegasus, Stewart Island, 40-46 m, MNZ M. 26617, shell height 32 7 mm 12. Off Poor Knights Islands, MNZ M. 83858, shell height 54.0 mm. 13. Orchard Bay, Marlborough Sounds. 29 m, MNZ M.451 10, shell height 28.0 mm. 14. Off Waiheke Island, 40 m, MNZ M 35635, shell height 57.5 mm. 15. Cliffs west of Whangaimoana, Palliser Bay, Early Pleistocene (Nukumaruan), MNZ M. 40361, shell height 32.8 nmi. 16, 17. Poirieria si/rini new species. 16. Cliffs west of Whangaimoana, Palliser Bay, Early Pleistocene (Nu- kumaruan), MNZ M. 91795, shell height 25.8 mm (incomplete,). 17. Holotype, MNZ Ml 17782, shell height 26.0 mm 18. Poir- ieria kopua Dell, 1956, paratype, Chatham Rise, 530 m, MNZ M. 10506, shell height 14.7 mm. B. A. Marshall and R. Houart, 1995 Page 31 Page 32 THE NAUTILUS, Vol. 108, No. 2 Outer lip thin at rim, internally slightly thickened, smooth, simple. Siphonal canal long, semitubular, spineless, gent- ly curved, 2 or 3 previous canals retained. Outer shell layer (intriticalx) hard, dense, weakly or locally obscurely spirally lirate, translucent, glossy, becoming chalky through erosion. Animal with small subparallel cephalic tentacles, eyes on outer edges at about midlength, subcylindrical behind them, tapered before. Male unknown. Radula (figure 2) similar to that of Poirieria zelandica. Type material: Holotype (M. 117782,) and 2 paratypes MNZ: BS761 {R119), 37°22.0'S, 176°40'E, 37 km E of Mayor I., New Zealand, alive, 616-666 m, January 24, 1979, R.V. Tangaroa (type locality). Other paratypes (10): 1364, 34°46'S, 174°05.8'E, NE of Cavalli I. alive, 492 m, November 20, 1977, R.V. Tangaroa (1 NZOI); 24 km NE of Plate I., alive, 622-585 m, October 29, 1962,M.V. Ikatere (1 MNZ); BS741 (R99), 37°20.6'S, 176°28.0'E, 17 km E of Mayor I., alive, 482-550 m, January 22, 1979, R.V. Tangaroa (3 MNZ); 37°31.64'S, 176°55.35'E, SW of White I., alive, 500-518 m, January 20, 1993, F.V. Kaharoa (1 MNZ); VUZ97, 41°33'S, 174°57'E, off Palliser Bay, alive, 786 m, August 28, 1957; (3 MNZ, 1 R. Houart collection). Other material examined: Cliffs west of Whangai- moana, Palliser Bay, New Zealand, 1971-73, B.A. Mar- shall, Early Pleistocene (Nukumaruan) (1 specimen). Distribution: (figure 6): Early Pleistocene (Nukumaruan) of Palliser Bay, and Recent off northeastern North Island and Cook Strait, New Zealand, living at 482-786 m on mud. Remarks: Compared with Poirieria zelandica, which it most resembles in shell morphology, P. syrinx n. sp. dif- fers in having the bases of the spines set further behind each varical rim, and in that the proximal parts of each spine become fully tubular after the third teleoconch whorl. Other shell differences include the smaller size (maximum length 45 mm vs. 71.5 mm), the larger pro- toconch (width 1.2 mm vs. of 0.9-1.0 mm), the more prominent spiral microlirae, and the absence or extreme weakness of spiral swellings between the bases of the spines. The animal (preserved material) differs markedly from that of P. zelandica in having considerably shorter cephalic tentacles that are almost parallel instead of strongly divergent, with eyes situated closer to the head. Unfortunately the male of P. syrinx n. sp. was not avail- able for comparison of the penial morphology. Their radulae are similar (figures 1, 2). Poirieria syrinx n. sp. and P. zelandica have overlapping geographic and bathymetric ranges, and the two species have been ob- tained living together (37°20.6'S, 176°28.0'E, off Mayor Island, 482-550 m, MNZ M.60247, 95064). Poirieria kopua differs from P. syrinx n. sp. in attain- ing a smaller size (to 19.0 mm high), in having a larger protoconch (width 1.5-1.7 mm), in having a fully open shoulder spine on each varix, and in lacking secondary spines. Judging from their protoconchs, both species have non-planktotrophic larval development (probably di- rect). They appear to be allopatric, though they could conceivably occur together off Cape Campbell. None of the New Zealand Tertiary species have tubular spine bases (Maxwell, 1971). The Early Pleistocene (Nuku- maruan) specimen (figure 16) is indistinguishable from Recent shells. Reus (1967) conclusion that the Whan- gaimoana beds were deposited at 400-600 meters depth is accordant with the known bathymetric range of this species. Etymology: Greek syrinx (pipe). Poirieria kopua Dell, 1956 (figures 3, 6, 9, 18) Poirieria kopua Dell, 1956:114, pi. 16, figs. 161, 162, 165; Max- well, 1971:771, figs. 26, 27; Powell, 1979:170 (in part), fig. 42/1. NOT Poirieria kopua Dell, 1962:76; Dell, 1963:212 (=P. syrinx n. sp.). Type material: Holotype MNZ M.9777 and 2 paratypes (MNZ and Canterbury Museum, Christchurch): CLE. sta. 59, 43°38'S, 177°19'E, Chatham Rise, alive, 530 m, February 11, 1954, M.V. Alert. Other material examined: (11 specimens) BS559, 43°14'S, 173°39'E, wall of Pegasus Canyon, off Kaikoura, New Zealand, dead, 1006-512 m, September 27, 1976, R.V. Acheron (2 MNZ); BS201, 44°45.6'S, 171°05'E, off Taia- roa Head, dead, c. 549 m, Januarv 23, 1957, M.V. Alert (3 MNZ); BS582, 45°46'S, 171°03'E, off Taiaroa Head, dead, 660 m, September 1, 1976, R.V. Munida (5 MNZ); Mu 70-45, 45°50'S, 171°01'E, off Taiaroa Head, dead, 540-490 m, October 22, 1970, R.V. Munida (1 NZGS). Distribution: (figure 6) Chatham Rise and off the east coast of South Island, from Kaikoura to Taiaroa Head, New Zealand, 490-1006 m, taken alive at 530 m. Remarks: The shell of P. kopua differs from that of P. zelandica and P. syrinx n. sp. in attaining smaller shell size (to 19.0 mm), in having a larger protoconch (width 1.50-1.70 mm), in lacking secondary spines below the peripheral spines, and in that the spines are not fully tubular at their bases. The radula is similar in all three Recent species of Poirieria (figures 1-3). ACKNOWLEDGMENTS We thank Philippe Bouchet (Museum National d'His- toire Naturelle, Paris,) and Kathie M. Way (The Natural History Museum, London) for the loan of material, and Alan G. Beu (Institute of Geological and Nuclear Sci- ences, Lower Hutt), Philip A. Maxwell (Waimate) and Emily H. Vokes (Tulane L'niversity, New Orleans) for comments on the manuscript. Thanks also to Michael Hall and Norman Heke (Museimi of New Zealand, Wel- lington) respectively for the photographic printing and photography, and Wendy St (ieorge (Institute of Geo- B. A. Marshall and R. Houart, 1995 Page 33 logiai and Nuclear Sciences, Lower Hutt) for assistance with the scanning electron microscopy- LITERATURE CITED Bayer, F. R.V Beu, Beu, M. 197L New and unusual mollusks collected by John Elliott Pillsbuby and R.V. Gerda in the tropical Western Atlantic. In: Bayer, F M and G.L. Voss (eds. ). Studies in tropical American Mollusks. University of Miami Press, Coral Gables, Florida! 11-236. A. G 1967 Deep-water Pliocene Mollusca from Palliser Bay, New Zealand. Transactions of the Royal Society of New Zealand, Geology 5(3):S9-122. A. G. and P. A. Maxwell 1990. Cenozoic Mollusca of New Zealand. New Zealand Geological Survey Paleonto- logical Bulletin 58, 518 pp. Bouchet, P and A. Waren. 1985. Revision of the Northeast Atlantic bathyal and abyssal Neogastropoda excluding Turridae (Mollusca, Gastropoda) Bollettino Malacologico, Supplemento 1:121-296 Cossmann, M. 1903 Essais de Paleoconchologie Comparee, 5. Cossmann, Paris 215 pp. R K 1956 The archibenthal Mollusca of New Zealand Dominion Museum Bulletin 18, 235 pp. R. K. 1962. Additional archibenthal Mollusca from New Zealand. Records of the Dominion Museum 4(6):67-76. R K. 1963. Archibenthal Mollusca from northern New Zealand Transactions of the Royal Society of New Zea- land. Zoology 3(201 );205-216. Harasewych, M G 1984 Comparativeanatomy of four prim- itive muricacean gastropods: Implications for trophonine phylogeny. American Malacological Bulletin 3: 11 -26 Jousseaume, F 1880. Division methodique de la famille des purpurides. Le Naturaliste 42:335-336. Kool, S. P. 1987. Significance of radular characters in recon- Dell, Dell, Dell, struction of thaidid phylogeny (Neogastropoda: Murica- cea). The Nautilus 101:117-132. Kool, S. P. 1993a. The systematic position of the genus Nucella (Prosobranchia: Muricidae: Ocenebrinae). The Nautilus 107:43-57. Kool, S. P. 1993b. Phylogenetic analysis of the Rapaninae (Neogastropoda: Muricidae). Malacologia 35:1.55-259. Maxwell, P. A. 1971. Notes on some Cenozoic Muricidae (Mollusca: Gastropoda) from New Zealand, with a review of the genus Poirieria Jousseaume, 1880. New Zealand Journal of Geology and Geophysics 14:757-781. Murray, J. 1895 A summary of the scientific results obtained at the sounding, dredging and trawling stations of H.M.S Challenger Challenger Report. A Summary of the Sci- entific Results 1. 796 pp Powell, A. W B 1979. New Zealand Mollusca. Marine, Land and Freshwater Shells. Collins, Auckland. 500 pp. Quoy, J C R and P. Gaimard. 1833. Voyage de decouvertes de I Astrolabe, execute par ordre du Roi pendant les annees 1826-1827-1828-1829 sous le commandement de M. J. Dumont d'Urville. Mollusques. Zoologie 2[ii]:321-686 Tizard, T H , H. N Moseley, J. Y Buchanan and J Murray. 1885. Narrative of the cruise of H. MS Challenger, with a general account of the scientific results of the expedition Challenger Report. Narrative 1(2), Appendix 11:1007-1015. Vokes, E. H. 1970. Cenozoic Muricidae of the Western At- lantic region. Part 5 - Pterynotus and Poirieria. Tulane Studies in Geology and Paleontology 8: 1-50 Vokes, E. H. 1992. Cenozoic Muricidae of the Western At- lantic region. Part 9 - Pterynotus, Poirieria, Aspella, Der- momurex, Calotrophon, Acantholabia, and Attiliosa, ad- ditions and corrections Tulane Studies in Geology and Paleontology 25:1-108. Watson, R B. 1886. Report on the Scaphopoda and Gaster- opoda collected by H.M.S. Challenger during the years 1873-76. Challenger Report, Zoology 15, 756 pp. THE NAUTILUS 108(2):34-36, 1995 Page 34 A New Species of Charitodoron (Gastropoda: Mitridae) from Mozambique, with Notes on C. veneris (Barnard, 1964) R. N. Kilburn Natal Museum P/Bag 9070 Pietermaritzburg, 3200, South Africa ABSTRACT Charitodoron rosadoi n. sp., from off Bazaruto Island, Mozam- bique, in 200-350 m, is described. This is the first Recent record of the genus Charitodoron Tomlin, 1932, from tropical waters. Mitromorpha veneris, Barnard, 1964, is referable to Charito- doron, and adult examples are described and figured for the first time. Key words: Mitridae, southern African, continental shelf, Charitodoron. INTRODUCTION The genus Charitodoron Tomlin, 1932, is distinguished from other genera of the Mitridae by the total absence of columellar pleats. Cernohorsky (1976), who revised the described taxa, recognized three species, all of which are endemic to the Agulhas Bank, the temperate-water continental shelf off South Africa. Subsequently, Kilburn (1986:635, fig. 21) pointed out that the holotype of Mi- tromorpha veneris Barnard, 1964, from off Zululand, is actually a worn, juvenile example of a species of Char- itodoron, but did not discuss its identity further. Most recently Lozouet (1991) recorded the genus for the first time from the Upper Oligocene of France, and illustrated several South African species. During the Natal Museum Dredging Programme, a numf)er of samples of a distinctive species of Charito- doron were collected off Natal and Zululand, confirming that the range of the genus extends into subtropical wa- ters. The early whorls in tiiis material agree with the holotype of the supposed turrid Mitromorpha veneris (Barnard, 1964). Subsequently, Mr. Jose Rosado of Ma- puto brought to my attention some specimens taken from crayfish traps off Mozambicjue, which prove to be an undescribed species of Charitodoron, and provide tlie first record of the genus from tropical East Africa. ABBREVIATIONS a/h = ratio of aperture length (measured along main axis) to total shell length. b/h = ratio of maximum protoconch breadth to its height. 1/h = ratio of shell breadth to total length. MHNM = Museu de Historia Natural, Maputo, Mozam- bique. NMSA = Natal Museum, Pietermaritzburg. NMDP = Natal Museum Dredging Programme. SAMC = South African Museum, Cape Town. TAXONOMY Family MITRIDAE Charitodoron Tomlin. 1932 Charitodoron rosadoi n. sp. Figures 1, 2 Diagnosis: Whorls almost flat, forming a very narrow shoulder immediately below suture, left side of base of body whorl shallowly concave; aperture with greatest width posterior to middle, columella medially convex, siphonal notch shallow; base of body whorl with 10-12 irregular spiral lirae, sculpture elsewhere weak and in- conspicuous, initially of extremely fine axial riblets, be- coming somewhat stronger on later whorls (20-30 on penultimate one), but remaining weak and irregular, forming a row of feeble nodules below suture; cream to brownish-white, with axial flames of brownish-orange, interrupted by a pale band at mid-bod\ whorl; base of body whorl white. Protoconch breadth 0.98-1.00 mm. Maximum length 16.9 mm. Description: Shell fusiform (1/h = 0.29-0.35), aperture narrow, a/h = 0.39-0.42; whorls almost fiat, forming a very narrow shoulder immediateK below suture, left side of base of bod\' whorl shallowK concave; base slightly oblique, outer lip slightly foreshortened relative to base of columella. Teleoconch whorls approximately 5.4 in number. Aperture w ith greatest width posterior to mid- dle, gradualK tapering anteriorly; parietal region shal- lowly concave, columella medialK con\ex, without trace of pleats; inner lip w ith callus deposit only in columellar region, bordered by a longitudinal depression (scarcely a false umbilicus); outer lip thin [edge damaged in all examples examined]; sipiional notch shallowly and obliquely U-shaped. R. N. Kilburn, 1995 Page 35 3 > ^^ Figures 1-4. Charitodoron rosadoi n. sp. and C. veneris, (Barnard, 1964), 1-2. Charitodoron rosadoi, holotype NMSA L1078, 16.9 X 5.5 mm 3-4. C. veneris, NMSA D4223, off Durban, 130 m, 24.0 x 7 2 mm Surface superficially smooth, other than spiral ridging on base of body whorl; under magnification whorls are seen to be axially ribbed, ribs initially extremely fine, indeed little stronger than growth lines, on later whorls strengthening, but remaining weak and irregular; ribs slightly arcuate, moderately prosocline, spacing irregu- lar, forming a row of feeble nodules where they termi- nate on the subsutural berm; 20-30 ribs on penultimate whorl, becoming obsolete on later part of body whorl. Extremely fine and faint spiral grooves visible on 2nd and 3rd whorls; base of body whorl with 10-12 irregular spiral lirae. Protoconch somewhat papilliform, limit ill-defined, evidently about 1.7 whorls; smooth, but fine axial riblets developing towards termination, and continuing onto 1st teleoconch whorl; dimensions: breadth 0.98-1.00 mm, height 0.80-0.88 mm (b/h = 1.14-1.23). Ground color cream to brownish-white, patterned with axial flames of brownish-orange, interrupted by a pale band at mid-body whorl; base of body whorl white Dimensions: 16.9 x 5.5 mm (holotype). Range: Known only from the type locality. Type material: Holotype NMSA L1078/T1 168, off Ba- zaruto Island (21°40'S; 35°25'E), Mozambique, in c. 200- 350 m, in a crayfish trap. Paratypes 1 and 2, same data. in MHNM and private collection J. Rosado respectively. All type specimens appear to have been inhabited by pagurids. Remarks: Of described species, Charitodoron rosadoi shows closest resemblance to C. agulhasensis , (Thiele, 1925) of the Agulhas Bank; in particular, the spire whorls of the latter species have a similar profile (although slight- ly more convex, with a weaker subsutural shoulder) and the color pattern is similar although much more muted; however, in C. agulhasensis the early whorls are char- acterized by much stronger spiral sculpture and the body whorl is much more convex. The ranges of C. rosadoi and C. agulhasensis are widely separated; the northern- most locality at which the latter species was found during the NMDP was off Mendu Point, Transkei (32°22.6'S; 29°00.4'E, in 250-260 m). Etymology: Named in honor of its discoverer, Mr. Jose Rosado. Charitodoron veneris (Barnard, Figures 3-4 1964) Mitromorpha veneris Barnard, 1964; 16. Type locality: Off Cape Vidai, Zululand, 80-100 fathoms [=' 145-180 m]. Charitodoron veneris: Kilburn, 1986:635, fig. 21. Page 36 THE NAUTILUS, Vol. 108, No, 2 Diagnosis: Whorls convex, suture deep, left side of base of body whorl markedly concave; aperture with greatest width anterior to middle, columella flattened, siphonai notch deep; sculpture of strong, flat-topped spiral lirae, their interstices with coarse collabral threads, feebly in- dicated on tops of lirae, 8-1 1 lirae on penultimate whorl, plus a thin thread below suture; white with axial blotches of brownish-orange, markings occasionally very pale. Protoconch breadth 1.08-1.15 mm. Maximum length 24 mm. Description: Shell fusiform (b/h = 0.29-0.30), with nar- row aperture (a/h = 0.42-0.49); whorls convex, with greatest width at basal 0.3 of each whorl, suture deep; left side of base of body whorl markedly concave; base slightly oblique, outer lip slightly foreshortened relative to base of columella. Teleoconch whorls up to 6. Aperture with greatest width anterior to middle; parietal region shallowK' concave, columella straight, callus deposit thin; siphonai notch deep and asymmetrically U-shaped; outer lip thin. Sculptured by flat-topped spiral lirae, their interstices with coarse collabral threads, feebly indicated on tops of lirae. First whorl with 6-9, close-set lirae, becoming more widely-spaced on later whorls (intervals often as wide as lirae); 8-11 lirae on penultimate whorl, plus a thin lira below suture, base of body whorl with 8-11 narrow lirae plus 13-18 somewhat angular and irregular lirae on ros- trum. Protoconch papilliform, 1.7 whorls, smooth with fine axial plicae near termination, termination sharply indi- cated; breadth 1.08-1.15 mm, height 1.00-1.25 mm (b/ h = 0.92-1.05). White with axial blotches of brownish- orange, markings occasionally very pale. Dimensions: 24.0 x 7.2 mm (largest individual exam- ined). Type material: Holotype SAMC A8750 (Kilburn, 1986: fig. 21). Distribution: Continental shelf of Zululand and Natal (Cape Vidal to Amanzimtoti), 98-320 m (fresh shells). Material examined: (all NM: NMDP unless otherwise stated): ZULULAND: S. E. of Neill Peak (Cunge), 320- 340 m, sandy mud (£4023); S. E. of Port Durnford, 310- 320 m, glutinous sandy mud (E3190); same locality, 153 m, mud, stones (E4585); off Matiguhi River mouth, 300 m, soft mud (E8903); same locality, 145 m, mud, shell rubble (E8791); same locality, 200-220 m, mud, coarse sand (P:9024). NATAL: off Tongaat Bluff, 100 m, coarse sand mud (E9817); same locality, 120 m, sandv mud (E9968); off Sheffield Ik-ach, 150 m, muddy sand (E9237); same locality, 1 10 m, muddy sand (E9303); S. E. of Sheffield Beach, 100-105 m, glutinous gray mud (E5038); N. E. of Umhianga liocks, 98 m. fine sand (S256); same locality, 107 m (B()297: A. Council); oH^ Durban, 130 m, sandstone gravel, some rocks (D4223); same locality, 95 m, fine, slightly muddy sand (D4059); same locality, 1 10- 120 m, coarse muddy sand (D3816); same locality, 100 m, very fine muddy sand (B5875); same locality, 104 m (B6287: A. Council); same locality, 104-110 m, muddy sand, broken shells (D4018); same locality, 150 m, sand- stone gravel and some sponge, living (D4172); same lo- cality, 100 m, slightly muddy sand (D3898); off Umlaas Canal, 150 m, coarse sand, numerous spatangoids, peb- bles (D789); same locality, 150 m, coarse sand (D865); off Amanzimtoti, 260-270 m, medium sand (Dl 188); same locality, 115-125 m, medium sand (Dl 285). Remarks: Juveniles of the present taxon were compared (Kilburn, 1986) with the worn and broken holotype of Mitromorpha veneris, and found to agree. Charitodoron veneris is characterized by stronger spi- ral sculpture than any other Recent member of the genus. There is a distinct but superficial resemblance in shape, sculpture and color pattern to a number of members of the conid genus Daphnclla Hinds, 1844 (s.l), but char- acters of protoconch and outer lip distinguish them at a glance. Empty shells of C. veneris are not rare, although in adults the outer lip is almost always damaged. The fore- gut anatom\ of the only example to be taken alive is presently being examined by Y. Kantor. ACKNOWLEDGMENTS This study was supported by a grant from the Foundation for Research Development (FRD). 1 thank Mr. Jose Ro- sado of Maputo for his hospitality and for making ma- terial available. For the loan of type material, 1 am in- debted to Ms. Michelle Van der Merwe (SAMC). Mrs. Linda Davis helped in the preparation of plates. Dr. D. G. Herbert kindly read the manuscript. LITERATURE CITED Barnard, K. H. 1964. The work of the s.s. Pieter Faure in Natal waters, with special reference to the Crustacea and Mollusca; with descriptions of new species of Mollusca from Natal Annals of the Natal Museum 16:9-29. Cernohorsky, W 0. 1976. The Mitridae of the World. Part 1. The subfamily Mitrinae. Indo-Pacific Mollusca 3(17); 273-.528. Kilburn, R. N. 1986. Turridae of southern Africa and Mo- zambique. Part 3. Subfamily Borsoniinae. .\nnals of the Natal Museum 27(2):633-720. Lozouet, P. 1991. Mollusca Gastropoda: Eumitra recents de la region neo-caledonieiine et Charitodoron fossiles de roligocene superieur d Aquitaine (Mitridae). In: Crosnier, A and P Bouchet (eds.) Resultats des Campagnes MU- SORSTOM 7 Memoires du Museum national d Histoire nalurelle [A] 150:205-222. THE NAUTILUS 108(2):37-38, 1995 Page 3 Taxonomic Notes on Kenyonia Brazier and Conopleura Hinds (Gastropoda: Conoidea) Donn L. Tippetl 10281 Gainsborough Road Potomac. Maryland 20854, USA John K. Tucker Illinois Natural History Survey Long Term Resource Monitoring Program-Pool 26 1005 Edwardsville Road Wood River, Illinois 62095, USA INTRODUCTION Kenyonia Brazier, 1896 is a monotypic genus that has been assigned to the families Conidae or Turridae by various authors. Brazier's description of the type species, Kenyonia pulcherrima, based on a single, unfigured. specimen, suggested a morphology so unique that he only tentatively placed the genus as a relative of Conus. He stated "This is connected with Conus and Pleurotoma and may be placed under the former genus for the pres- ent untill the animal is known." Thought for some time to be lost, the holotype and only known specimen of K Figures 1-3. Kenyonia pulcherrima Brazier, 1896 I. Holotype (SAM D6181), New Hebrides, shell length = 27 8 mm. 2. Radiograph of holotype showing resorption of internal shell structure. 3. Apical view of holotype showing pitting of shoulder sulcus. Figures 4-7. Conopleura striata Hinds, 1844. 4. USNM 869535, Panglao, Bohol Islands, Phhilippines, shell length = 14 7 mm. 5. Radiograph showing absence of internal shell resorption. 6. Apical view showing honeycomb pitting on shoulder sulcus. 7. Operculum, length = 3 mm Page 38 THE NAUTILUS, Vol. 108, No. 2 pulcherrima was located and figured by Zeidler (1985: 71, fig. 2b), who considered its family allocation uncer- tain. Powell (1966, with query) included Kenyonia in the synonymy of Conopleura, also a monotypic genus, t)ecause he presumed the pits on the shoulder slope re- ported in the description of Kenyonia to be similar to those of Conopleura (see figures 3,6.) Regarded as a turrid unequivocally, the relationships of Conopleura within the family have been uncertain. This genus had been grouped with the drilliine genera by older authors such as Fischer (1883) and Tryon (1884). However, Thiele (1929) removed it to Cytharinae ( = Mangeliinae), commenting that this assignment was un- certain. Powell (1966) referred Conopleura to the sub- family Mangeliinae. A specimen of C. striata Hinds, 1844, originally in the junior author's collection (USNM 869535), was discovered to contain an operculum. This finding together with a reevaluation of the shell mor- phology of this species suggests a different subfamilial affinity Radiographic examination of the holotype of Kenyonia indicates a reclassification of that genus is re- quired. The purpose of this paper is to report the results of these findings. RESULTS An X-ray of the holotype of Kenyonia pulcherrima (fig- ure 2) clearly demonstrates that it has undergone exten- sive interior shell remodeling (sensu Kohn et ai, 1979). Its columella is exceedingly thin. In contrast, a similar X-ray reveals the columella of Conopleura striata to be thick and stout (figure 5). The internal portions of body wall in K. pulcherrima are also resorbed and thin, while those of C. striata are thick. The operculum of Cono- pleura striata (figure 7) is leaf-shaped, with a terminal nucleus. DISCUSSION Among the toxoglossate mollusks, extensive interior re- modeling of the shell is a derived character state (Kohn, 1990) found in the Conidae (sensu Cossmann, 1896 = Conorbinae/Coninae of Taylor et a/., 1993). Therefore, K. pulcherrima and the genus, Kenyonia, belong in the family Conidae since they share this state. The presence of an operculum in Conopleura striata supports removal of the genus from Mangeliinae since members of that subfamily lack opercula. The characteristics of the oper- culum of C. striata are similar to opercula in members of the subfamily Drilliinae. Furthermore, the general shape of the shell and the presence of a subtubular sinus with parietal tubercle are typical of that subfamily. Based on these characters, we transfer the genus Conopleura to the subfamily Drilliinae. ACKNOWLEDGEMENTS We thank Karen Gowlett-Holmes of the South Australian Museum for loan of the holotype of Kenyonia pulcher- rima. Marc Moore of the Armed Forces Radiobiology Research Institutute, provided the radiographs. Photo- graphic prints were made by Richard Harris. LITERATURE CITED Brazier, J. 1896. A new genus and three new species of Mol- lusca from New South Wales, New Hebrides, and Western Australia. Proceedings of the Linnean Society of New South Wales 21:345-347. Cossmann, M 1896 Essais de Paleoconchologie comparee. Cossmann, Paris. 21-179, figs 1-48, pis. 1-8. Fischer, P. 1880-87. Manuel de conchyliologie et de paleon- tologie conchyliologique. F. Savy, Paris, 1369 pp., 23 pis. Hinds, R. B. 1844-45. The zoology of the voyage of the H.M.S. Sulphur Volume II. Mollusca. Smith, Elder & Co., London, v + 72 pp., 21 pis. Kohn, A. J. 1990. Tempo and mode of evolution in Conidae. Malacologia 32(l):55-67, figs. 1-4. Kohn, A. J, E. R. Myers, and V. R. Meenakshi. 1979 Interior remodeling of the shell by a gastropod mollusc. Proceed- ings of the National Academy of Sciences (USA)76(7): 3406-3410, figs. 1-4. Powell, A. W. B. 1966. The molluscan families Speightiidae and Turridae. Bulletin of the Auckland Institute and Mu- seum 5:1-184, pis. 1-23, figs. 1-179. Taylor, J. D., Y. I. Kantor, and A. V. Sysoev. 1993. Foregut anatomy, feeding mechanisms,relationships and classifi- cation of the Conoidea (=Toxoglossa) (Gastropoda). Bul- letin of the Natural History Museum (Zoology) 59(2): 125- 170, figs. 1-27. Thiele, J 1929. Handbuch der systematischen weichter- kunde. Gustav Fisher, Jena. I(l):l-376. Tryon, G. W. 1884. Family Pleurotomidae. Manual of Con- chology 6(23-24):151-413, pis. 1-.34 Zeidler, W 1985 Mollusc type-specimens in the South Aus- tralian Museum 2 GastropodaConidae. Records of the South Australian Museum 19(5):69-75, figs. 1-3. THE NAUTILUS 108(2);39-41, 1995 Page 39 Four New Genera for Northeastern Pacific Prosobranch Gastropods James H. McLean Los Angeles County Museum of Natural History 900 Exposition Blvd. Los Angeles, CA 90007 USA ABSTRACT In order to make the names available for use in a faunistic revision of the northeastern Pacific gastropod fauna, four new genera are proposed Naticidae: new genus Benthobulbus. type species Choristes carpenteri Dall, 1896 Eulimidae; new genus Pseudosabinella. type species Sabinella bakeri Bartsch, 1917. Muricidae, Trophoninae: new genus Ocenotrophon, type spe- cies Murex (Ocinebra?) painei Dall, 1903. Buccinidae: new genus Retimohnia, type species Mohnia frielei Dall, 1891. Key words: Northeastern Pacific Gastropoda; systematics; new- genera. INTRODUCTION In the course of preparing a checklist of the northeastern Pacific gastropod fauna, in which all generic assignments are being re-evaluated, I have been unable to reconcile the allocation of a number of species to the genera in which they have previously been placed. The following new genera have been recognized and are here intro- duced in advance of that work. Family NATICIDAE Forbes, 1838 Benthobulbus McLean, new genus Type species: Choristes carpenteri Dall, 1896. Included species: Type species and Choristes coani Mar- incovich, 1975. Marincovich (1977) treated both species in detail and illustrated their radulae; both were further discussed by McLean (1992). Diagnosis (modified from that of Marincovich, 1977:338, for "Choristes" Carpenter): Shell small to medium in size, globose, thin; whorl moderately inflated, spiral sculpture of weak lirae, suture narrowly channeled. Um- bilicus narrow to broad, simple. Inner lip slightly thick- ened, simple, lacking umbilical callus. Parietal callus thin. Operculum chitinous, entirely filling aperture. Radula with monocuspate rachidian, one monocuspate lateral tooth and two monocuspate marginal teeth per half row. Remarks: McLean (1992:291) assigned the type species of Choristes Carpenter in Dawson, 1872, to the synon- ymy of Atnauropsis islandica (Gmelin, 1791), leaving two living eastern Pacific species treated by Marincovich without a genus. Marincovich placed Choristes next to Bulbus Brown in J. Smith, 1839 (type species Bulbus smithii Brown in J. Smith, 1839; = Natica fragilis Leach, 1819), noting that only Bulbus and the two deep-water eastern Pacific "Choristes" species had the monocuspate rachidian tooth. Further remarks on the naticid radula are given by Bouchet and Waren (1993:753), who noted that the first marginal of Bulbus has three terminal den- ticles. The unsatisfactory alternatives to placement of these two species in a new genus would be to place them in Bulbus. in which the umbilicus is narrow and filled with callus, or in Euspira Agassiz, 1838, in which the rachidian is tricuspate and shells may be larger and thick- Family EULIMIDAE Troschel, 1853 Pseudosabinella McLean, new genus Type and only known species: Sabinella bakeri Bartsch, 1917 (synonyms: Alaba catalinensis Bartsch, 1920; Alaba serrana Smith & Gordon, 1948). See Abbott (1974:126, fig. 1382). Diagnosis: Shell thin, whorls rounded, with faintly an- gulate shoulder; suture deep; final lip inflated and pro- duced anteriorly; umbilical chink present; lip scars slight- ly raised, variable in position; early whorls dark brown, surface of teleoconch tan. Remarks: Regarding "Sabinella" bakeri, Waren (1992: 189) commented: "This species probably is an eulimid despite having a rather fragile and irregular shell. I have examined a specimen with dried soft parts and it has a ptenoglossate radula, similar to species of Eulirna. It can provisionally be placed in Eulitna." Pseudosabinella bakeri differs from typical Eulima in having rounded whorls, a projecting and flared final whorl and a brown cast to the shell. In contrast, typical species of Eulima, as defined by Waren (1984:43; 1992:179), have straight- sided shells, often with color banding. Waren elected not Page 40 THE NAUTILUS. Vol. 108, No. 2 to provide a genus for this species until the echinoderm host becomes known, but now endorses the need for a genus (pers. comm.). The species is a fairly common and characteristic member of the Californian faunal prov- ince, for which the provision of a genus may perhaps provide impetus toward the eventual discovery of its host. Familv MURICIDAE Rafinesque, 1815 Subfamily TROPHONINAE Cossmann, 1903 Ocenotrophon McLean, new genus Type and only known species: Miirex {Ocinebra?) pai- nei Dall, 1903. See Radwin & D'Attilio (1976:123, pi. 20, pi. 20, fig. 8). Diagnosis: Shell small (length about 15 mm), sculpture of numerous lamellar cords of unequal prominence, crossed by sharply raised ribs, forming nodes at inter- sections; canal short, closed. Remarks: The generic allocation of the type species has been uncertain, starting with Dall's original assertion that '"it resembles one of the austral trophons in miniature." Dall (1921:159, pi. 6, fig. 1) retained it in Tritonalia (ICZN rejected name for what is now Ocenebra Gray, 1847), evidently because of the sealed canal, which is characteristic of Ocenebra but not of trophonine genera. Radwin & D'Attilio (1976:123) followed Dall and re- tained it in Ocenebra but noted that: "The lamellose shell sculpture and short, stout form suggest affinity with the southern trophons (e.g. T. geversianus Pallas, 1774); the fused canal implies an ocenebrine relationship; and the radula favors neither of these possibilities." D'Attilio (1980:6) again treated the species and illustrated the rad- ula, concluding that the radula is in fact close to that of Trophon geversianus, and reallocated the species in Tro- phon. That allocation is less than satisfactory because T. geversianus , type species of Trophon, is large, has a dark colored interior, has an open canal and occurs in the intertidal zone of Argentina in the southern hemisphere. Kool (1993) treated the radula and anatomy of Trophon geversiansus. Ocenotrophon differs from species of Bor- eotrophon Fischer, 1884, (in which spiral sculpture is lacking or does not override the axial lamellae) and from species of Trophonopsis Bucquoy & Dautzenberg, 1882 (in which spiral sculpture overrides the axial lamellae) in having a thick final lip and in having a shorter, sealed canal rather than a relatively long open canal. Species of Boreotrophon and Trophonopsis tend to merge at the subgeneric level, whereas Ocenotrophon painei stands well apart from all other boreal trophonine species. Family BUCCINIDAE Rafinesque, 1815 Retimohnia McLean, new genus Type species: Mohnia frielei Dall, 1891. The holotype has been well figured by Kosuge (1972, pi. 13, fig. 3), showing also the characteristic pattern of the operculum. Included species: Urosalpinx carolinensis Verrill, 1884; Neptunea caelata Verrill & Smith, 1880; Sipho glyptus Verrill, 1882; Mohnia clarki Dall, 1907; Mohnia micra Dall, 1907; Mohnia hondoensis Dall, 1913; Mohnia ja- ponica Dall, 1913; Mohnia robusta Dall. 1913; Mohnia vernalis Dall, 1913; Cohis (Latisipho) clementinus Dall, 1919. Diagnosis: Shell small, fusiform; whorls rounded, canal short, twisted to left; periostracum thick, adherent; dom- inant sculpture of strong to faint axial ribs on upper half of whorl, becoming obsolete in some species at later growth stages; spiral sculpture of fine to moderate spiral incisions. Initial whorl of protoconch small, not project- ing; operculum subspiral, nucleus away from anterior edge. Remarks: Species grouped in Retimohnia have previ- ouslv been assigned to Mohnia Friele in Kobelt, 1878, by Dall (1913), Radwin (1972), and Tiba & Kosuge (1992). Bouchet & Waren (1985:205) only tentatively included in Mohnia three northeastern Atlantic species "for which we could not find a better genus, although we are not satisfied with their position here " The latter three species (Urosalpinx carolinensis Verrill, 1884, Neptunea caelata Verrill & Smith, 1880, Sipho glyptus N'errill, 1882) were treated in detail by Bouchet and Waren and are here allocated Retimohnia. The type species of Mohnia. Fusus mohni Friele, 1877, has been illustrated by Bouchet & Waren (1985:205, figs. 435, 482, 530-531) and by Tiba & Kosuge (1992:1). In species of Mohnia (and the subgenus Tacita Lus, 1971, as treated by Bouchet & Waren, 1985:210), the whorls are rounded and the suture deep; spiral sculpture pre- dominates, whereas axial sculpture is limited to fine axial riblets that produce a fine clathrate effect in early stages. The characteristic low protoconch was illustrated by Bouchet & Waren (1985). Axial sculpture of projecting ribs is the principal sculp- ture in Retimohnia, although in fi. clementinus it may be weak. The protoconch of Pacific species of Reti- mohnia is eroded in specimens I have examined, but is known from the three Atlantic species illustrated by Bouchet & Waren. The operculum is similar in both genera and the genera are evidently related. As is true of the Atlantic species treated by Bouchet & Waren, the Pacific species of both genera occur in deep water, at abyssal depths or the lower continental slope. Tiba and Kosuge (1992) placed three species in Mohn- ia that are assignable neither to Mohnia or Retimohnia: "Chrysodomus" brunneus Dall, 1877, "C." virens Dall, 1877, and "Mohnia" siphonoides Dall, 1913. I retain the first two in the genus Retifusus Dall. 1916 (type species Tritonium jessoense Schrenck, 1867, Japan), in which they were retained by Dall (as a subgenus of Plicifusus Dall, 1902). These two species are not restricted to deep water and have stronger spiral sculpture than species of Retimohnia. The operculum of Retifusus brunneus, as figured by Tiba & Kosuge (1992, fig. 10) does not have the nucleus sufficiently distant from the margin to be considered comparable to that of Retimohnia. Dall (1913: 502) remarked that the species he then described as J. H. McLean, 1995 Page 41 Mohnia siphonoides from 987 fathoms off the Pribilof Islands "would certainly be referred to Tritonofusus" [now Coins Roding, 1798] if it were not for the oper- culum. I assign the latter species to Coins, and assume that Dall mixed the operculum with that of a different species. With the e.xceptions above and the type species of Mohnia. all other species illustrated b> Tiba & Kosuge (1992) as Mohnia, are here considered t\pical species of Retimohnia. Bouchet & Waren (1985) described and illustrated several typical species of Mohnia. New species of Mohnia that I have recognized from the northeastern Pacific will be described separately. LITERATURE CITED Abbott, R. T. 1974. American Seashells, second edition. New York: Van Nostrand Reinhold, 663 pp. Bouchet, P. & .\. Waren 1985. Revision of the northeast Atlantic bathyal and abyssal Neogastropoda excluding Turridae (Mollusca, Gastropoda) Societa Italiana di Malacologia, Bollettino Malacologica, Suppiemento 1, 12.3- 269. Bouchet, P. & A. Waren. 1993 Re\ision of the northeast At- lantic bathyal and abyssal Mesogastropoda. Societa Ualiana di Malacologia, Bollettino Malacologico, Suppiemento 3, 579-840. Dall, W. H. 1903. Diagnoses of new shells from the Santa Barbara Channel, California. Proceedings of the Biological Society of Washington 16:171-176. Dall, W. H. 1913. New species of the genus Mohnia from the North Pacific. Proceedings of the Academy of Natural Sciences, Philadelphia, 65(2):501-504. Dall, W. H. 1921. Summary of the marine shell-bearing mol- lusks of the northwest coast of America, from San Diego, California, to the polar sea, mostly contained in the col- lection of the United States National Museum, with illus- trations of hitherto unfigured species. United States Na- tional Museum, Bulletin 112, 217 pp , 22 pis. D'Attilio, A. 1980 Trophon painei (Dall, 1903): An anom- alous murex. The Festivus 12(l):6-9. Kool,S. P. 1993. Phylogenetic analysis of the Rapaninae (Neo- gastropoda: Muricidae). Malacologia 35(2): 155-260. Kosuge, S. 1972. Illustrations of type specimens of molluscs described by William Healey Dall (North- Western Pacific Gastropoda). National Science Museum, Tokyo, 29 pis. and captions. Marincovich, L. N. 1977. Cenozoic Naticidae (Mollusca: Gas- tropoda) of the northeastern Pacific. Bulletins of American Paleontology 70(294): 165-494, pi. 17-42. McLean, J. H. 1992. Systematic review of the family Chor- istellidae (Archaeogastropoda: Lepetellacea) with descrip- tions of new species. The Veliger 35(4):273-294. Radwin, G. E. 1972. The systematic position of Urosalpinx carolinensis Verrill, 1884, with comments on the genus Mohnia Friele, 1878. Transactions of the San Diego So- ciety of Natural History 16(16):339-342. Radwin, G. E. and A D'Attilio. 1976. Murex shells of the world. Stanford University Press, 284 pp Tiba, R. and S. Kosuge. 1992. Genus Mohnia Friele, 1877. North Pacific Shells, no. 18, Occasional Publication of the Institute of Malacology, Tokyo, 26 pp. Waren, A. 1984. A generic revision of the family Eulimidae (Gastropoda, Prosobranchia). The Journal of Molluscan Studies, Supplement 13, 96 pp. Waren, A. 1992. Comments on and descriptions of eulimid gastropods from Tropical West America. The Veliger 35(3): 177-194. THE NAUTILUS 108(2):42-47, 1995 Page 42 Population Structure, Growth and Fecundity of Melampus bidentatus (Say) from Two Regions of a Tidal Marsh Complex in Connecticut Jessica A. Spelke Paul E. Fell' Lucille L. Helvenston Department of Zoology Connecticut College New London, CT 06320, USA ABSTRACT We examined papulations of the tidal marsh snail, Melampus bidentatus Say, within two regions of the Barn Island Wildlife Management Area in Stonington, Connecticut: a Spartina pat- ens-dominaied bayfront marsh and a restored impounded val- ley marsh covered by Spartina alterniflora and forbs. On the bayfront marsh, 87% of the snails collected from late May through early November were less than 8 mm in shell length, whereas 99% of the snails on the restored impounded marsh exceeded 8 mm in shell length. Snails from both marsh regions reared in the laboratory on Spartina alterniflora /(orh turf from the restored impounded marsh grew more rapidly than snails reared on Spartina patens turf from the bayfront marsh. In laboratory culture, Melampus exhibited a regular lunar (4- week) egg-laying periodicity with cycles of egg-laying occur- ring in late May, late June and late July. Not only did large snails (9.1-10.0 mm) from the restored impounded marsh de- posit nearly twice as many egg masses as small snails (6.1-7.0 mm) from the bayfront marsh, but the egg masses produced by the large snails were significantly larger than those laid by the small snails. Key words: Melampus bidentatus, tidal marsh snail, popu- lation structure, growth, fecundity. INTRODUCTION Melampus bidentatus Say, 1822, is a pulmonale snail that inhabits the higher elevations of tidal salt marshes along the Atlantic and Gulf of Mexico coasts of North America (Hausman, 1932; Holle & Dineen, 1957; Russell- Hunter et al., 1972). This snail often occurs in large numbers in regions of high marsh covered by Spartina patens (Ait) Muhi, 1817 (saltmeadow hay), Distichlis spicata (L.) Greene, 1887 (spikegrass) and Juncus spp.. ' To whom correspondence should be addressed. as well as in well-drained regions dominated by stunted Spartina alterniflora Loisel., 1807 (saltwater cordgrass). It is frequently a dominant member of the high marsh community (Russell-Hunter et al., 1972; Price, 1980; Hilbish, 1981; Fell et al, 1982; Joyce & Weisberg, 1986). It has been shown that on a restored impounded tidal marsh dominated by stunted Spartina alterniflora , the size of Melampus bidentatus is substantially larger than on bayfront, Spartina pa^fns-dominated marshes in the same system. On the restored impounded marsh, 83% of the snails exceeded 8 mm in shell length and the modal size was 9.1-10.0 mm. In contrast, on the undisturbed Spartina patens -dominated marshes, 95% of the snails were 8 mm in length with a modal size of 6.1-7.0 mm (FeW etal., 1991). Considering specific areas within these marshes, the same difference in the sizes of snails was observed. During the summer of 1990 and the late spring/ early summer of 1991, the most abundant size class of Melampus within the Spartina patens meadows of an undisturbed bayfront marsh was 6.1-7.0 mm, whereas the most abundant size class of this snail in well-drained areas of the restored impounded marsh covered by stunt- ed Spartina alterniflora and forbs (herbaceous plants other than grasses, rushes and sedges) was 9.1-10.0 mm. Because of this size difference, the shell-free biomass of Melampus in the first area was only about half that in the second, even though snail density was somewhat higher (Peck et al., 1993). Such size differences could be due in large part to quantitative and/or qualitative dif- ferences in the food resources available in the two marsh regions (Rietsma et al., 1988; Peck et al., 1993). How- ever, the size differences could also result to some extent from greater size specific predation (Vince et al., 1976; Joyce & Weisberg, 1986) and/or factors that promote greater longevity of snails (Crowe & Covich, 1990) on the restored impounded marsh. Melampus feeds exten- sively on detritus derived from the marsh grasses but also consumes algae and animal tissues (Rietsma et al., 1982; Thompson, 1984). One objective of the present study was to examine growth of Melampus from the J. A. Speike et al, 1995 Page 43 Spartina patens meadows of an undisturbed bayfront marsh and from stunted Spartina alterni flora -dominat- ed regions of tfie restored impounded marsh when sam- ples from both groups of snails were maintained in the laboratory on marsh turf from these two areas. Melam- pus is a simultaneous hermaphrodite with an ovotestis. It deposits egg masses on the surface of the marsh peat, as well as on the stems and leaves of grasses and on the shells of other Melampus. These egg masses are flattened hemispheres about 1 to 2 mm in diameter and approx- imately 0.5 mm high. Each egg mass may contain from about 500 to more than 1200 eggs (Hausman, 1932; Apley, 1970; Russell-Hunter et al., 1972). A preliminary study (Balboni-Tashiro et al., 1985) indicates that Melampus exhibits age (size) specific fecundity. Older snails pro- duce more egg masses containing more eggs. A second objective of this study was to compare egg mass depo- sition by large Melampus from stunted Spartina alter- nt^Jora -dominated regions of the restored impounded marsh and by smaller snails from the Spartina patens meadows of an undisturbed bayfront marsh. MATERIALS AND METHODS The study areas are part of the Barn Island Wildlife Management Area in Stonington, Connecticut, which is located in the southeastern corner of the state (figure 1). The marsh complex consists of a series of valley marshes that border Little Narragansett Bay. One study area was the westernmost valley marsh which was impounded in 1946-1947 in an attempt to create waterfowl habitat. This marsh (figure 1, IP) subsequently converted to a Typha angustifolia L., 1753 (narrow-leaved cattail- dominated brackish marsh. In 1978 a 5 ft. -diameter cul- vert was placed in the impoundment dike, and in 1982 a 7 ft. -diameter culvert was added. As a result of the re- established tidal exchange, typical tidal salt marsh an- giosperms have recolonized much of the area, with stunt- ed Spartina alterniflora and forbs providing most of the plant cover (Sinicrope et al., 1990). The other study area was an undisturbed Spartina patens -dominated bayfront marsh situated below the westernmost impoundment (Wequetequock Cove marsh, figure 1, WC). In order to examine the size structure of Melampus populations in the Spartina patens meadows of the bay- front marsh and stunted Spartina alterniflora-dominai- ed areas of the restored impounded marsh, all of the snails contained within two randomly placed 0.25m' quadrats were collected from each marsh area at about 30-day intervals from late May until early November. Shell lengths were measured to the nearest 0. 1 mm with vernier calipers. For experimental studies, small (6.1-7.0 mm) Melam- pus were collected from Spartina patens -dominated regions of the bayfront marsh and large (9.1-10.0 mm) snails were gathered from an area of the restored im- pounded marsh covered by stunted Spartina alterniflora and forbs. The snails were maintained in the laboratory on clipped marsh turf. They were placed in covered 11 IITTIE NARRAGANSETT BAY Figure 1. Map of the western portion of the tidal marsh complex within the Barn Island Wildlife Management Area in Stonington, Connecticut (short vertical arrow on inset at upper left) IP marks the collecting site on the restored impounded marsh and WC indicates the location of the Wequetequock Cove Marsh. cm-diameter finger bowls containing a piece of turf ap- proximately 6 X 4 X 1.5 cm situated on a 9 cm disc of Whatman No. 1 filter paper moistened with 15 ppt sea water. In most cases, 8 snails were placed in each bowl. The cultures were kept at about 21°C and exposed to natural photoperiod. The filter paper and bowls were changed every 2 days and the marsh turf was replaced every 6 days. For growth experiments, some of the snails from each marsh region were placed on Spartina patens turf taken from the bayfront marsh and others were cultured on Spartina alterniflora /forh turf from the restored im- pounded marsh. Each treatment was replicated 5 times (a total of 40 snails per treatment). The snails were mea- sured to the nearest 0.1 mm with vernier calipers at monthly intervals. The experiments were begun 8 June 1992 and continued until 1 November 1992. In order to quantify egg mass deposition, snails from each of the two marsh regions were cultured separately on Spartina patens turf from the bayfront marsh (15 snails per bowl in 1991 and 8 snails per bowl in 1992). Cultures were initiated 23 May 1991 and 20 May, 10 June and 25 June 1992. Each set of cultures was repli- Page 44 THE NAUTILUS, Vol. 108, No. 2 o SHELL LENGTH (mm) Q 7 8 9 10 11 12 13 SHELL LENGTH (mm) Figure 2-3. Size-frequency distribution of Melampus biden- tatus 2. In Spartina patens meadows of the bayfront marsfi during the reproductive period (20 May to 3 August 1992). N= 1024. 3. In regions of the restored impounded marsh covered by stunted Spartina alterniflora and forbs during the repro- ductive period (20 May to .3 August 1992). N = 702. cated 3 times. Newly deposited egg masses were removed and counted each day through 31 July. RESULTS On the bayfront marsh, 87% of the snails collected throughout the study were less than 8 mm in shell length, and the most abundant size class was 6.6-7.5 mm. In contrast, 99% of the snails on the restored impounded marsh exceeded 8 mm in shell length, and 10.6-1L5 mm was the most abundant size class. During the reproduc- tive period, the minimum length of Melampus on the bayfront marsh was 4.1-4.5 mm and the most abundant size class was 6.6-7.5 m (figure 2). Minimum length of snails from the restored impounded marsh during the reproductive period was 6.6-7.0 nmi, and less than 1% of the snails were of this size. The majority of snails (61%) were 10.1-11.5 mm in shell length (figure 3). Young-of- the-year snails were not observed on either marsh region during the study. Snails from both marsh regions reared in the laboratory on Spartina alterniflora /f orb turf from the restored im- pounded marsh grew more rapidly than snails reared on Spartina patens turf from the bayfront marsh (Table 1). Mann Whitney tests with values of tied rank (Zar, 1974) indicated that the difference in growth of Melampus from the bayfront marsh on turf from the two marsh regions was statistically significant (U = 0, df = 2, p = 0.01) as was the difference in growth of snails from the restored impounded marsh on these substrata (U = 23.5, df = 2, 0.05 > p > 0.02). In laboratory culture, Melampus exhibited a regular lunar egg-laying periodicity with cycles of egg-laying occurring in late May, late June and late July (figure 4). Egg-laying was usually initiated at the time of the full moon and continued for about 9 to 14 days. Deposition of egg masses ceased shortly before the new moon, and little or no egg-laying was observed between the new moon and subsequent full moon. Aggregation of snails was noted 2-3 days before egg-laying began. Egg masses were deposited most commonly on the surface of the peat, but they were also attached to the stems of Spartina patens and occasionally to the shells of other Melampus. Egg-laying by Melampus collected from the bayfront marsh on 20 May 1992 was not in synchrony with that of other snails gathered from the two marsh regions. For this group, egg-laying began 9 June, six days before the full moon, and continued at a low rate until 25 June, ten days past the full moon. A second cycle of egg-laying was initiated 6 July, eight days before the full moon, and lasted until 15 July. The snails from the restored impounded marsh de- posited nearly twice as many egg masses as those from the bayfront marsh (Table 2). During the summer of 1992, large snails (9.1-10.0 mm) from the restored im- pounded marsh maintained under laboratory conditions produced an average of 128 egg masses per animal dur- ing the reproductive season, whereas small snails (6.1- 7.0 mm) from the bayfront marsh deposited an average of 63 egg masses per animal over the same span of time. Analysis of the data from the third spawning period, for which there were three different sets of snails from each marsh region, indicated that the difference in the number of egg masses deposited by the larger snails of the restored impounded marsh and the smaller snails of the bayfront marsh is statistically significant (two-sample t-test: t = 2.9, df = 15, p = 0.012). The mean size of egg masses produced by snails from the restored impounded marsh was 3.3 mm ± 0.42 mm SD (N = 31) and that of snails from the bayfront marsh was 2.5 mm ± 0.39 mm SD (N = 28). Using a two- .sample t-test, the difference was found to be highly sig- nificant (t = 7.58, df = 56, p < 0.0001). During 1991, large snails from the restored impounded marsh, maintained at a density of 15 snails per bowl. J. A. Spelke et al., 1995 Page 45 Table 1. Growth (increase in shell length) of Melampus hidentatus (Say) from a restored impounded marsh (IP) and an undisturbed bayfront marsh (Wequetequock Cove Marsh, WC) on turf from these two marshes in the laboratory. The experiment extended from 8 June to 1 November 1992. Mean size Origin of Mean initial size (mm) ± SD at end M ?an growth snails Type of marsh turf (mm) ± SD of experiment (mm) % increase WC S. patens (WC) 6.6 ± 0.27 7.7 ± 0.35 11 17 S. alterni flora /loihs. (IP) 6.6 ± 0.28 8.2 ± 0.54 16 24 IP S. patens (WC) 9.6 ± 0.20 10.2 ± 0.37 0,6 6 S. alterniflora/iorhs (IP) 9.7 ± 0.29 10,5 ± 0,46 0,8 8 deposited an average of 96 egg masses per animal during the first cycle of egg-laying and the study was terminated before the cycle was completed. In contrast, during 1992, large snails cultured 8 per bowl produced an average of no more than 71 egg masses per animal for any cycle. The relationship between snail density and egg-laying should be systematically investigated. DISCUSSION On the Barn Island marshes in southeastern Connecticut, Melampus exhibits a clear lunar periodicity in egg-lay- ing. Egg mass deposition begins at about the time of the full moon and continues until shortly before the time of the following new moon. This pattern of egg-laying dif- fers from that described for snails at the Little Sippew- isset Marsh in Falmouth, Massachusetts (Cape Cod). Sev- eral studies conducted over a 10-yr. period at that location have demonstrated a semilunar pattern of egg-laying by Melampus (Apley, 1970; Russell-Hunter et al., 1972; Table 2. Production of egg masses by Melampus hidentatus (Say) in the laboratory Small snails were collected from an undisturbed bayfront marsh (Wequetequock Cove Marsh, WC) and large snails were gathered from a restored impounded marsh (IP). Spawn- ing period Mean number of egg masses/snail ± SD Date of collection 91- 10 0 mm (IP) 6.1- 7 0 mm (WC) Ratio 23 May 1991 1 96 ± 6.0 44 ± 1.7 2.2 20 May 1992 1 2 3 all 10 ± 6.1 69 ± 9.3 50 ± 7.4 129 ± 22.7 0 18 ± 7.0 41 ± 7.1 58 ± 14.0 2.2 10 June 1992 2 3 all 70 ± 6.6 37 ± 11,0 107 ±17 4 39 ±3,1 32 ± 2.7 71 ± 4.0 1.5 25 June 1992 3 56 ± 2.7 31 ± 13.5 1.8 All cultures 388 204 1.9 Price, 1979). The semilunar periodicity is well defined with egg-laying normally taking place for about 4 days beginning on about the second day following a new or full moon (figure 5). Spring tides that occur at these times often immerse the newly deposited egg masses and cover them with a layer of fine detritus that keeps the eggs/ embryos moist during the period of neap tides (Russell- Hunter et al, 1972). The pattern of egg-laying described here does not appear to be an artifact resulting from laboratory con- LUNAR PHASES HEIGHT OF HIGH TIDES EGG-LAYING PERIODS n FULL o A LITTLE SIPPEWISSET MARSH B BARN ISLAND MARSH Figures 4-5. Egg-laying by Melampus hidentatus 4. Snails from the restored impounded marsh in relation to the occur- rence of full moons. Each point represents the average number of egg masses produced by three bowls of snails. Cultures were initiated 23 May 1991 and 20 May, 10 June and 25 June 1992. 5. Snails on the Little Sippewisset Marsh in Falmouth, Mas- sachusetts (after Russell-Hunter et al., 1972) and on the Barn Island marshes in Stonington, Connecticut in relation to phases of the moon and the relative height of high tides (marsh sub)- mergence). Page 46 THE NAUTILUS, Vol. 108, No. 2 ditions. First, egg-laying by Melampus maintained in the laboratory follows the same rhythm as that of field populations of this snail (Russell-Hunter et ai, 1972 and our unpublished observations). Second, cultures of Me- lampus were initiated a few days before both new and full moon phases, and the pattern of egg-laying was the same in both cases: egg-laying began at the time of the full moon but not at the time of the new moon. It would be interesting to simultaneously compare egg-laying by Melampus from Barn Island and Cape Cod under iden- tical laboratory conditions. How the timing of reproduction by these animals is set is unknown. When Melampus spawns with a lunar periodicity, each spawning cycle is about 2-3 times lon- ger than when it spawns with a semilunar rhythm and the period between spawning cycles is nearly doubled. Evidently more eggs are produced per spawning cycle and more time may be required to recover before com- mencing another cycle of egg-laying. Melampus is a largely terrestrial snail; its marsh hab- itat may be unsubmerged for more than 95% of the time. However, this snail possesses an aquatic veliger larva (Russell-Hunter et al., 1972). Although egg-laying is usu- ally initiated during spring tides both on the Little Sip- pewisset and Barn Island marshes, flooding of the marsh surface is not required for egg-laying to occur. On the other hand, the hatching of veligers and their subsequent settlement back on to the marsh depend upon inundation of the marsh during spring tides (Russell-Hunter et al., 1972). The observed differences in the patterns of egg- laying by the two populations of Melampus do not ap- pear to necessarily affect later stages in the life cycle of the snail. For example, larval hatching may occur from about 10 days to 24 or more days after egg mass depo- sition, depending upon tidal submergence (Russell-Hun- ter, 1972). The minimal reproductive size of Melampus is from about 5 mm shell length (Apley, 1970) to 5.8 mm shell length (Balboni-Tashiro et al., 1985). However, not all snails of this minimal size reproduce (Balboni-Tashiro et al., 1985). Most of the Melampus collected from both marsh regions at Barn Island were greater than 6 mm in shell length. In the present study, Melampus maintained in labo- ratory culture produced an average of from 63 to 128 egg masses per animal during the reproductive season, depending upon snail size. On the other hand, Russell- Hunter e< a/., (1972) reported that under their laboratory conditions, snails taken from the Little Sippewisset Marsh produced on average only 39 egg masses during the reproductive period. This is consistent with the obser- vation that egg-laying by the Cape Cod population of Melampus is typically restricted to fewer days compared to the populations of this snail at Barn Island. On Cape Cod, there are about 12 to 16 days of egg-laying by Melampus each year (Ru.ssell-Hunter et al., 1972), whereas at Barn Island snails deposit egg masses during about 20 to 30 days within the reproductive period. How- ever, the snails normally aggregate prior to mating and egg-laying (Apley, 1970; Russell-Hunter et al., 1972); and the number of egg masses produced may also be influenced by snail density in laboratory cultures and under field conditions. Not only did the large snails from the restored im- pounded marsh at Barn Island deposit more egg masses than the smaller snails from the nearby bayfront marsh, but the egg masses were significantly larger. In gastro- pods for which size specific fecundity has been well doc- umented, the larger egg capsules/ masses produced by large individuals contain more eggs than the smaller egg capsules/masses produced by small individuals (Hendler & Franz, 1971; Spight & Emlen, 1976). In addition, a preliminary study by Balboni-Tashiro et al. (1985) has shown that older (larger) Melampus deposit more eggs per mass than younger (smaller) snails. Therefore even though egg numbers were not determined in the present study, it seems highly probable that large Melampus from the restored impounded marsh produced more eggs than smaller ones from the bayfront marsh, indicating that fecundity is size specific for this species. However, interpretation of the data is complicated by the fact that the different sized snails were taken from different marshes. Although snails from the two marshes were maintained on the same substratum (food resources) in the laboratory, an influence of prior nutrition on repro- duction beyond a larger size, resulting from a more rapid growth rate, cannot be ruled out (Spight & Emlen, 1976). Differences in growth rates of snails in the laboratory on turf from the two marsh regions were significant. Melampus from both marsh regions grew more rapidly on Spartina alterniflora/iorh turf from the restored im- pounded marsh than on Spartina patens turf from the bayfront marsh. These results are consistent with the hypothesis that the snails on the restored impounded marsh grow more rapidly than the snails on the undis- turbed bayfront marsh because of a larger/better food resource. It would be interesting to determine which potential foods are being used by Melampus on the two marshes by examination of gut and fecal pellet contents (Thompson, 1984). It is well documented that Melampus prefers old detritus with a low phenolic content and grows more rapidly on this type of diet (Rietsma et al., 1988). However, other factors, such as the structural characteristics (toughness and texture) of the food and/ or its moisture content, may also importantly affect pal- atability. Finally, large microbial populations are often found on detritus particles and they constitute a rich energy source for organisms capable of removing them (Odum & De La Cruz, 1967). In conclusion, snails inhabiting a restored impounded tidal marsh in southeastern Connecticut are larger, grow more rapidly and produce more eggs than snails occu- pying an undisturbed bayfront marsh within the same marsh complex. The rapid growth and high fecundity of Melampus on the restored impounded marsh indicate that the marsh has become a highly favorable habitat for this snail and provide additional evidence that res- toration efforts have been successful. J, A. Spelke et al., 1995 Page 47 ACKNOWLEDGMENTS The authors are indebted to Dr. Robert Askins for help with the statistical analysis and to Myron Peck for pro- ducing figures 4 and 5. This study was supported by a grant from the Andrew Mellon Foundation. LITERATURE CITED Apley, M. L. 1970. Field studies on life history, gonadal cycle and reproductive periodicity in Melampus bidentatus (Puimonata: Ellobiidae) Malacologia 10(2):381-397. Baiboni-TashircJ.. P. WalbornandB. Crise. 1985. Diapause, degrowth, and age of first reproduction in Melampus bi- dentatus Biological Bulletin 169533 Crowe, T. A. and A. P. Covich. 1990. Predator-induced life- history shifts in a freshwater snail. Science 247949-951. Fell, P. E., N C Olmstead, E. Carlson, W Jacob, D Hitchcock and G. Silber. 1982 Distribution and abundance of mar- coinvertebrates on certain Connecticut tidal marshes, with emphasis on dominant molluscs. Estuaries 5:234-239. Fell, P. E , K A Murphy, M. A. Peck, and M. L. Recchia. 1991. Re-establishment of Melampus bidentatus (Say) and other macroinvertebrates on a restored impounded tidal marsh: comparison of populations above and below the impoundment dike. Journal of Experimental Marine Biology and Ecology 15233-48 Hausman, S. A 1932. A contribution to the ecology of the salt marsh snail, Melampus bidentatus Say. The American Naturalist 66:541-545. Hendler, G. and D R. Franz. 1971 Population dynamics and life history of Crepidula convexa Say (Gastropoda: Pros- obranchia) in Delaware Bay. Biological Bulletin 141:514- 526. Hilbish, T. J 1981. Latitudinal variation in freezing tolerance of Melampus bidentatus (Say) (Gastropoda: Puimonata). Journal of E.xperimental Marine Biology and Ecology 52: 283-297. Holle, P. A. and C. F. Dineen. 1957. Life history of the salt- marsh snail, Melampus bidentatus Say. The Nautilus 70: 90-95. Joyce, A. A. and S. B. Weisberg. 1986. The effects of predation by the mummichog, Fundulus heteroclitus (L), on the abundance and distribution of the salt marsh snail, Me- lampus bidentatus (Say). Journal of Experimental Marine Biology and Ecology 100:295-306. Odum, E P and A. A. De La Cruz 1967. Particulate organic detritus in a Georgia salt marsh ecosystem. Estuaries 83: 383-388. Peck, M. A., P. E. Fell, E. A. Allen, J. A. Gieg, C. R. Guthke, and M. D. Newkirk 1994. Evaluation of tidal marsh restoration: comparison of selected macroinvertebrate populations on a restored impounded valley marsh and an unimpounded valley marsh with the same salt marsh sys- tem in Connecticut, USA, Environmental Management 18: 283-293. Price, C. H. 1979 Physical factors and neurosecretion in the control of reproduction in Melampus (Mollusca: Puimon- ata). Journal of Experimental Zoology 207:269-282. Price, C. H. 1980. Water relations and physiological ecology of the salt marsh snail Melampus bidentatus . Journal of Experimental Marine Biology and Ecology 45:51-67. Rietsma, C S., I. Valiela, and A. Sylvester-Serianni 1982. Food preferences of dominant salt marsh herbivores and detritivores. Marine Ecology 3(2): 179-189. Rietsma, C. S., I. Valiela, and R. Buchsbaum. 1988. Detrital chemistry, growth, and food choice in the salt marsh snail (Melampus bidentatus). Ecology 69:261-266. Russell-Hunter, W D., M. L. Apley, and R. D. Hunter 1972. Early life history of Melampus and the significance of semilunar synchrony. Biological Bulletin 143:623-656. Sinicrope, T L., P G Hine, R S. Warren, and W. A. Niering. 1990 Restoration of an impounded salt marsh in New England. Estuaries 13(l):25-30. Spight, T. M. and J. Emien. 1976. Clutch sizes of two marine snails with a changing food supply. Ecology 57: 1 162-1 178. Thompson, L. S. 1984. Comparison of the diets of the tidal marsh snail, Melampus bidentatus and the amphipod, Orchestia grillus. The Nautilus 98(l):44-53 Vince,S., I. Valiela, N. Backus and J. M. Teal. 1976. Predation by the salt marsh killifish Fundulus heteroclitus (L.) in relation to prey size and habitat structure: consequences for prey distribution and abundance. Journal of Experi- mental Marine Biology and Ecology 23:255-266. Zar, J. H. 1974. Biostatistical analysis. Prentice-Hall, Engle- wood Cliffs, New Jersey, 620 pp. NOTICE THIS MATERIAL MAY BE PROTECTED BY LAW (TITLE 17 U.S. CODE) THE NAUTILUS 108(2):48, 1995 Page 48 News and Notices SMITHSONIAN FUNDS FOR MALACOLOGY STUDENTS The Division of Mollusks, Department of Invertebrate Zoology, National Museum of Natural History, Smith- sonian Institution announces the availability of the Houb- rick-Rosewater Fellowship (up to $750) to be awarded to graduate students of systematic malacology This award provides support for students conducting systematic stud- ies of Mollusca (leading to publication) who require ac- cess to collections and libraries of the Division of Mol- lusks, National Museum of Natural History. Funds can be used for travel, subsistence, and research costs. Inter- ested students should submit a succinct proposal (1-2 pages), including budget, with indication of any match- ing funds, and a support letter from faculty advisor(s). The application deadline is March 15, 1995. Award(s) will be announced on April 15, 1995. Applications should be sent to: Dr. Clyde F. E. Roper Division of Mollusks/NHB stop 118 National Museum of Natural History Smithsonian Institution Washington, DC 20560 USA THE AMERICAN MALACOLOGICAL UNION ANNUAL MEETING The 1995 meeting of the American Malacological Union will be held at the University of Hawaii at Hilo, in Hilo, Hawaii, June 8-12, 1995 (arrive Hilo June 7 — leave June 13). The meeting is a celebration of islands, beginning with a keynote address by Hawaii's well known naturalist and photographer Bill Mull, who will describe with tales and slides some of the wonders of Hawaiian animals and plants, and especially land shells. Three symposia are scheduled: biogeography convened by Gustav Paulay, cephalopods convened by Richard Young, and conser- vation convened by Elaine Hoagland. Plan to contribute a paper and/or poster for other sessions. The annual auction of books and shells will be exciting. Social events include the President's Welcome, a sunset reception at Lyman House Museum, a barbecue, and final banquet. Field trips on June 1 1 include tidepools and snorkeling, lava tubes, rainforest, kipukas (oases with snails on Mauna Kea), and the volcano. Accommodation will be in University of Hawaii at Hilo dormitories on campus with a meal plan. Accommoda- tion will run $25 per night per bed in a four room suite, and the meal plan, covering most meals and coffee breaks from arrival June 7 to leaving June 13 will be about $150. If you prefer a hotel, you may make reservations from a list with rates to be included with registration forms. Van service to and from the airport will be provided. Special day trips for accompanying guests and family are scheduled. Please indicate on a postcard, by FAX or e-mail if you wish to receive a registration form and plan to attend: E. 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THE NAUTILUS Volume 108, Number 3 April 19, 1995 ISSN 0028-1344 A quarterly devoted to malacology. ,7/ c Iruiiiution Hoio, IbM 043 EDITOR-IN-CHIEF Dr. M. G. Harasev\Tch Division of Molliisks National Museum of Natural Histor\' Smithsonian Institution Washington, DC 20560 ASSOCIATE EDITOR Dr. R. Tucker Abbott .■\merican Malacologists, Inc. P.O. Box 2255 Melbourne, FL 32902 CONSULTING EDITORS Dr. Riidiger Bieler Department of Invertebrates Field Museum of Natural History Chicago, IL 60605 Dr. Robert T. Dillon, Jr. Department of Biology College of Charleston Charleston, SC 29424 Dr William K. Emerson Department of Living Invertebrates The American Museum of Natural History NewYork, NY 10024 Dr Robert Hershler Division of Mollusks National Museum of Natural History Smithsonian Institution Washington, DC 20560 Mr. Richard I. Johnson Department of Mollusks Museum of Comparative Zoology Harvard University Cambridge, MA 02138 Dr. Aurele I^a Rocque Department of Geology The Ohio State University Columbus, OH 43210 Dr. James H. McLean Department of Malacology Los Angeles ("ounty Museum of Natural History 900 Exposition Boulevard Los Angeles, C A 90007 Dr. Arthur S. Merrill % Department of Mollusks Museum of Comparative Zoology Harvard University Cambridge, MA 02138 Dr. Paula M. Mikkelsen Department of Malacology Delaware Museum of Natural Histor\' P.O. Bo.x 3937 Wilmington, DE 19807 Dr. Donald R. Moore Division ol Marine Geology and Geophysics Rosenstiel School of Marine and Atmospheric Science University of Miami 4600 Rickenbacker Causeway Miami, FL 33149 Dr. Ciustav Paulay Marine Laboratory University of Guam Mangilao, Guam 96923 Mr. Richard E. Petit RO. Box 30 North Myrtle Beach, SC 29582 Dr. Edward J. P(>tuch Department of Geology Florida Atlantic University Boca Raton, FL 33431 Dr. David H. Stansbery Museum of Zoology The Ohio State University Columbus, OH 43210 Dr. Ruth D. Tinner Department of Mollusks Museum of C^omparative Zoology Harvard University Cambridge. MA 02138 Dr. Geerat J. Vermeij Department of Geology University of C'alifornia at Davis Davis, CA 95616 SUBSCRIPTION INFORMATION The subscription rate per volume is US $28.00 for individuals, US $45.00 for institutions. Postage outside the United States is an additional US $5.00 for surface and US $12.00 for air mail. All orders should be accompanied by payment and sent to: THE NAUTILUS, P.O. Box 7279, Silver Spring, MD 20907-7279, USA. Change of address: Please inform the publisher of your new address at least 6 weeks in advance. All communications should include both old and new addresses (with zip codes) and state the effective date. THE NAUTILUS USSN 002S-1344) is published quarterly Iiy Trophon Corporation, 891 1 Alton Parkway, Silver Spring, MD 20910. Second Class postage paid at SiKer Spring. MD and additional mailing offices. POSTMASTER: Send address changes to: THE NAUTILUS PO. Box 7279 Silver Spring. MD 20907-7279 TH E€:7NAUTI LUS CONTENTS Volume 108, Number 3 April 19, 1995 ISSN 0028-1344 Nathan S. Dayan Robert T. Dillon. Jr. Florida as a Biogeographic Boundary: Evidence from the Population Genetics of Littorina irrorata 49 Mark E. Gordon Venustaconcha sima (Lea), an Overlooked Freshwater Mussel (Bivalvia: Unionoidea) from the Cumberland River Basin of Central Tennessee 55 M. G. Harasewych Alan R. kabat Richard S. Houbrick (1937-1993): Biographical Sketch and Malacologicai (contributions 61 Yu. A. Filippova D. N. Khromov K. N. Nesis I. V. Nikitina New Data on the Distribution and Morphology of some Western Indian Ocean Sepiid Cuttlefishes (Cephalopoda: Sepiida) 67 James H. McLean Three Additional New Genera and two Replacement Names for Northeastern Pacific Prosobranch Gastropods 80 News and Notices WESTERN SOCIETY OF MALACOLOGISTS ANNUAL MEETING The twenty-eighth Annual Meeting of the Western So- ciety of Malacologists will be held at the Resort at Chena Hot Springs, near Fairbanks, Alaska, from June 2 to June 6, 1995. The agenda will include contributed papers on all areas of molluscan studies, freshwater, marine, ter- restrial, living and fossil. Symposia on ecology and pa- leoecology are being organized \s ith the help of Howard Fader and David Hopkins. Also in the planning stages are an auction, reprint sale, and banquet. The University of Alaska Museum Aquatic Collection will be available for visitors before and after the meeting. For more in- formation contact WSM President, Nora R. Foster, Uni- versity of Alaska Museum, 907 Yukon Drive, Fairbanks, Alaska 99775 USA. Phone: (907) 474-9557. E-Mail: FYA- QUA(a) aurora.alaska.edu. Or contact Conference and Special Events, 117 Eielson Building, University of Alas- ka Fairbanks, Fairbanks, Alaska 99775. INDO-PACIFIC MALACOLOGICAL MEETING PRELIMINARY NOTICE The Council of the Malacological Society of Australasia has agreed to hold an international meeting on mollusks of the Indo-Pacific region in Perth, Western Australia, in January or February 1997. This notice is intended to alert malacologists worldwide of the upcoming meeting and to solicit suggestions for possible symposia; all sug- gestions of possible topics are welcome. The meeting will include all aspects of malacology, including mollusks of marine, freshwater and terrestrial habitats. If sufficient interest develops a post-conference field trip to the south- west of Western Australia can be arranged. As plans develop the conference will be advertised in malacolog- ical journals. To be placed on the list for direct receipt of future notices please write: Dr. F. E. Wells, Western Australian Museum, Perth 6000. Western Australia (Fax: 61-9-328-8686). NEW MOLLLISK JOURNAL LAUNCHED The Malacological Society of Australia has a history of over forty years of service to the study of mollusks in Australia, and has published the Journal of the Mala- cological Society of Australia since 1957. Like most sci- entific societies, including AMSA, the Malacological So- ciety has been continuously evolving. In recent years there has been increased emphasis and membership of the Society in New Zealand and countries to the north of Australia. In recognition of its changing role the So- ciety changed its name to the Malacological Society of Australasia at the annual general meeting held in Sydney in November 1993. With the change in name and emphasis of the Society a re-examination of the role oi the Journal of the Mala- cological Societtj of Australia was appropriate. After extensive di.scussion within the MSA Council it was de- cided to broaden the content of the Journal and change its name to Molluscan Research to reflect the modifi- cations. The cover and pages have been redesigned. To ensure continuity, the volume numbering remains the same. Theoretical papers considering any molluscan topic are welcome for consideration for publication in Molluscan Research. Papers considering specific geographical areas or new taxa should be restricted to the Indo-West Pacific region, which includes all of Australia. The Journal has previously been an annual publication, but if sufficient quality papers are available, Molluscan Research will appear twice a year. The first issue of Molluscan Research has been published in early November, 1994. For further information about Molluscan Research please contact the editor. Dr. F. E. Wells, Western Australian Museum, Perth, WA 6000, Australia. For information afjout the Malacological Society of Australasia contact the Department of Malacology, Australian Museum, P.O. Box A285, Svdnev south, NSW 2000, Australia. THE NAUTILUS 108(3):49-54, 1995 Page 49 Florida as a Biogeographic Boundary: Evidence from the Population Genetics of Littorina irrorata^ [Nathan S. Dayan Graduate Program in Marine Biology Grice Marine Biological Laboratory 205 Ft. Johnson. Charleston, SC 29412 USA Robert T. Dillon. Jr.- Department of Biology C;ollege of Charleston C;harleston. SC 29424 USA ABSTRACT The marsh periwinkle, Littorina irrorata (Say, 1822), ranges from New York to Texas with an apparently large disjunction around southern Florida. We examined gene frequencies at eight polymorphic enzyme loci in populations from Virginia, South Carolina, Atlantic Florida, Gulf Florida, and Louisiana. The within-population deviation from Hardv -Weinberg ex- pectation was small at all loci examined (F|,,=0 049) Small but significant gene frequency differences among populations were detected at four loci, indicating some isolation by distance Hierarchical gene diversity analysis suggested, however, that very little of the population divergence present (Fy, =0.033) is attributable to a division between coasts (Ft ;h=0.004). Nei ge- netic distances calculated between pairs of sites sharing the same coast were comparable to such distances between Atlantic and Gulf pairs Thus we find no evidence that the F"lorida peninsula constitutes a significant barrier to L irrorata, in spite of the apparently extensive gap in its range. We offer three hypotheses, not mutually exclusive: that the barrier is not real, that it was broached in the not-too-distant past, or that bal- ancing selection may be ongoing to hold polymorphisms con- stant at multiple enzyme-encoding loci simultaneously. Keij words: Gastropoda, electrophoresis, allozymes, range dis- junction, balancing selection INTRODUCTION Shallow, protected regions along the southern Atlantic coast of the United States and the northern coast of the Gull of Mexico are characterized by temperate climate, depositional environment, and the dominant salt marsh cordgrass, Spartina. The molluscan faunas of these two regions are described as "Carolinian (Rehder, 1954; Coomans, 1962), sharing over 60% of their shallow water gastropod species (query to database of Rosenberg, 1993). However, striking climatic and geological variation along ' Contribution number 120 from the Grice Marine Biological Laboratory ^ Address for Correspondence the Florida peninsula seems to impose a biogeographic boundary between these regions (Briggs, 1974). Due to tropica! conditions, mangroves replace salt marshes in protected bays and estuaries around 27°- 29° N latitude (Kangas & Lugo, 1990). This transition occurs between Cedar Key and Tampa Bay on the Gulf coast of Florida, and between St. Augustine and Cape Canaveral on the Atlantic coast. Coralline sands become the dominant sed- iment type along south Florida coasts, replacing terrig- enous silt and mud. Scheltema (1989) surveyed the ranges of 88 mesogas- tropod and neogastropod species, dividing the western Atlantic coast into eight regions from Arctic Canada to Brazil. He reported that 58 ol 72 species inhabiting his "region IV ' (Beaufort, North Carolina to Miami, Flor- ida) also occurred in his "region V" (Gulf of Mexico). However, 18 of the 58 shared species did not occur in Scheltema s "region VI", encompassing the southern tip of Florida and the Greater Antilles. For at least these 18 species (including such common species as Fasciolaria hunteria. Poliniccs duplicatus, and Littorina irrorata) Florida would seem to constitute a potential barrier This general distributional pattern extends beyond the near- shore molluscan fauna to include many other elements of the flora and fauna of the southeastern United States. Recent molecular techniques have commonly detected substantial genetic differentiation between animal pop- ulations of the southern Atlantic coast and those of the northern Gulf of Mexico. Most of the 19 such species for which mtDNA surveys have been completed, including horseshoe crabs, toadfish, black sea bass, diamondback terrapins, and seaside sparrf)ws, show distinct differen- tiation associated with the Florida peninsula (Avise, 1992). To this list could be added the coastal North American tiger beetle, Cicindela dorsalis, where mitochondrial DNA sequence data were used to assign four subspecies to either an Atlantic coastline lineage or a Gulf of Mexico coastline lineage (Vogler & DeSaile, 1993). Significant divergence at enzyme-encoding loci has been detected between Atlantic and Gulf populations of such diverse taxa as the sea anemone, Bunodosoma cavernata (McCommas, 1982), and the marsh crab Sesarnia reti- culatum (Felder & Staton, 1994). Page 50 THE NAUTILUS, Vol. 108, No. 3 Figure 1. The range of Littorina irrorata (shaded), showing sample sites for the present study. No Florida disjunction is apparent in the range of the American oyster, Crassostrea virginica, nor was any di- vergence in allozyme frequencies detected by Buroker (1983) in his survey of 19 populations from Massachusetts to Texas. Thus the report of substantial mtDNA diver- gence between Atlantic and Gulf oyster populations by Reeb and Avise (1990) was greeted with unusual interest. Karl and Avise s (1992) re-e.\amination of the issue using restriction fragment length polymorphisms in anony- mous single-copy nuclear genes confirmed the mtDNA results, suggesting that Florida does, in fact, constitute a barrier to the disper.sal of oysters. Karl and Avise went on to propose that the similarity in allozyme frequencies reported by liuroker might result from balancing selec- tion at multiple enzyme-encoding loci. The importance of natural selection to the preservation of enzyme poly- morphism has been a central question in evolutionary biology for over 25 years (Lewontin, 1991). Thus if the findings of Karl and Avise can be generalized beyond oysters, there will be implications for our understanding of evolution as a whole. The purpose of this study is to document divergence in allozyme frequency between Atlantic and (iulf pop- ulations of an heretofore genetically unsurveyed inter- tidal mollusk, the marsh periwinkle Littorina (or Lit- toraria) irrorata (Say, 1822). The snail is primarily an inhabitant of salt marshes dominated by the cord grass Spartina altcrniflora, leaving the marsh surface with the incoming tide to climb the vegetation. It ranges from Jamaica Bay, Long Island, New York (Jacobson, 1965) to Port Isabel, Texas (Bequaert, 1943), with a disjunction around southern Florida (Figure 1). As such it would seem an excellent candidate for an attempt to confirm the Karl and Avise phenomenon. Littorina has been the object of considerable popu- lation genetics research worldwide, with much effort directed toward questions of systematics (Ward & War- wick, 1980; Maestro et al . 1991; Boulding el ai, 1993; Zaslavskaya et al., 1992). Other workers have prospected for environmental dines (Newkirk & Doyle, 1979;Janson & Ward, 1984; Johannesson et al., 1993), founder effects (Janson, 1987), or correlates of heterozygosity (Noy et al., 1987; Foltz et al , 1993). However, the only previous examination of L. irrorata allozymes, prior to this report, was that of Berliner (1981) in a Virginia salt marsh. He found no significant difference in heterozygosity between young and old size classes, but noted a lower value of heterozygosity in snails of median age. Berliner's study did not extend beyond his single population. MATERIALS AND METHODS Approximately 50-60 L. irrorata per site were collected from three sites on the Atlantic coast of the southeast United States and two from the Gulf of Mexico: the Virginia Institute of Marine Science Laboratory (VIMS) at Wachapreague, Virginia, the Grice Marine Biological Laboratory (GMBL) at Charleston, South Carolina, Cres- cent Beach, Florida (CBFL), Tampa Ba\ , Florida (TBAY), and the Louisiana University Marine Consortium Lab- oratory (LUMC) at Cocodrie, Louisiana (Figure 1) All sites except TBAY were typical salt marshes dominated by the salt marsh cord grass, S. alterniflora. The Tampa Bay site was a small patchy area of S. alterniflora growing on sand, rather than mud. At all sites, the snails were found on the stalks of the salt marsh cord grass or on the substrate at the base of the stalks, generalK in the mid- marsli to high marsh areas. The snails were transported to Charleston alive, where tissues were frozen at — 60°C in 150-300^1 of 0.05 M Tris Tissue buffer pH 7.5 (Dayan, 1994). Horizontal protein electrophoresis was conducted us- ing methods and equipment previously described (Dil- lon, 1985; 1992; Dayan, 1994). The 12% starch gels were a 1:1 mixture (by volume) of Electrostarch (Otto Hillar, Madi.son, Wl) and Sigma starch (Sigma C'hemical, St. Louis, MO). We initially compared the zymograms of individuals from GMBL and LUMC using 19 enzyme stains and 9 buffer systems. Results were poor or unin- terpretable for six enzyme-s and appeared invariable for seven others. LUtimately we were able to resolve allo- zymes interpretable as the products of codominant Men- delian alleles at eight loci (encoding six enzymes) using N. S. Dayan and R. T. Dillon, Jr., 1995 Page 51 Table 1. Allele frequencies at eight enzyme-encoding loci in five populations of Liltorina irrorata. (n) = Sample size Locus Estl (n) Est2 (n) Pgm (n) Mpi (n) Isdhl (n) Isdh2 (n) Sdh (n) Lap All- ele VIMS Population GMBL CBFL TBAY LUMC A B C A B A B C D A B A B C 0.571 0 429 0.000 56 0,984 0.016 61 0.648 0.270 0.082 0.000 61 0 992 0 008 63 0 934 0.057 0.008 61 A 0.992 B 0.008 60 A B C A B C D (n) 0814 0.161 0.025 59 0917 0.075 0.008 0 000 60 0.689 0.292 0.019 53 0,989 0.011 44 0 441 0.480 0.078 0.000 51 0.952 0.048 83 0.965 0.035 0.000 57 0.990 0.010 50 0.788 0.205 0.008 66 0.700 0.255 0.036 0.009 55 0.638 0.362 0.000 58 0 992 0,008 62 0 490 0.288 0.202 0.019 52 0.992 0.008 62 0.877 0.098 0.025 61 1 000 0.000 62 0.856 0.136 0.008 59 0 900 0 067 0 025 0.008 60 0 556 0 444 0.000 62 0.968 0.032 62 0.411 0.218 0.371 0.000 62 0.968 0.032 62 0 966 0.017 0017 59 1 000 0.000 62 0.805 0.161 0.034 59 0 860 0 061 0 061 0.018 0.507 0,485 0.007 67 1 000 0.000 63 0 490 0.127 0 343 0 039 51 0.971 0.029 70 0.918 0 061 0.020 49 0 990 0010 49 0.698 0.250 0.052 48 0.862 0.078 0.043 0.017 58 three buffer systems. The AP6 buffer (Clayton & Tretiak, 1972) was used to resolve esterases {Est, two loci), and phosphoglucomutase (Pgrii). The VVWI (Ward & War- wick, 1980) buffer was also used to resolve Pgm, as well as isocitrate dehydrogenase {Isdh, two loci), and sorbitol dehydrogenase (Sdh). The TC6 buffer (Dillon, 1985) was used to resolve leucine aminopeptidase (Lap) and man- nose-phosphate isomerase (Mpi). Working with L. sax- atilis. Ward et al. (1986; 1991) have confirmed Men- delian inheritance at all these loci except Est and Sdh Data analysis was by Biosys-1 (Release 1.7, Swofford & Selander, 1981) unless otherwise specified. We tested the fits to Hardy- Weinberg expectation for each locus at each population using goodness of fit x" statistics, com- bining rare genotypic classes as necessary. We then per- formed two separate gene diversity analyses using Wrights (1978) F-statistics. In the more conventional analysis, the total deviation from Hardy- Weinberg ex- pectation over all loci (Fi^) was divided into a component Table 2. Wright's (1978) F-statistics, averaged over eight loci, measuring gene diversity attributable to individuals (/), pop- ulations (S ), and coast (C) Values from the hierarchical analysis of coastal variance are set under Fsj, the variance attributable to population structure C^omparison Coefficient F,s FsT Fsc FcB >^sn Fn 0.049 0.033 0.020 0.004 0.024 0.080 within populations (F,^) and a component between the five populations (Fsy). We also performed a hierarchical analysis (Dillon & Manzi, 1992; Dillon & Wethington, 1995), grouping the three Atlantic and two Gulf popu- lations to determine the proportion of gene diversit\' attributable to coast. Mean F-statistics calculated in this way we labeled F^^ (between populations within coasts), F( B (between coasts) and Fsb Note that Fjb is expected to be less than F^-,. since that proportion of the variance between populations between coasts remains unattribut- ed in the hierarchical analysis. For each locus at which n alleles were identified, di- vergence among populations was tested with a 5 x n x" contingency test. To avoid the necessity of combining or eliminating rare alleles for this analysis, we estimated the significance of our values of x" using the Monte Carlo approach of Roff and Bentzen (1989). Unbiased genetic identity and distance between all pairs of populations was calculated using the method of Nei (1978). RESULTS Gene frequencies at eight enzyme-encoding loci from five populations of Littorina irrorata are shown in Table 1. The fits to Hardy- Weinberg expectation within pop- ulations were very close in most cases. Goodness-of-fit values of x' nominally significant at the 0.05 level were obtained only at Pgm in VIMS, and at Isdh in TBAY, well within expectation for type I error. The mean value of F|s over all loci, measuring deviation from Hardy- Weinberg within-sites over the entire studv, was small (F,s=0.049, Table 2). The mean value of F^t = 0.033, measuring deviation from Hardy- Weinberg expectation between sites, was lower than the deviation within-sites. However, the data of Table 1 reflect significant divergence among the five populations at four loci. The value of x' testing homo- geneity in Pgm allele frequencies was 103.6 (p<0.001), with notably high frequencies of Pgm-A in VIMS, Pgm- B in GMBL, and Pgm-C in the three most southern populations. The GMBL population was distinguished by a significantly high frequency of Lap-B (overall X^=46.4, p<0.001), the LUMC population by high Sdh- B (overall x^=21.7, p=0.034), and the CBFL population Page 52 THE NAUTILUS, Vol. 108, No. 3 Table 3. Neis (1978) unbiased genetic identity (above diag- onal) and distance (below diagonal) between all pairs of five L. irrorata populations. Popula- tions VIMS GMBL CBFL TBAY LUMC VIMS — 0.986 0.998 0.990 0.991 GMBL 0.014 — 0.990 0.981 0974 CBFL 0 002 0 011 — 0 996 0.993 TBAY 0010 0019 0 004 — 0,999 LUMC 0 009 0,026 0,007 OOOl — by high Isdhl-B (overall x-=19.6, p=0.047). Differences at the remaining four loci were not significant. The contribution of the coastal level to the hierarchical gene diversity analysis was negligible. Table 2 shows that F^;i5, the deviation from Hardy-Weinberg expectation between coasts, was 0.004. This was lower than the vari- ance between populations within coasts. The values of Neis unbiased similarity and distance among all pairs of sites are shown in Table 3. The four within-coast values of Neis distance ranged from 0.001 to 0.014, only slightly less than the range for the six values between coasts (0.004 to 0.026). DISCUSSION The data of Table 1 constitute substantial evidence of low-level genetic divergence among populations of Lit- torina irrorata separated by distances on the order of hundreds of kilometers. A general relationship between interpopulation divergence, mode of embryonic devel- opment, and dispersal capability has often been noted in marine mollusks (Burton & Feldman, 1981; Scheltema, 1989; Yamada, 1989). Littorina irrorata spawn at high tide, females releasing eggs just beneath the water level (Gallagher & Reid, 1974). Egg capsules are planktonic, but slightly negatively buoyant in calm seawater (Bing- ham, 1972). They hatch into swimming veliger larvae after one to two days of further development, but the time to settlement is unknown. Settlement occurs on Spartina shoots (Boothe, 1969), where juveniles remain hidden in curled blades until they reach about 5 mm (Stiven & Hunter, 1976; Crist & Banta, 1983). Berger (1973) compared population divergence in three North American Littorina species, L. littorea (pelagic larval development), L. ohtusata (juveniles hatch from gelat- inous egg masses), and L. saxatilis (ovoviviparous). The levels of genetic differentiation we report here in L. irrorata are, as might be expected, comparable to those of L. littorea and much less than those of L. ohtusata or L. saxatilis. Such interpopulation genetic divergence as we have identified does not, however, seem to reflect a barrier to dispersal around the Florida peninsula. Table 1 shows no allele uni(|uc to cither coast; even the live rarest alleles {Est2-R. !sdh2-B, Lap-C, Lap-D. and Mpi-R) were found in both Atlantic and Gull populations. Nor did our hi- erarchical gene diversity analysis or our inspection of interpopulation genetic distances suggest any evidence of a barrier to gene flow corresponding to the Florida peninsula. The divergence between Atlantic and Gulf populations of L. irrorata is indistinguishable from dif- ferences among populations sharing the same coast. We offer three (not mutually-exclusive) hypotheses for this unexpected result: no-barrier, past-dispersal, and bal- ancing selection. It is possible that, in spite of the climate shift in south- ern Florida and the disappearance of salt marsh habitat, no barrier currently exists between Atlantic and Gulf populations of L. irrorata. We are unaware of any col- lections of adult snails south of Ft. Pierce, Florida. But it is possible that sparse populations do exist on man- groves or concrete bulkheads along the remainder of the southern Florida coast. Long distance dispersal of larvae or juveniles (perhaps on dead Spartina rack) is also pos- sible. Only a few days might be required to transport larvae on the Gulf Stream from the west coast of Florida into the eastern Atlantic, although onshore currents would still be necessary to carry larvae back into estuaries. It is difficult, however, to imagine how such a passive dis- persal mechanism might be effective for Littorina larvae and not for toadfish, black sea bass, horseshoe crabs, sea anemones, or oyster larvae, all of which do exhibit ge- netic divergence between coasts. A second possible explanation for the absence of di- vergence between Atlantic and Gulf L. littorina popu- lations would invoke higher levels of gene flow in the not-too-distant past. The Suwannee Straits (or Okeefen- okee Trough) most recently connected the two coasts of northern Florida during the Pliocene epoch (Avise, 1992). Fossil L. irrorata found in North Carolina, South Car- olina, and Florida have been dated from the upper Mio- cene and Pliocene, while fossils found in Louisiana and Texas are Pleistocene in age (Bequaert, 1943). Faunal exchange between Atlantic and Gulf may also have oc- curred in the Pleistocene epoch, although it is difficult to predict under what environmental conditions. L ir- rorata could certainly have extended its range southward during glacial periods, but lowering sea levels would have elongated the Florida peninsula. The distance between Atlantic and Gulf shortened as interglacial seas rose, but the snail may have been driven north by the advance of tropical conditions. It would appear, however, that the past-dispersal hypothesis is no better than the no-barrier hypothesis in accounting for the difference betv\een Lit- torina and those (many) s[)ecies that do show intercoastal divergence. The third hypothesis would recall Karl and Avise's (1992) work with oysters. Although allozyme frequencies were homogeneous, Karl ami .\\ ise were able to establish that a barrier has existed between Atlantic and Gulf oyster populations using nilDNA and anonymous nuclear DNA markers. They concluded that balancing selection may be holding Atlantic and Gulf cnster populations undifferentiated in their enzyme poK niorphism in the absence of gene flow, and that caution should be used N. S. Dayan and R T Dillon, Jr . 1995 Page 53 regarding the assumption of neutrality for ailozyme markers. The present data are insufficient to distinguish among these three hypotheses. Further, more detailed surveys of L. irrorata's range in south Florida would be helpful, along with an expansion of isozyme studies if necessary. Some examination of the larval behavior and develop- ment time of L. irrorata would clarifv the likelihood of long distance gene Dow. And further surveys of genetic divergence among Atlantic and Gulf samples of L. ir- rorata using mtDNA or nuclear DN A markers are strong- ly indicated. ACKNOWLEDGMENTS We thank E. W. McMillan, K. Metzner-Roop, and A. R Wethington for assistance in the laboratory, and C. Choi- let and C. Walton for help in the field. Drafts of this manuscript have been read to good effect by C. K. Biern- baum, R. C. Brusca, R. A. McCarthy, and J. W. Smiley Gary Rosenberg provided insights on the distribution of mollusks, R. W. Chapman wrote the program for the Roff-Bentzen analysis, and Dr. & Mrs. M. Dayan pro- vided moral support. This work is based on a thesis sub- mitted in partial fulfillment of the requirements for an M.S. in Marine Biology from the University of Charles- ton, SC. LITERATURE CITED Avise, J. C. 1992. Molecular population structure and the biogeographic history of regional fauna; a case history with lessons for conservation biolog) . Oikos 6362-76 Bequaert, J. C. 1943. 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Journal of Mol- luscan Studies 58: 377-384 THE NAUTILUS 108(3):55-60, 1995 Page 55 Ventistaconcha sima (Lea), an Overlooked Freshwater Mussel (Bivalvia: Unionoidea) from the Cumberland River Basin of Central Tennessee Mark E. Gordon Zoology Section Campus Box 315 University of Colorado Museum Boulder, CO 80309 USA ABSTRACT Unio simus Lea, 1838, generally has been considered to be a junior synonym of Villosa nebulosa (Cxinrad, 1834) Exami- nation of recently collected specimens indicates that it is a valid species of Venustaconcha restricted to the upper Caney Fork River system in central Tennessee. Venuslacuncha sima may represent the Cumberlandian counterpart of V. pleasii (Marsh, 1891) of the southern Ozark Plateaus. Key words: Bivalvia, Unionoidea, Venustaconcha. INTRODUCTION During recent studies of the mussel fauna of the Caney Fork River basin (e.g., Farzaad, 1991; Layzer et ai, 1993), a major tributary system of the Cumberland River, a diminutive purplish-nacred mussel was found, which possessed a posterioventral emargination with an asso- ciated radial sulcus in female shells and relatively heavy hinge dentition. Stansbery (personal communication) considered it to be an undescribed species of Villosa. however, the above morphological characters match di- agnostic criteria in Haas (1969) for Venustaconcha. A review of the multitudinous species descriptions of Isaac Lea and subsequent examination of type specimens in the National Museum of Natural History, Smithsonian Institution, resulted in the identification of this mussel as Venustaconcha sima (Lea, 1838). SYSTEMATICS Tribe Lampsilini Venustaconcha sima (Lea, 1838) (Figures 1-7; Table 1) Margarita (llnio) simus Lea (1836:29) nomen nudum. Unio simus Lea (1838:26, figure 20 on plate V'llI). Margaron (Unio) simus Lea (1852:31). Lampsilis simus Simpson (1900:556). Lampsilis sima Simpson (1914:123). Eurynia (Micromya) nebulosa (in part) Ortmann (1918:577). Micromija nclndosa (in part) Ortmann (1924:102). Lampsilis (Liguniia) nebulosa (in part) Frierson (1927:78). Villosa nebidosa (in part) Burch (1975:173). Description: Shell small, broadly elliptical (males) to somewhat ovate (females), barely inflated (W/HsO.63; for interpretation of shell proportions, see Wu, 1978); solid, thinner posteriorly; anterior rounded; dorsal mar- gin slightly convex, oblique, consequently shell may ap- pear humped or somewhat winged, juncture with an- terior margin sometimes angular; ventral margin slightly convex to virtually straight; posterior bluntly pointed to biangulate; posterio-ventral region of female shells may be swollen with some distention of the extreme posterior of the ventral margin, an emargination and associated radial sulcus may develop posteriorly to the distention; posterior ridge low, vaguely double, somewhat flattened between ridges but may appear rounded; posterior slope slightly concave with a shallow radial furrow, furrow may cause an indentation of the posterio-dorsal margin; ligament low, short; umbo compressed, low, barely ele- vated above dorsal margin; umbonal sculpture ". . ir- regular, somewhat doubly-looped ridges. . ." (Simpson, 1914); periostracum rather smooth and somewhat shiny in younger specimens, may be dull and rough in older shells, annual growth lines well-marked, yellowish to dark brown or black with narrow dark green rays, raying most prevalent on posterior half of shell and may be wavy and clustered closely together. Pseudocardinals thick, heavy, serrated, double in left valve, single in right valve but often with small anterior and posterior denticles opposite adjacent sulci; interden- tum moderately wide, relatively short; lateral teeth short, straight, lamellar, may be slightly serrate, double in left valve, single in right valve; anterior adductor and re- tractor muscle scars confluent, rather small, deeply im- pressed; protractor muscle scar distinct, straight to cres- cent-shaped; posterior muscle scars typically confluent, impressed, adductor large, retractor small and positioned under distal base of lateral tooth; dorsal muscle scars Page 56 THE NAUTILUS, Vol. 108, No. 3 Figures 1-11. Shells of Venustacoiicha. 1-5. Typf sptvuiR-ii!, ul \ cntislaconcha sinm 1. Hnidtype (male, I'SNM 85342) Mai.' paratypes (ANSP .56474). 5. Female paratype (ANSP 56474). 6-7. Venustaconcha sima from C:ollins River, Mt. Olive, Grundy County, Tennessee, collected 15 Julv 1989. 6. Male. 7. Female. 8-9. Venustaconcha ellipsiformis from Osage Fork of Gasconade Hivcr, Dryknoh, Laclede Countv. Missouri, collected 3 October 1983. 8. Male. 9. Female 10-11. Venustaconcha pleasii from James River, east of Springfield, Greene County, Missouri, collected 4 October 1983 10. Male 11. Female Scale bars = 1 cm. M. E. Gordon, 1995 Page 57 Table 1. Shell dimensions of type specimens and representative shells of Venustaconcha sima (L = length, H = height, W = width, M = male, F = female; specimens other than types were collected from the Collins River, Tennessee, Grundy County, Mt. Olive, 15 July, 1989). Specimen Se\ H W H L \V/H Holotype Paratype Paratype Paratype Collins River Collins River Collins River Collins River Collins River Collins River Paratype Collins River Collins River Collins River Collins River Collins River Collins River Collins River Collins River Collins River M 43 9 25.4 15.2 0.57 0 60 M 43.4 26.7 15.6 0 62 0.58 M 42.0 24.3 15.2 0.58 0.63 M 37.8 21.8 14.1 0.58 0.65 M 47.3 27.1 17.5 0.57 0.65 M 42.2 24.7 15.7 0.59 0.64 M 46.5 27.5 15.8 0.59 0.58 M 45.0 25.5 16.6 0.57 0.65 M 42.3 24.0 14.9 0.57 0.62 M 41 I 23,4 13.4 0.57 0.57 M X = 0.58 X = 0 62 F 29.5 17.7 10 3 0.60 0.58 F 42.8 23.4 14.5 0.55 0.64 F 36.4 21.7 14.2 0.60 0.65 F 33.0 19.6 13.4 0.59 0.68 F 31.3 18.4 11.2 0.59 0.61 F 34.7 20.7 12.2 0.60 0.59 F 36.7 21 4 14.8 058 0.69 F 30.5 18.3 11.8 0.60 0.65 F 32.9 19.9 12.3 061 0.62 F 30.6 18.3 12.3 0.60 0,67 F X ± 059 X = 0.64 Overall X = 0.59 X = 0.63 deeply impressed on underside of interdentum, occa- sionally causing a notch in interdentum, extend from just posterior of the umbo to base of pseudocardinals; pallial line impressed, lighter posteriorly; beak cavity moder- ately developed; nacre variable, tends to be purple in hving individuals but may be lighter, pinkish or whitish, and blotched with brow n, color fades rapidly in dead shells. Type locality: "Cumberland River, Tennessee . . . This shell was procured by Professor Troost from the Cum- berland River, but whether near Nashville or not, I am not informed (Lea, 1838). Type specimens: Holotype, National Museum of Nat- ural History, Smithsonian Institution (USNM) 85342 (male). Paratypes, Academy of Natural Sciences of Phil- adelphia (ANSP) 56474 (three males, one female). All type specimens have whitish nacre as noted by Lea (1838)', evidently reflecting the tendency for the purple coloration to fade in dead shells. Only one additional lot of this species was located at the USNM (782358) labelled "Villosa trahalis perpurpurea (Lea, 1861) ', but contain- ' Lea (1862:62) stated that V. sima had a salmon colored nacre. Since examination of available specimens at LISNM and ANSP indicated that Lea only had access to specimens in the type lots (all with white nacre), he may have confused this particular attribute of the similarly-shaped Unio noiatus Lea, 1838 (in- certae sedis) with .sima. ing a mixed collection of Venustaconcha sima and Pleu- robema gibberum (Lea, 1838) from the Collins River, Grundy County, Tennessee. Additional lots are located at the Carnegie Museum, Museum of Comparative Zo- ology, and Ohio State University Museum. Voucher spec- imens from the present study have been deposited in the USNM, University of Michigan Museum of Zoology, Mu- seum of Comparative Zoology, Carnegie Museum, and University of Colorado Museum. Distribution: The distribution of this species is difficult to delineate due to the lack of historical records; however, recent collecting indicates that it is restricted to the Ca- ney Fork River basin above Great Falls. Much of its presumed former habitat is inundated by Great Falls Reservoir. Habitat: Venustaconcha sima appears to be associated with riffle systems in small to medium-sized streams, including headwaters. It is most abundant in sandy sub- strate between cobbles and boulders with slow to mod- erate current, although it also occurs in courser substrate and faster currents (see Farzaad, 1991). Etymology: Venustaconcha- L. venusta [adj.] -I- con- cha [n., feminine]), pretty shell; sima-L. [adj.], snub- nosed. DISCUSSION Following its original description, Venustaconcha sima was known only by its inclusion in synoptic lists of fresh- Page 58 THE NAUTILUS, Vol. 108, No. 3 Figure 12. Distribution of Venustacuncha sima within the Caney Fork River system, Tennessee. water mussels (e.g., Troschel, 1839; Conrad, 1853; Lea, 1870). Call (1885) alluded to an affinity with -Xhuo iris. Lea," 1829, which Simpson (1900, 1914) subsequently followed. Simpson (1914) did note its resemblance to "Lampsilis" nelmlosa (Conrad, 1834) and "L. ' ellipsi- formis (Conrad, 1836). Ortmann (1918) relegated sima as a junior subjective synonym inider "Eunjnia {Micro- mya)' nebidosa. Veimstaconcha sima is a heavier shell than Villosa iris or V. nebidosa. It tends to be relatively shorter and broader than either of the latter and, par- ticularly in the case of females, more closely resembles Venustaconrha ellipsiformis by virtue of its posterior- ventral swelling with associated emargination and radial sulcus, heavy p.seudocardinal teeth, wide interdentum, and fine, wavy, closely-spaced rays (Figs. 8-9). These characteristics were among those employed by Haas (1969) to distinguish Venttstaamcha Frierson, 1927-, from Villosa Frierson, 1927. Within the Cumberland River basin, Venustaconcha sima is similar in appearance to and occasionally may be confused with species in the Villosa iris / nebiilosa comple.x (see above) and V. trabalis (Conrad, 1834) (e.g., (USNM 782358). Compared to Venustaconcha sima, the latter species is a relatively heavier, larger and more elongate shell. Its periostracum tends to be darker, the posterior-ventral emargination of the female is not as acute, pseudocardinal teeth are relatively larger, and the nacre tends not to be blotched. These two species also appear to be distributed allopatrically. Interestingly, the shells of V. trabalis and V. ellipsiformis are more similar to each other than either is to V. sima, and Frierson (1927) included trabalis in his original list of species under Venustaconcha. Although further investigation into generic relationships of advanced lampsilines is re- quired (Hoeh and Frazer, personal communications), classification of trabalis under Venustaconcha may be more representative of actual relationships that an as- sociation with Villosa. The distribution of Venustaconcha sima appears to be restricted to the portion of the Caney Fork River basin draining the Highland Rim upstream from Great Falls, the cataract at the escarpment between the High- land Rim and Nashville Basin (Fig. 12). Call (1885) listed its range as the "Cumberland river, Tennessee, and Swamp Creek, Whitfield County, Georgia." To this, Simpson (1900, 1914) added '. . .Tennessee river sys- tem(s); Othcalooga Creek, northwest Georgia." As noted above, only two lots of V. sima were observed at the USNM. It is unclear upon which specimens Simpson based this range. Call (1885) and Simpson (1900, 1914) possibly may have confused V. sima with species of Vil- losa (e.g., V. iris complex; V. vaniixemii [Lea, 1838]', particularly the umbrans Lea, 1857, morph). Lea (1838) noted the Cumberland River, Tennessee, as the type locality for U. simus, but commented that he did not know the exact collection site. Lea's locality information often was increadibly vague (e.g., Unio grayanus Lea, 1834: type locality - China) or reflected the address of the collector (e.g., Lampsilis reeveiana Lea, 1852: see Gordon & Kraemer, 1984). The latter may be the case with Venustaconcha sima. Since the type specimens were sent by Prof. Troost from Nashville, Lea may have assumed that they had been collected from the Cumberland River. With respect to its known dis- tribution (Fig. 12), the type specimens may have origi- nated from the Collins River or possibly its principle - On several occasions (e.g., Vokes, 1980; Oesch, 1984; Stan.s- bery, xeroxed ephemera), authorship of Venustaconrha has been attributed to Thieie, 1934 or 1935. Venustaconcha was proposed as a replacement name by E'>ier.son (1927) in the "errata el corrigenda" for Vcnusta Frierson, 1927 (nan Boett- ger, 1877, nee Barrande, 1881). ^ Lea (1838) staled that he named this species after Prof Lardner Vanuxem, yet the original spelling ended with the suffix for a locality name rather than the masculine genetive. The spelling of vanuxeniensis was corrected to vanuxemii by Lea (1858). This change is in accordance with Articles 19a (i), 32c (ii), 32d, ;33b (ii) and .Appendix D of the International Code of Zoological Nomenclature (thirtl edition) John.soii (1974) pre- viously noted this correction of the lapstis calami. M. E. Gordon, 1995 Page 59 tributary. Barren Fork, in the vicinity of McMinnville, Warren Count>, Tennessee (the largest town in that area in 1838). Vcniistaconcha sima does not appear to have a coun- terpart in tlie Tennessee River or drainages northward into Kentucky. As previously observed, Simpson (1914) noted similarities between V. sima and V. ellipsijormis. However, in both shell morphology and habitat prefer- ence, V. sima seems to be allied more closely to V. plcasii (Marsh, 1891), a small mussel endemic to the Ozark Plateau drainages of the White River system, Arkansas and Missouri (Figs. 10-11) (personal observations; Gor- don, 1980). This hypothetical association would be con- sistent with previously observed molluscan affinities be- tween the fauna of the Cumberlandian and Interior Highlands regions (e.g., Ortmann, 1917; van der Schalie & van der Schalie, 1950). .\lthough major surveys of the Cumberland River have been conducted (Wilson & Clark, 1914; Neel & Allen, 1964), its mussel fauna has received relatively little at- tention in comparison with that of the Tennessee River system. Starnes and Bogan (1988) listed 85 species from the CAuriberland River drainage, while Gordon and Lav - zer (1989) reported 94 species. Previous faunal compi- lations for the Caney Fork included 27 species (Miller, 1984) and 14 species (Starnes & Bogan , 1988); however, Layzer et al. (1993) have found that the historical fauna was considerably more diverse than the former accounts indicate. It is apparent that the species richness of the Cumberland River mussel fauna is greater than previ- ously considered, Ortmann (1924) concluded that the Cumberland River system lacked an endemic mussel fauna. Starnes and Bogan (1988) similarly stated that "all of the mussel species recorded from the Cumberland River occur in the Tennessee River system despite the inclusion in their synoptic table of two mussels (Alasmidonta atropiirpii- rea [Rafinesque, 1831]^ and Pleurobema gibberum) that are restricted to portions of the Cumberland River drain- age. A distinct endemic mussel fauna did evolve in the upper CAmiberland River system. Despite massive hab- itat destruction within the basin (e.g., reservoir construc- tion, acid coal mine run-off), a fragment of this fauna persists, as represented by A. atropnrpttrea, Quadrula tiiberosa (Lea, 1840: possilby e.xtinct), P. gibberum. and Vcniistaconcha sima. ■* Clarke (1981) reported A. atropurpurea from the Collins River at a site within the Highland Rim province. This speci- men appears to have been a misidentified shell of A. marginata Say, 1818 (Anderson, personal communication) Alasmidonta atropurpurea is restricted to the Cumberland River drainage on the Cumberland Plateau upstream from the hypothesized pre-erosional localilv of Cumberland Falls (Gordon & Lavzer, 1993). ACKNOWLEDGMENTS Partial funding for this study was provided by the Na- tional Ecology Research Center, U.S. Fish and Wildlife Service, Fort Collins, Colorado. I would like to thank Andrew G. Gerberich, Paul R. Greenhall, and Robert Hershler for their assistance and for access to the collec- tions at the National Museum of Natural History. Robert M. Anderson, Indiana Department of Natural Resources; Kenneth S. Frazer, Bar Harbor, Maine; Walter R. Hoeh, Dalhousie l^niversity; and David H. Stansbery , Ohio State University Museum, provided pertinent informa- tion. LITERATURE CITED Burch, J. B. 1975. Freshwater unionacean clams (Mollusca: Pelecypoda) of North America. Malacological Publica- tions, Hamburg, Michigan. 204 pp. Call, R E. 1885. A geographic catalogue of the Unionidae of the Mississippi Valley Bulletin of the Des Moines Acad- emy of Science 1:5-57. Clarke, A H 1981 The tribe Alasmidontini (Unionidae: An- odontinae), part 1: Pegias. Alasmidonta, and Arcidens. Smithsonian Contributions to Zoology 326:1-101. Conrad, T. A. 1853. A synopsis of the family of naiades of North America, with notes, and a table of some of the genera of the family, according to their geographical dis- tribution, and descriptions of genera and subgenera of the familv. Proceedings of the Academy of Natural Sciences of Philadelphia 6:243-269. Farzaad, M 1991. Microhabitat utilization of Venustaconcha sima (Bivalvia; Unionidae) and its host, Cottus carolinae, in the Collins River, Grundy County, Tennessee MS the- sis, Tennessee Technological LIniversity, Cookeville. 94 pp. Frierson, L. S. 1927. A classified and annotated check list of the North American naiades. Baylor University Press, Waco, Texas 111 pp Gordon, M E 1980 Freshwater Mollusca of the Elk River, White River above Beaver Reservoir, and Frog Bayou drainages of the southwestern Ozarks. M.S. thesis. Uni- versity of Arkansas, Fayetteville. 366 pp. Gordon, M. E. and L. R. Kraemer. 1984. Lanipsilis reeveiana and Lampsilis strecheri (Bivalvia: Unionacea): some clar- ifications. Malacological Review 17:99-100. Gordon, M. E. and J. B. Layzer. 1989. Mussels (Bivalvia: LInionoidea) of the CAimberland River: review of life his- tories and ecological relationships. U.S. Fish and Wildlife Service Biological Report 89(15):l-99. Gordon, ME. and J. B. Layzer 1993 Glochidial host of Alasmidonta atropurpurea (Bivalvia: Unionoidea: Union- idae). Transactions of the American Microscopical Society 112:145-1.50. Haas, F. 1969. Das Tierreich, Lieferung 88: Superfamilia Unionacea. Walter de Gruyer & Co., Berlin. 663 pp Johnson, R. I. 1974. Lea's unionid types or recent and fossil taxa of Unionacea and Mutelacea introduced by Isaac Lea including the location of all the extant types. Special Oc- casional Publication of the Department of Mollusks, Mu- seum of Comparative Zoology 2:1-159 Layzer, J B , M. E. Gordon, and R M Anderson. 1993. Mus- sels: the forgotten fauna of regulated rivers Regulated Rivers: Research and Management 8:63-71. Page 60 THE NAUTILUS, Vol. 108, No 3 Lea. I. 1836 A s\ nopsis of ttic family of naiades. Carey, Lea, and Blaiuhard, Fhiladelpliia 59 pp. Lea, I. 1838. Descriptions of new fresh- water and land shells Transactions of the American Philosophical Society 6:1- 154. Lea, I. 1852. A synopsis of the family of naiades. Blanchard and Lea, Philadelphia. 88 pp. Lea, I. 1858. New Unionidae of the LInited States Journal of the Academy of Natural Sciences of Philadelphia 451- 95. Lea, I. 1862. New Unionidae of the United States. Journal of the Academy of Natural Sciences of Philadelphia 5:53- 109. Lea, 1. 1870. A synopsis of the famiU of naiades Henry C Lea, Philadelphia. 184 pp. Miller, A. C. 1984. A survey for mussels on the lower Caney Fork River, Tennessee, 18-20 October, 1983 U.S. Army Waterways E.xperiment Station, Vicksburg, Mississippi 18 pp. Neel, J. K. and W. R. Allen. 1964. The mussel fauna of the upper Cumberland River before its impoundment Malacologia 1:427-459. Oesch, R. D. 1984. Missouri Naiades: a guide to the mussels of Missouri. Missouri Department of Conservation, Jeffer- son City. 270 pp. Ortmann, A. E. 1917. A new type of the nayad-genus Fus- conaia. Group of F. barncsiana Lea. The Nautilus 31:58- 64. Ortmann, A. E. 1918. The nayades (freshwater mussels) of the upper Tennessee drainage with notes on synonymy and distribution Proceedings ol the American Philosoph- ical Society 57:521-626. Ortmann, A. E. 1924. The naiad-fauna of Duck River in Tennessee. American Midland Naturalist 9:18-62. Simpson, C. T. 1900. Synopsis of the naiades, or pearly fresh- water mussels. Proceedings of the U.S. National Museum 22:501 -1044. Simpson, C. T. 1914. A descriptive catalogue ot the naiades or pearly fresh-water mussels. Brvant Walker, Detroit, 1540 pp. Starnes, L. B. and A. E. Bogan. 1988. The mussels (Mollusca: Bivalvia: Unionidae) of Tennessee. American Malacolog- ical Bulletin 6:19-37. Troschel, F H. 1839. Die Familie der Najaden. Archiv fiir Naturgeschichte 5:235 van der Schalie, H and A van der Schalie 1950 The mussels of the Mississippi River American Midland Naturalist 44: 448-466. Yokes, H E 1980. Genera of the Bivalvia: a systematic and bibliographic catalogue (revised and updated). Paleonto- logical Research Institute, Ithaca, New York, 307 pp. Wilson, C. B., and H. W. Clark. 1914. The mussel fauna of the Cumberland River and its tributaries. Report of the U.S. Commissioner of Fisheries for 1912, Special Paper: 1-63. Wu, S.-K. 1978. The Bivalvia of Colorado, part 1. The fingernail and pill clams (family Sphaeriidae). University of Colorado Museum Natural History Survey of Colorado 2:1-39. THE NAUTILUS 108(;3):61-66, 1995 Page 61 Richard S. Houbrick (1937-1993): Biographical Sketch and Malacological Contributions M.C. Harasewyeh Alan R. Kabal Department of Invertebrate Zoology National Museum of Natural History Smithsonian Institution Wasfiington, DC. 20560 USA I. BIOGRAPHY The malacological community lost one of its most emi- nent and respected members and prolific authors with the death of Richard S. Houbrick on August 26, 1993 at the age of 56, after a long and valiant struggle with leukemia and hepatitis Richard Steven Houbrick was born on March 16, 1937 in Trenton, New Jersey to Stephen J. and Barbara A. Houbrick. The family moved to Hollywood, Florida when Joe was 12. As a young student, Joe was an avid field biologist, shell collector, and occasional shell show ex- hibitor. After completing Broward High School, he en- tered St. John's Seminary in Little Rock, Arkansas, then transferred to St. Bernard College, Cullman, Alabama, where he graduated with a Bachelor of Arts degree in 1959. He then entered St. Leo Theological School, St. Leo, Florida, and was ordained to the priesthood in March, 1963. As a member of St. Leo Abbey (Benedictine Order) he took the name Joseph, and has since been known as "Joe' to his many friends and colleagues. He was an Instructor in general biology, zoology, invertebrate zo- ology, embryology and botany at St. Leo College. He began attending summer sessions of the University of Miami and the University of Florida, taking courses in Biology and Chemistry. In 1964, Joe entered the Grad- uate School of the L'niversity of Miami. He earned a M.S. degree from the Rosenstiel School of Marine and At- mospheric Science, University of Miami in 1967, having completed his thesis on "A survey of the littoral marine mollusks of eastern Costa Rica. In the summer of 1968, he attended the Biology of Mollusks Training Program, in Hawaii, where he studied moUuscan anatomy with Vera Fretter and others. This course had a profound influence on his future research, as shown by his extensive use of anatomy and repro- ductive biology in his systematic publications. Joe enrolled in the newly established LIniversity of South Florida in 1968, and received the first Ph.D. to be awarded by this University in 1971, for his dissertation on "Taxonomy, anatomy, and life history studies on the genus Cerithium (Gastropoda: Prosobranchia) in the western Atlantic. His thesis advisor was Dr. Joseph L. Simon, a marine ecologist. Upon completing his doctor- ate, Joe joined the staff of the Smithsonian Oceanograph- Richard S Houbrick at the Smithsonian Marine Station at Link Port, Florida, February, 1983. ic Sorting Center, and was subsequently appointed Su- pervisor of the Benthic Section, and later Acting Director. In 1977, Joe was appointed Associate Curator in the Division of Mollusks at the National Museum of Natural History, where he became Curator in 1981, a position in which he remained active until shortly before his death. Noted primarily for his expertise in the systematics, comparative anatomy, fimctional morphology and re- productive biology of the Cerithioidea, he contributed significantly to the body of descriptive and synthetic literature on this group. Since his work emphasized the Page 62 THE NAUTILUS, Vol. 108, No. 3 study of living animals and llieir life-histories, he traveled wideK , conducting field work in many parts of the world, including the tropical western Atlantic, the tropical Pa- cific and .\ustralia. He was a frequent invited participant in s\niposia and workshops. Since joining the Division of Mollusks, Joe spent an extremely productive portion of each year working with live animals at the Smithsonian Marine Station at Link Port, Florida. The many colleagues who had an oppor- tunity to work with Joe at the Marine Station soon dis- covered his special interest in and enormous knowledge of the flora and fauna of Florida that he had accumulated since childhood. Joe was an active supporter of the Ma- rine Station, serving on its Task Force and Advisory Group. All who knew Joe recognized that he was above all an educator. He was an Adjunct Professor at George Wash- ington University, and served on thesis committees of graduate students at this and other universities. He also served as Advisor to several Smithsonian Post-doctoral Fellows, who have since risen to prominent positions in the field of systematic malacology. Whether student, post- doc or professional colleague, his own or someone else's, Joe would always take the time to show an interest in their work, to learn from them as well as to make helpful comments and offer his own insights. Joe was active in numerous professional organizations, among them the American Malacological Union (of which he was president in 1980-81), the American Society of Zoologists, the California Malacozoological Society, the Malacological Society of London, Unitas Malacologia, the Biological Society of Washington, and the Smithsonian Senate of Scientists. Joe will be remembered as a quiet scholar whose re- search and publications stand as a lasting monument to his scientific dedication and insight. As a person, he was known for his sense of humor, for his enjoyment of the fine arts along with music, and he was invariably most congenial to all. He is survived by his parents, and by his brother James and his family. ACKNOWLEDGMENTS We thank James Houbrick, Robert Hershler, and CHyde Roper for their helpful comments on this manuscript. II MALACOLOGICAL TAXA Of R.S. Houbrick s fifty-six publications from 1967 to 1993 (not counting meeting abstracts or notes), twelve included descriptions of twenty-two new taxa. These comprise two family-level taxa, three genus-level taxa (two in the C>erilhiidae and one in the Plana.\idae) and seventeen species-level taxa (twelve in the C^erithiidae and the remainder in other prosobranch gastropod fam- ilies). Two of the species-level taxa are replacement names for junior homonyms. Abbreviations AMS Australian Mu,seuni, Sydney ANSP Academy of Natural Sciences of Philadelphia LACM Los Angeles County Museum of Natural His- tory MNHNP Museum National d'Histoire Naturelle, Paris NM Natal Museum, Pietermaritzburg NMFS National Marine Fisheries Service [USA] USBF U.S. Bureau of Fisheries USNM U.S. National Museum [= National Museum of Natural History, Smithsonian Institution] WAM Western Australian Museum, Perth Family-level taxa CERITHIDEIDAE Houbrick 1988, Malacological Re- view, Supplement 4:88 jf. (diagnosis on page 118). For (by implication) Cerithidea Swainson 1840. Subsequent- ly svnonymized with the Potamididae H. & A. Adams 1854 by Houbrick (1991, Malacologia, 33(l-2):333). Cer- ithioidea. PLESIOTROCHIDAE Houbrick 1990, Proceedings of the Third International Marine Biological Workshop: The Marine Flora and Fauna of Albany, Western Australia, pages 237 ff. (diagnosis on page 248). For Plesiotrochus Fischer 1878. Originally placed in the Cerithioidea. Sub- sequently transferred to the Campaniloidea by Healy (1993). Genus-level ta.xa Ittibittium Houbrick 1993, Malacologia, 35(2):283. Type species Bittium parcuni Gould 1861; monotypy. Ceri- thiidae. Longicerithium Houbrick 1978, Monographs of Marine Mollusca, 1:85. Type species Cerithium longicaudata A. Adams and Reeve 1850; original designation. As a sub- genus of Rhinoclavis Swainson 1840. Cerithiidae. Simulathena Houbrick 1992, The Veliger, 35(l):64-65. Type species Simulathena papuenais Houbrick 1992; monotypy. Planaxidae. Species-level taxa abditum, Cerithium Houbrick 1992, Smithsonian Con- tributions to Zoology, 510:18-20, figures 6-8. Holotvpe USNM 286404; five paratypes USNM 860482. USBF'a/- batross Station 5569, 554 meters, off Simaluc Island, Tawitawi, Philippines. Cerithiidae. admirabilis, Clypeontorus Houbrick 1985, Smithsonian C:ontributions to Zoology, 403:99-104, figures 47-49. Ho- lotype WAM 3346-83; seven paratypes USNM 784652; thirty-eight paratypes ANSP 233431; three paratvpes AMS 139779. False Cape Creek (near mouth), 100 miles (161 kilometers) S of Broome, Western Australia, .Aus- tralia. Cerithiidae. africanum, Cerithium Houbrick 1992, Smithsonian Contributions to Zoology, 510:20-23, figures 9-11. Ho- lotype NM HI 861; five paratypes USNM 629034. South- western Conducia Bay, NW of Choca, Mozambique. Cerithiidae. M. G. Harasewych and A. R. Kabat, 1995 Page 63 argentea, Strombina (Cotonopsis) Houbrick 1983, Pro- ceedings of the Biological Society of Washington, 96(3): 349-352, Bgures 1-2. Holotvpe USNM 810345; nine para- types USNM 821851. NMFS R/V Oregon. Station 5432, 384-430 meters, 20°48'N, 70°46'W, near Silver Bank, 60 miles NE of Luperon, Dominican Republic. Columbel- lidae. diadema, Rhinoclavis (Rhinoclavis) Houbrick 1978, Monographs of Marine Molkisca, 1:64-66, figures 33-35. Replacement name for Cerithitim nitidum Honibron and Jacquinot 1854 non MacAndrew and Forbes 1847 et al.. Lectotype and one paralectotype (of CerUhiiim nitidum), MNHNP (not numbered), selected by Houb- rick (1978: 66). Mangareva, Gambier Islands. Cerithi- idae. exiinium, Ataxocerithium Houbrick 1987, The Nautilus, 101(4): 157-159. figures 1-16. Holotype AMS C142394; five paratvpes AMS C153005; two paratypes L'SNM 862328. 421 meters, 34°21-16'S, 151°24-28'E, SE of Bot- any Bay, New South Wales, Australia. Cerithiopsidae. garciai. Cerithioclava Houbrick 1986, Proceedings of the Biological Society of Washington, 99(2):257-260, figure 1. Holotype USNM 849023; two paratypes USNM 849024. 24 meters, off Great Corn Island, Nicaragua [Caribbean Sea]. Cerithiidae. gloriostun. Cerithium Houbrick 1992, Smithsonian Con- tributions to Zoology, 510:94-97, figures 61-63. Holo- type, MNHNP (not numbered); 200 paratypes MNHNP (not numbered); seventy-nine paratypes USNM 862327. Benthedi Station DR-06, 500-600 meters, 11°28'S, 47°12'E, lies des Glorieuses, Indian Ocean. Orithiidae inflatiis, Acamptochetus Houbrick 1984, Proceedings of the Biological Society of Washington, 97(2):421-423, fig- ure 1. Holotype USNM 229183; one paratype USNM 820185. USBF Albatross Station 5419, 320 meters, 9°58'30"N, 123°46'E, off Lubigon, Bohol, Philippines. Buccinidae. nympha, Clypeomorus Houbrick 1985, Smithsonian Contributions to Zoology, 403:113-120, figures 56-59. Replacement name for Cerithium variegatum Quoy and Gaimard 1834 non Fischer 1807 et al.. Lectotype and one paralectotype (of Cerithium variegatum), MNHNP [not numbered]; selected by Houbrick (1985: 113). Ton- gatapu [Tonga Islands]. Cerithiidae. pacificum Cerithium Houbrick 1992, Smithsonian Con- tributions to Zoology, 510:143-146, figures 103-104. Ho- lotype USNM 584662; four paratypes USNM 862491. 55-61 meters, Bikini lagoon, Bikini Atoll, Marshall Is- lands. Cerithiidae. papuensis, Simulathena Houbrick 1992, The Veliger, 35(l):65-67, figures 1-13. Holotype AMS C166326; two paratypes USNM 859456. 8°50'S, 146°32'E, Kairuku, Yule Island, Central District, Papua New Guinea. Planaxidae. persica, Clypeomorus bifasciata (Sowerby 1855) netv subspecies Houbrick 1985, Smithsonian Contributions to Zoology, 403:41-43, figures 17-18. Holotype, ANSP 263194; eight paratypes USNM 838028. Ras fanura. Per- sian Gulf, Saudi Arabia. Cerithiidae. purpurastoma, Clypeomorus Houbrick 1985, Smithson- ian Contributions to Zoology, 403:83-89, figures 38-40. Holotype L'SNM 774749; sixteen paratypes USNM 784650 Napot Point, Morong, Bataan, Luzon, Philip- pines. Cerithiidae. rehderi, Cerithium Houbrick 1992, Smithsonian Con- tributions to Zoology, 510:156-158, figures 112-113. Ho- lotype USNM 859954; five paratypes USNM 798574. 56- 72 meters, Haava Straits, between Tahuata and Hiva Oa, Marquesas. Cerithiidae. reidi. Cerithidea Houbrick 1986, The Veliger, 28(3):280 ff., figures 1-16. Holotype WAM 3380-84; one paratype AMS C144144; ten paratypes USNM 828823. Willies Creek, N of Broome, Western Australia, Australia. Po- tamididae. scobiniforme, Cerithium Houbrick 1992, Smithsonian Contributions to Zoology, 510:177-179, figures 129-130. Holotype USNM 859930; two paratypes USNM 859931; six paratypes LACM 76-702. 1.5 miles S of Estango Is- land, Port Dimalosan, Cagayan Province, Luzon, Phil- ippines. Cerithiidae. III. PUBLICATIONS R.S. Houbrick authored or co-authored a total of fifty- six research papers and notes, along with eighteen ab- stracts, two "President's Messages " (AMLI) and two grad- uate theses over a twenty-seven year period (1967-1993). Of the research papers, thirty-eight (68%) treated mem- bers of the Cerithioidea (twenty-five specifically on the Cerithiidae), thirteen (23%) were on other prosobranch gastropod families, and five (9%) were on general topics. As for publication venue, the following statistics were obtained. Thirty-one (55%) of his research papers were published in malacological serials, twenty (36%) were published in non-malacological serials (primarily the Proceedings of the Biological Society of Washington and the Smithsonian Contributions to Zoology), and the re- maining five (9%) were published as book chapters. Note: his papers for 1967-1971 were authored as "JR. Houbrick"; those for 1972-1974b as either "R.S. Houb- rick or "R.S. (JR.) Houbrick", and the remainder as ■R.S. Houbrick". Abbreviations AMU American Malacological Union WSM Western Society of Malacologists 1967 [24 April]. Notes on Cyctostremiscus .whramnni The Nautilus 80(4):131-133. 1968a [1 Jan ] New record of Conu4c/?rae2i,s in Costa Rica The Veliger 10(3):292. 1968b [1 July]. A survey of the littoral marine mollusks of the Caribbean coast of Costa Rica. The Veliger 11(1):4- 23. Page 64 THE NAUTILUS, Vol. 108, No. 3 1969 [.^ug.] Houbriok, J R and Fretter, V Some a.spects of the functional anatomy and biology of Cymatium and Bursa. Proceedings of llie Malacological Society of London 38(5):415-429. 1970 [June]. Houbrick, J.R. and Fretter, V. Some aspects of the functional anatomy and biology of Cymatium and Bursa. In: (Kay, E.A., ed.). The Biology of Molluscs. A collection of abstracts from the National Science Foun- dation Graduate Research Training Program June - Sep- tember 1968, pages 12-13. University of Hawaii, Hawaii Institute of Marine Biology, Technical Report 18, 30 pages. [Abstract] 1971a "1970 [18 Feb ] Reproduction and development in Florida Cerithium AMU, Key West, Florida, 16-20 July 1970. Annual Report of the American Malacological L'nion for 1970, page 74. [Abstract] 1971b [26 Oct.]. Some aspects of the anatomy, reproduction, and early development of Cerithium nodulosum (Bru- guiere) (Gastropoda, Prosobranchia) Pacific Science 25(4): 560-565. 1972 [Nov.]. The Smithsonian Institution - Great Lakes role. Great Lakes Basin Commission, Communicator 3(5):3-4. 1973 [10 Oct! CcnV/iium Bruguiere, [1789], (Gastropoda); Proposed preservation by designation of a type-species under the plenary powers. Bulletin of Zoological Nomen- clature 30(2)404-107. 1974a [29 Jan ]. Growth studies on the genus Cerithium (Gastropoda: Prosobranchia) with notes on ecology and microhabitats. The Nautilus 88(1); 14-27. 1974b "1973" [7 May]. Studies on the reproductive biology of the genus Cerithium (Gastropoda: Prosobranchia) in the Western Atlantic. Bulletin of Marine Science 23(4); 875-904. 1974c [29 July]. The genus Cerithium in the Western At- lantic (Cerithiidae; Prosobranchia) Johnsonia 5(50)33-84. 1975 [29 Oct.]. Clavocerithium (Indocerithium) taeniatum, a little-known and unusual cerithiid from New Guinea The Nautilus 89(4);99-105. 1976 "1975" [30 Jan.]. Preliminary revision of supraspecific taxa in the Cerithiinae Fleming, 1882 (Cerithiidae: Pros- obranchia) Bulletin of the American Malacological Union, Inc. for 1975, pages 14-18, 18a. 1977 [1 Oct.]. Reevaluation and new description of the genus Bittium (Cerithiidae). The Veliger 20(2); 101-106. 1978a [31 Jan ] Redescription of Bittium proteum (Jous- seaume, 1930) with comments on its generic placement. The Nautilus 92( I ):9-ll, 1978b [April]. Form, function, and adaptive radiation in snails of the family Cerithiidae. AMU, Naples, Florida, 10-15 July 1977. Bulletin of the American Malacological Union, Inc. for 1977, page 84. [Abstract] 1978c [6 Dec.]. Reassignment of Batillaria sordida (Gmelin) from the Cerithiidae to the Potamididae (Gastropoda: Prosobranchia). Proceedings of the Biological Society of Washington 91(3);642-649 1978d [15 Dec ] The family Cerithiidae m the Indo-Pacific Part 1; The genera Rhinoclavis. Pseudovertagus and Chiv- ocerithium. Monographs of Marine Mollusca 1:1-130 1979 [5 June]. Classification and systematic relationships of the Abyssochrysidae, a relict family of bathyal snails (Pro.s- obranchia; (Jastropoda) Smith.sonian Ointributions to Zo- ology 290:1-21. 1980a [24 March]. Reproductive biology of Modttlu.s mod- ulus Linnaeus. 1758 (Prosobranchia: Ceritliiacea). AML' and WSM, (;orpus Christi, Texas, 5-11 August 1979. Bul- letin of the American Malacological L'nion, Inc for 1979, page 71 [title only]; The Western Society of Malacologists Annual Report 12:10. [Abstract] 1980b [30 June] Form, function and adaptive radiation in the Cerithiidae. Symposium on the biology and evolution of Mollusca, Australian Museum, Sydney, 21-25 May 1979. Journal of the Malacological Society of Australia 4(4):230- 231. [Abstract] 1980c [23 July] Review of the deep-sea genus Argyropeza (Gastropoda: Prosobranchia: Cerithiidae). Smithsonian Contributions to Zoology 321:1-30. 1980d [22 Aug ]. Observations on the anatomy and life his- tory of Moduhis modulus (Prosobranchia: Modulidae). Malacologia 20(1):1 17-142. 1980e [Aug.]. Anatomy and biology of Campanile symhul- icum Iredale with reference to adaptive radiation of the Cerithiacea. Unitas Malacologica, Perpignan, France, 31 Aug. - 7 Sept. 1980. In: (Gaillard, J. M., ed). [AbstractsJ Septieme Congres International de Malacologie. Haliotis 10(2): 168. [Abstract] 1980f [6 Nov,]. Reapprai.sal of the gastropod genus Varico- peza Griindel (Cerithiidae: Prosobranchia). Proceedings of the Biological Society of Washington 93(3):525-535. 1980g Systematics, anatomy and biology of Campanile sym- bolicum. a relict cerithiacean snail. AMU, Louisville, Ken- tucky, 19-25 July 1980 Bulletin of the American Mala- cological Union for 1980, page 72. [Abstract] 1980h Presidents Message. The .American Malacological Union, Inc., Newsletter, lI(2-A):l-2. 1981a [29 Jan ]. .Anatomy and systematics of Gourmya gour- mi/i (Prosobranchia: Cerithiidae), a Tethyan relict from the Southwest Pacific. The Nautilus 95(1):2-1I. 1981b [20 July] Anatomy of Dias/oma nip/anioirfes (Reeve, 1849) with remarks on the systematic position of the family Diastomatidae (Prosobranchia: Gastropoda). Proceedings of the Biological Society of Washington 94(2):598-621. 1981c [16 Oct ]. Systematic position of the genus C.lyptozaria Iredale (Prosobranchia: Gastropoda) Proceedings of the Biological Society of Washington 94(3):838-847. 1981d [8 Dec ]. .Anatomy, biology and systematics of Cam- panile symbolicum with reference to adaptive radiation of the Cerithiacea (Gastropoda: Prosobranchia). Malaco- logia 21(l-2):263-289. 1981e What is Diastema? Systematic position of the Dias- tomatidae AMU. Fort Lauderdale, Florida, 19-25 July 1981 Bulletin of the American Malacological L'nion for 198L page 31 [Abstract] 198 If President's Message The American Malacological Union, Inc., Newsletter 12(1):1. 1983 [31 Oct.]. A new Strombina species (Gastropoda: Pros- obranchia) from the tropical Western Atlantic. Proceed- ings of the Biological Societ\ of Washington. 96(3):349- 354. 1984a [Feb.] Revision of higher taxa in genus Cerithidea (Mesogastropoda: Potamididae) based on comparative morphology and biological data American Malacological Bulletin 2:1-20, 95. 1984b [29 March]. Functional morpholog) and ontogeny of Mollusca as related to higher category systematics; Intro- duction. Malacologia 25(I):1. 1984c [April], (Joing collecting? Look for a live Royella si- mm Hawaiian Shell News 32(4); 12. 1984d [31 Ma\ ] Ke\ision of higher taxa in genus CVn7/!i(ipa based on comparative morphology and biological data. AMLI and WSM, I'niv. of Washington, Seattle, 7-13 Aug. M. G. Harasewvch and A. R Kabat, 1995 Page 65 1983 The Western Society of \4alacologists. Annual Re- port 16:23. [Abstract] l*)84-e [6 Jub ] A new "Mctula" species from the Indo-West Pacific (Prosobranchia; Bucciniclae). Proceedings of the Biological Society of Washington 97(2)420-424. 1984f [19 July]. The Giant Creeper, Campanile symbolicum Iredale, an Australian relict marine snail. In: (Eidredge, N. and Stanley, S.M eds. ), Living Fossils, pages 232-235 New York, Springer Verlag, xii + 291 pages. I'JBtg [19 Jub] Diastoma mclaniuidcs (Reeve), a relict snail Iriiui South Australia. In: (Eidredge, N. and Stanley, S.M. eds.), Living Fossils, pages 236-239. New York, Springer Verlag, xii + 291 pages. 1081-h [19 July] The relict cerithiid prosobraiich, Gourmya gutirniyi (Crosse). In: (Eidredge. N. and Stanley, S.M. eds.). Living Fossils, pages 240-242. New York, Springer Verlag, xii + 291 pages. IQSSa "1984" [Feb.]. Comparative reproductive biology of P/aiiflxis species. AMU, Norfolk, Virginia, 22-27 July 1984. American Malacological Bulletin 3(1)96. [Abstract] 1985b [3 April] Genus CUjpcomorus Jousseaume (Cerithi- idae: Prosobranchia). Smithsonian Contributions to Zool- ogy 403:1-131 1986a [2 Jan.]. Cerithidea reidi. spec, nov., from Western Australia. The Veliger 28(3):280-286. 1986b [Feb.] C/i/pfomoru.s, a genus of littorinid-like cerith- ids. AMU, Univ. of Rhode Island, Kingston, 28 July - 2 Aug. 1985. American Malacological Bulletin 4( 1 ) 109, [Ab- stract] 1986c [1 April]. The systematic position of Royella sinon (Bavle) (Prosobranchia: Cerithiidae). The Veliger 28(4): 429-435. 1986d [4 June]. Discovery of a new living Cerithioclava species in the Caribbean (Mollusca: Prosobranchia: Ceri- thiidae). Proceedings of the Biological Society of Wash- ington 99(2):257-260. 1986e [4 June] Transfer of Quadrasia from the Planaxidae to the Buccinidae (Mollusca: Gastropoda: Prosobranchia). Proceedings of the Biological Society of Washington 99(2): 359-362. 1986f [June] Observations on the anatomy and life histories of cerithiacean prosobranchs. In: (Rice, M., ed). Abstracts, Symposium on Current Research at the Smithsonian Ma- rine Station at Link Port, June 24 - 25, 1986. Fort Pierce (Florida), Smithsonian Marine Station [unpaginated]. [Ab- stract] 1986g [Aug ] Functional anatomy and systematicsof Litiopa and Alaba (Prosobranchia: Orithiacea). AML' and WSM, Monterey, California, 1-6 July 1986, American Malaco- logical Bulletin 4(2):235. [Abstract] 1986h [Aug ]. Phylogenetic relationships in the superfamily Cerithiacea. Unitas Malacologica, Edinburgh, Scotland, 31 Aug. - 6 Sept 1986 In: (Heppell, D , ed). Abstracts, Unitas Malacologia. Ninth International Malacological Congress, page 38. Edinburgh, National Museums of Scotland, 104 pages. [Abstract] 1986i Prosobranch Mollusca of Carrie Bow Cay In: (Riitzler, K , ed), Caribbean Coral Reef Ecosystems, Progress Re- port 1986, page 42. Washington, DC, National Museum of Natural History, Smithsonian Institution, 49 pages. [Ab- stract] 1987a [30 Jan.]. Anatomy of Alaba and Litiopa (Prosobran- chia: Litiopidae): systematic implications. The Nautilus 101(1):9-18. 1987b [18 March]. Anatomy, reproductive biology, and phy- logeny of the Planaxidae (Cerithiacea: Prosobranchia). Smithsonian C^jntributions to Zoology 445:1-57. 1987c [28 .'Vpril] Transfer of Cerithiopsifi crystallina Dall to the genus Varicupeza Griindel, family Cerithiidae (Prosobranchia: Gastropoda). The Nautilus 101(2):80-85. 1987d [31 July]. Houbrick, R.S., Robertson, R , and Abbott, R T. Anatomy and systematic position of Fasligiella car- inata Reeve (Cerithiidae: Prosobranchia) The Nautilus 101(3):10I-110 1987e [6 Nov.], Description of a new, giant Ataxoccrithium species from Australia with remarks on the systematic placement of the genus (Prosobranchia: Cerithiopsidae). The Nautilus 101(4):155-161. 1988a [15 Oct.]. Houbrick, R S., Stiirmer, W. and Yochelson, EL. Rare Mollusca from the Lower Devonian Hunsriick Slate of southern Germany. Lethaia 21(4):395-402. 1988b [20 Dec]. Cerithioidean phylogeny In: (Ponder, WF., et ai, eds ), Prosobranch Phylogeny. Malacological Re- view, Supplement 488-128. 1989a [April] Campanile revisited: implications for ceri- thioidean phylogeny. American Malacological Bulletin 7(1): 1-6. 1989b [Aug.]. Functional morphology and systematics of mangrove snails Terebralia and Telescopium (Potamidi- dae; Prosobranchia). Unitas Malacologica, Universitat Tii- bingen. West Germany, 27 Aug. - 3 Sept. 1989 In: (Meier- Brook, C, ed). Abstracts, Unitas Malacologica, Tenth In- ternational Malacological Congress, page 106 Tiibingen, Tropenmedizinischen Institijt der Universitat Tubingen, li + 287 pages. [Abstract] 1989c [29 Sept ]. Comments on the proposed conservation of Iphinoe Bate, 1856 (Crustacea, Cumacea). Bulletin of Zoological Nomenclature 46(3):191. 1 990a [ 1 1 June]. Anatomy, reproductive biology and system- atic position of Fossarus ambiguus (Linne) (Fossaridae: Prosobranchia). AML' and WSM, Univ. of Southern Cal- ifornia, Los Angeles, 25-30 June 1989 Western Society of Malacologists Annual Report 2222. [Abstract] 1990b [6 Sept ] Review of the genus Co/ina H and A. Ad- ams, 1854 (Cerithiidae: Prosobranchia). The Nautilus 104(2):35-52. 1990c [Oct.] Anatomy, reproductive biolog) and systematic position of Fossarus ambiguus (Linne) (Fossarinae: Plan- axidae; Prosobranchia) In: (Frias Martins, A.M., ed). The Marine Fauna and Flora of the Azores Proceedings of the First International Workshop of Malacology, Sao Miguel, Azores, 11-24 July 1988, pages 59-73. A^oreana, Suple- niento, 173 pages. 1990d [31 Dec.]. Aspects of the anatomy of Plesiotrochus (Plesiotrochidae, fam. n.) and its systematic position in Cerithioidea (Prosobranchia, Caenogastropoda). In: (Wells, F.E., Walker, D.I., Kirkman, H & Lethbridge, R., eds). Proceedings of the Third International Marine Biological Workshop: The Marine Flora and Fauna of Albany, West- ern Australia, pages 237-249. Perth, Western Australian Museum, volume 1 (of 2), x -t- 437 pages. 1991a [15 July] Functional inference from gastropod shell morphology - some caveats Lethaia 24(3):265-270. 1991b [6 Sept.], Systematic review and functional morphol- ogy of the mangrove snails Terebralia and Telescopium (Potamididae; Prosobranchia). Malacologia 33(l-2):289- 338. 1991c [16 Dec.]. Anatomy and systematic placement of Fai/- nus Montfort 1810 (Prosobranchia: Melanopsinae). Mal- acological Review 24(l-2):35-54. Page 66 THE NAUTILUS, Vol. 108, No. 3 1992a [2 Jan ]. Sinitilathcna pajmcnsis. a new planaxid genus antl species from the Imlo-West Pacific. The Vehger 35(1): 64-69. 1992b [8 June]. Anatomy and prehminary generic review of the Bittium group. AMLI and WSM, Univ. of California, Berkeley, 30 June - 5 July 1991. Western Society of Ma- iacologists, Aniuial Report 24:27. [Abstract] 1992c [29 Sept.]. Monograph of the genus Cerithiuiu Bru- guiere in the Indo-Pacific (derithiidae: Prosobranchia) Smithsonian C'ontributions to Zoology 510, iv + 211 pages. 1993a [24 March]. Two confusing Indo-Pacific cerithiids. The Nautilus 107(l):14-23. 1993b [2 Dec.]. Phylogenetic relationships and generic re- view of the Bittiinae (Prosobranchia: Cerithioidea). Malacologia 35(2):261-313. NOTE: Dr. Winston F. Ponder (Australian Museum) is editing and revising several uncompleted manuscripts left by R.S. Houbrick for eventual publication. Graduate Theses 1967 [May]. A survey of the littoral marine mollusks of East- ern Costa Rica. Master of Science, University of Miami, Coral Gables, Florida, iv + 90 pages, 5 plates. See Houb- rick, 1968b. 1971 [May]. Taxonomy, anatomy and life history studies on the genus Cerithium (Gastropoda: Prosobranchia) in the Western Atlantic. Doctor of Philosophy, University of South Florida, Tampa, Florida xii -I- 182 pages. See Houbrick, 1974a, 1974b and 1974c. OTHER LITERATURE CITED Healy, J M. 1993. Transfer of the gastropod family Plesi- otrochidae to the C^ampaniloidea based on sperm ultra- structural evidence Journal ol Moiluscan Studies 59(2): 135-146. THE NAUTILUS 108(3):67-79, 1995 Page 67 New Data on the Distribution and Morphology of some Western Indian Ocean Sepiid Cuttlefishes (Cephalopoda: Sepiida) Yu. A. Filippova D. N. Khroniov Russian Research Institute of Marine Fisheries & Oceanography (VNIRO) 1071-10 Moscow RUSSIA K. N. Nesis I. V. Nikilina P F. Shirsliov Institute of Oceanology Russian Academy of Sciences 117218 Moscow RUSSIA ABSTRACT New data are presented on the distribution and/or morphology of 21 species of the family Sepiidae, based on the collections of various expeditions during 1956-1989 in the Western Indian Ocean from the Northern Arabian Sea to Southern Mozam- bique, Agulhas Bank and Cargados-Carajos Islands Previously neglected or inadequately described characters are provided for Sepia joulnni, S. confusa. S. typica and others. New dis- tribution records are provided for S. ornani off Pakistan, S. acuminata off Somalia, Kenya and Madagascar, S. hieronis and S. simoniana off Kenya, S. joubini, S. incerta, S. confusa. S. burnupi, S. hieronis, S. simoniana, S. officinalis vermiculaia and S. zanzibarica on Saya-de-MaIha Bank The Eastern At- lantic species S. elegans was recorded from the SW Indian Ocean on the Agulhas Bank Four species were found at shal- lower depths and seven at greater depths than recorded pre- viously Five species are recorded at depths greater than 300 m Key words Sepiida, Sepia, cuttlefish, species, morphology, distribution, new records INTRODUCTION The cuttlefish (Sepiidae) fauna of the Indian Ocean is incompletely known The area has been relatively well studied in the northern part of the ocean (Adam, 1959; Adam & Rees, 1966; Silas ct ai, 1986), in the south- western part off South Africa (Voss, 1962; Roeleveld, 1972) and to some extent in the southeastern part off Western Australia (Adam, 1979). However, the Western Indian Ocean between the Gulf of Aden and Southern Mozambique, including many islands, archipelagoes and shallow banks (Socotra, Seychelles, Madagascar, Saya-de- Malha, Cargados-Carajos) remain almost unexplored. This broad area was studied during the last several years by various Soviet expeditions organized by many USSR in- stitutions; All-Union (now Russian) Research institute of Marine Fisheries and Oceanography (Moscow), Southern Research Institute of Fisheries and Oceanography (Kerch), Institute of Oceanology of the LiSSR (now Rus- sian) Academy of Sciences (Moscow), Institute of the Biology of the Southern Seas, Ukrainian Academy of Sciences (Sevastopol), Moscow State University, "Hy- dronaut' Base (Sevastopol), etc. As a result of these ex- peditions large amounts of data were gathered, partic- ularly on the taxonomy, distribution and biology of cut- tlefishes (Zuev ct al., 1968; Zuev, 1971; Druzhinin & Filippova, 1974; Homenko, 1976, 1983, 1989; Nesis, 1980, 1993; Homenko & Khromov, 1984; Khromov, 1982, 1987, 1988a; Filippova & Khromov, 1991; Khromov et ai, 1991). As a result of these expeditions, the sepiid fauna of the Western Indian Ocean was found to be richer and more diverse than was previously supposed. Ten new species have been described to date, and others are await- ing description. Some species are common and may have commercial potential. The data gathered permit a fresh examination of the composition, distribution and rela- tionships of the sepiid fauna of the Indian Ocean as a whole. We present new data on the distribution of several known species, and on the morphology of some species described incompletely or inadequately. All but one are inhabitants of the Indian Ocean — Sepia elegans is an Atlantic species recorded for the first time in the Indian Ocean off South Africa. This work is based primarily on the catches made in 1976 by the R/V "Professor Mesyatzev" off East Africa between 2°S and 25°S, off the Cargados-Carajos Islands and on Saya-de-Malha Bank, and by the R/V "Vityaz" in 1988-89 oft Socotra, Madagascar, Mozambique and on Saya-de-Malha Bank. We have also studied collections of many other expeditions, and are grateful to the col- lectors of the following material: G. M. Belyaev ("Ob", 1956), V. N. Semenov ("Akademik Knipovich ', 1965- 66), the late V. I. Chekunova ("Akademik Knipovich", 1967), the late S. V. Mikhailin ("Gizhiga", 1975), B. G. Ivanov ("Professor Mesyatzev", 1975-76), L. S. Hom- enko ("Nauka", 1976, "Zheleznyakov", 1977, "Yelsk", 1977-78), N. S. Khromov ("Fiolent", 1976, "Sabah", 1979), V. A. Spiridonov ("Odissey", 1980), M. G. Kar- pinsky ("Professor Mesyatzev", 1983), D. L. Ivanov ("Akademik Petrovsky", 1984), Yu. S. Korzun ("Skif", 1986), V. A. Bizikov and D. O. Alekseyev ("Akademik Knipovich", 1984, "Odissey", 1985, "Gidrobiolog", 1987). All specimens are either in the Russian Research Institute THE NAUTILUS, Vol. 108, No. 3 ^f^c: -"ii Yii. A. Filippova et al , 1995 Page 69 of Marine Fisheries and Oceanography (VNIRO), the Zoological Museum of Moscow llniversity (ZMML'), or in the Institute of Oceanology of the Russian Academy of Sciences (lOAS), Moscow. SYSTEMATICS Family Sepiidae 1. Sepia (Doratuscpion) jotibini Massv, 1927 (Figs. 1, 2, 20) Material examined: "Vitvaz", 25/11/1988, sta. 2634a, bottom trawl, 90-92 m, 25°05'S, 34°50'-34°44'E: 1 fe- male ML 42 mm, lOAS; "Professor Mesyatzev", 24/01/ 1976, sta. 64, bottom trawl, 167-170 m, 24°23'S, 35°29'E: 1 female ML 64 mm, ZMMU 327. Description and remarks: This species was described by Massy (1927), redescribed (syntypes) by Adam and Rees (1966), and again redescribed (topotypes) by Roe- leveld (1972), In spite of the large number of described specimens of both se.xes, the sepions of S. joubini remain nearly unknown because they were damaged and de- calcified in both the syntypes and the topotvpes. Adam and Rees (1966) doubted the validity of S. joub- ini. According to Roeleveld (1972) the cuttlebones of the topotypes were narrow and elongated, pointed anteri- orly, with a median longitudinal ridge dorsally and groove ventrally. The striated zone is convex on both sides of the median groove, angular anteriorly, becoming flat- tened posteriorly, with convex and wavy striae posteri- orly. The inner cone has narrow limbs that form a ledge posteriorly. The outer cone has posterior wings. Roelev- eld (1972) also mentioned a peculiar structure at the base of the spine, but it is not known whether it occurs in all specimens. Roeleveld's figure of S. joubini illustrates an expansion of the posterior part of the imier cone that is not described in the text With this discrepancy and her description of a decalcified sepion, unambiguous iden- tification of this species is difficult. There are two females of S. joubini in our material, one (sta. 2634a) properly fixed and in good condition. Therefore we are able to add several important details to the description of this species. The dorsal surface of the mantle of the living animal has small red spots and interrupted light-colored stripes along the fins that form a series of elongate, raised bolsters as in S. (D.) peterscni Appelloef, 1886, and S. (D.) viet- namica Khromov, 1987. The fins are narrow and slightly expanded posteriorlv The characteristic arms of the fe- male differ in length (lateral arms longer than the dorsal and ventral), have attenuated tips and well developed protective membranes (Fig. 1), and correspond to Roe- leveld s (1972) description. Longitudinal red stripes are found on the dorsal side of the head and on dorsal and lateral arms of living animals, but are not evident on fixed specimens. The sepion is dark-cream colored, nar- rowly elongate, width 24*^(, thickness 9% of length. A metlian rib, not limited b> grooves, lies on the dorsal surface of the sepion. Only the median third of the dorsal shield is calcified and bordered by wide, chitinous mar- gins. The sepion is semi-rounded in cross-section, with a flattened middle part of phragmocone. The median groove, shallow on the striated zone and deep on the last loculus, is clearly marked along the whole length of the phragmocone. The striated zone occupies the entire width on the ventral side, and is bordered by smooth marginal zones. Anterior striae are reversed V-shaped. The inner cone forms a ledge that is slightly expanded posteriorly, but without any blade-like structure, as in Roeleveld s figure. Inner cone limbs are close to the chitinous mar- gins. The outer cone forms a regular, cup-like, decalcified structure posteriorly. The spine is long, narrow at its base, slightly expanded in the middle part, and spindle-like in lateral view. We confirm Roeleveld s opinion on the validity of S. joubini and consider this species to be fully described now. A female (ML 64 mm) in these collections (ZMMU) is the largest known specimen. Distribution: Sepia joubini was recorded off South- eastern Africa, from the Tugela River to Cape Natal (Roeleveld, 1972). We foimd this species off Southern Mozambique, 24-25°S. The depth range of 66-134 m (Roeleveld, 1972), is expanded to 66-170 m 2. Sepia (Doratosepion) incerta Smith, 1916 (Figs. 3, 4, 20) Material examined: "Professor Mesyatzev , 18/01/1976, sta. 41, bottom trawl, 100 m, 18°29'S, 37°17'E: 3 males, ML 44, 115, 118 mm, 2 females. ML 82, 88 mm; "Pro- fessor Mesyatzev ', 19/01/1976, sta. 44, bottom trawl, 152-172 m', 19°07'S, 36°59'E: 1 male, ML 123 mm; "Pro- fessor Mesyatzev", 24/01/1976, sta. 64, bottom trawl, 167-170 m', 24°23'S, 35°29'E: 2 males, ML 53, 111 mm, all specimens are in the collections of ZMMU, No 185, 212, 230, 232, 334. "Professor Mesyatzev", 31/01/1976, sta. 77, bottom trawl, 330-345 m, 25°21'S, 34°19'E: 1 female, ML 82 mm, VNIRO. "Vityaz", 25/11/1988, sta. Figures L 2. Sepia (Doratosepion) jotihini Massy, 1927. MI. 42 mm, 2.5°05'S. 34°.50'-34°44'E, 90-92 mm, lOAS, uncatalogued 1. Head of female. 2. Sepion of female. Figures 3, 4. Sepia {Duratusepion) incerta Smith, 1916, ML 11.5 mm, 1S°29'S, 37°17'E, 100 m, ZMMU No. 212. 3. Dorsal and 4. lateral views of sepion of male. Figures 5, 6. Sepia (Doratosepion) confusa Smith, 1916. ML 103 mm, 24°23'S, .35°29'E, 167-170 m, VNIRO, uncatalogued 5. Sepion of male 6. Posterior part of male sepion Figure 7. Sepia (Doratosepion) trygonina Rochebrune, 1884. ML 65 mm, Saya-de-MaIha Bank, VNIRO, uncatalogued Sepion of male. All scale bars = 1 cm. as, anterior striae; ds, dorsal shield; ic, inner cone; lie, limbs of inner cone; II, last loculus; mg, median groove (or furrow); oc, outer cone; p, phragmocone; s, spine; smz, smooth marginal zone; sz, striated zone; s, spine; w, wings of outer cone Page 70 THE NAUTILUS, Vol. 108, No. 3 cm Yu. A. Filippova et al., 1995 Page 71 2634a, bottom trawl, 90-92 m, 25°05'S, 34°50'-34°44'E: 1 male ML 45 mm. 1 female, ML 91 mm, lO.^S. Description and remarks: All our specimens are quite typical and agree completely with the descriptions of Adam and Rees (1966) and Roeieveld (1972) Large males are characterized by a peculiar transformation of the dorsal arms. Males and females differ from all other species in having a furrow on the posterior ventral part of the sepion inner cone. In the largest female (ML 91 mm) the eggs from the ovary are very elongate (8.9 x 3.2 mm). Distribution: Sepia inccrta was recorded off eastern Cape Province and Natal, East London to Durban (Massy, 1925). Sepions were also found from Port Elizabeth to Southern Mozambique (Roeieveld, 1972). We found the animals from North to South Mozambiciue, 1S°29' to 25''2rS, at depths of 90-345 m, significantly deeper than previous records for this species (70-79 m; Roeieveld, 1972). 3. Sepia (Doratosepion) confusa Smith, 1916 (Figs. 5, 6, 20) Material examined: "Professor Mesyatzev", 06/01 / 1976, sta. 25, bottom trawl, 335 m, 05°19'S, 39°09'E: 3 females, ML 64, 69, 69 mm; "Professor Mesyatzev". 22/01/1976, sta. 57, bottom trawl, 150-170 m. 21°27'S, 35°32'E: 1 male, ML 70 mm; "Professor Mesyatzev ', 24/01/1976, sta. 64, bottom trawl, 167-170 m, 24°23'S, 35°29'E: 5 males, ML 91-148 mm; ""Professor Mesvatzev ', 25/01/ 1976, sta. 65, bottom trawl, 256-258 m, 24°43'S, 35°26'E: 5 males, ML 62-117 mm; "Professor Mesyatzev ", 25/ 01/1976, sta. 66, bottom trawl, 200-210'm, 24°57'S, 35°07'E: 90 males, ML 48-135 mm; ""Professor Mesy- atzev", 25/01/1976, sta. 67, bottom trawl, 56-53 m, 24°51'S, 35°05'E: 1 male, ML 120 mm, some of the spec- imens are in the collection of ZMMU, Nos. 206, 215. ""Vityaz", 25/11/1988, sta. 2635, bottom trawl, 225-228 m, 25°05'-25°06'S, 35°15'E: 5 females ML 63-68 mm, 30 males 75-115 mm; some of the specimens are in the collection of lOAS, (there were nearly 600 specimens of both sexes of this species in this catch, but males greatly outnumbered females). Description and remarks: Sepia confusa males differ from all other species in having a long "tail" (Adam & Rees, 1966; Roeieveld, 1972). The length of this structure. formed by fins fused or not fused posteriorly, depends on the stage of maturity. Mature males have long tails that are absent in immature males and in females. The tail first appears at the 2nd maturity stage, and begins to grow exponentially at ML about 90 mm, at which size males change from the 3rd to the 4th maturity stage. In females, the dorsal arms have expanded membranes and two rows of suckers distally as in S. joubini. The lateral arms have the same two rows of suckers distally, but the protective membranes do not expand. Dorso-lateral arms are not attenuated and are subequal in length. The sepion differs from other species in having a dorsal posterior keel and a ventral margin of the outer cone that is in- verted to the outer side, triangular and thickened me- dially. Egg size in a female (ML 72 mm) was 5.0-5,1 x 2.6- 2.7 mm. Distribution: Sepia confusa occurs off eastern and southeastern Africa (5° to 30°S). The northernmost re- cords are off Zanzibar (Adam & Rees, 1966) and Pemba (our data). We found it to be very common along the coast of Mozambique, 21°30'-25°S. The record of this species from the Arabian Sea (Zuev, 1971), is erroneous, while a record from Saya-de-Malha Bank (Nesis, 1982) was not confirmed by more recent expeditions (Nesis, 1993). This species was recorded at depths of 53-352 m (Roeieveld, 1972; our data). 4. Sepia {Doratosepion) burnupi Hoyle, 1904 (=Sepia exsignata Barnard, 1962) (Fig. 20) Material examined: ""Professor Mesvatzev", 23/01/1976, sta. 62, bottom trawl, 235-240 m,' 23°29'S, 35°33'E: 2 males, ML 54, 56 mm, ZMMU 221. Description and remarks: This species was hitherto known from three sepions and three specimens taken off Natal (Massy, 1925; Barnard, 1962; Roeieveld, 1972). The two males of S. Inirntipi in our collection are practically identical with Roeieveld s (1972) description and figures of this species. Distribution: Sepia burnupi is a very rare species. Our collections enlarge its known range to Southern Mozam- bique, and known depth distribution from 40-48 m to 235-240 m. Figure 8. Sepia (Anomalosepia) omani Adam, 1939 Posterior part of female sepion. Redrawn from Adam & Rees, 1966, pi. 22, fig. 144. Figures 9-1 L Scpia (Rhomhosepion) acuminata Smith, 1916. 9. ML 84 mm. 05°19'S. 39°09'E, 335 m, ZMMU No. 204. Atypical sepion of female with furrow 10-1 1. ML 87 mm, 24°25'S. 3.5°29'E. 170 m, VNIRO uncatalogued 10. Ventral and 11. dorsal views of typical sepion of female with rib. Figures 12, 13. Sepia (Rhonthosepion) hieronis Robson, 1924 ML 62 mm, 04°47'S. 39°24'E, 180-185 m, ZMMU No. 227. 12. Ventral and 13. lateral views of sepion of a male. Figure 14. Sepia (Acanthosepion) zanzilmrica Pfeffer, 1884. ML 92 mm, 12°22'S,54°28'E, 50 m, ZMMU No, 317, Sepion of male. Figure 15. Sepia (Acanthosepion) prashadi Winckworth, 1936, ML 90 mm, 04°47'S, 39°24'E, 180-185 m, ZMMU No. 249, Sepion of male. Figures 16, 17. Sepiella cyanea Robson, 1924. ML 60 mm, 17°13'S, 38°39'E, 14 m, ZMMU No. 207. 16. Ventral and 17. lateral views of sepion of male. Figures 18, 19. Sepiella inermis (Ferussac & d'Orbigny, 1835). ML 66 mm, Kuwait territorial waters, 0-20 m, ZMMU No. 198. 18. Ventral and 19. lateral views of sepion of male. Scale bars = 1 cm. cm, chitinous margin of sepion; ds, dorsal shield; gr, grooves beside median ridge; mr, median ridge (or rib). Page 72 THE NAUTILUS, Vol. 108, No. 3 40°E 60°E 20° I I I I I I I I -r- r I I 1 I 1 II 20" 40-E 60°E f'igure 20. Distribution of Sepia (Doratoscpion) in the western Indian Ocean. Triangles = Sepia (Doratosepiun) jouhini, cir- cles = S. (D.) incerta, crosses = S. (D.)co7ifusa, stiuare = S.(D.) burnupt. 5. Sepia (Doratosepion) trygonina Rochebrune, 1884 (Figs. 7, 21) Material examined: "Ob", 08/06/1956, sta. 150, bot- tom trawl, 50 m, 11°50'N, SPIO'E: 8 females, ML 53- 85 mm; "Ob", 10/06/1956, sta. 152, bottom trawl, 30 m, 12°05'N, 44°12'E: 1 male, ML 48 mm, 1 female, ML 36 mm; "Professor Mesyatzev", 19/12/1975, sta. 2, bot- tom trawl, 67-77 m, 2°'24'S, 40°56'E: 2 males, ML 110, 41 mm; "Yelsk", December 1977 - January 1978, Saya- de-Malha Bank, 205 m, some sepions; "Professor Mesy- atzev", 27/03/1983, sta. 14, bottom trawl, 200 m, 1 r30'S, 60°49'E: 2 females, ML 46, 53 mm; "Akademik Petrov- sky", 20/01/1984, sta. 67, Sigsbee trawl, 55 m, 15°07'N, 41°41'E: 1 female, ML 28 mm; "Odissey", 27/05/1984, sta. 9, bottom trawl, 28 m, 12°19'N, 53°22'E: 2 males, ML 51, 59 mm, 8 females, ML 51-74 mm; "Odissey", 27/05/1984, sta. 10, bottom trawl, 27 m, 12°14'N, 53°46'E: 1 female, ML 31 mm; "Odissey", 19/04/1985, sta. 95, Sigsbee trawl, 80 m, 12°21'N, 54°28'E: 1 male, ML 30 mm; "Odissey", 19/04/1985, sta. 96, bottom trawl, 100 m, 12°19'N, 54°25'E: 2 males, ML 47-60 mm; "Skif", 01/11/1986, sta. 140, bottom trawl, 20 m, 15°35'N, 52°32'E: I male, ML 47 mm, 1 female, ML 45 mm, 1 juv, ML 25 mm; "Gidrobiolog", 01/05/1987, sta. 2, bot- tom trawl, 50 m, 13°37'N, 47°23'E: 3 femaie.s, ML 47, 49, 61 mm, 5 juv, ML 30-20 mm; "Gidrobiolog", 16/ 05/1987, sta. 6, bottom trawl, 33 m, 14°47'N, 50°01'E: 2 females, ML 52, 62 mm; "Gidrobiolog", 29/05/1987, sta. 10, bottom trawl, 50 m, 13°55'N, 48°29'E: 5 females, ML 52-64 mm; "(iidrobiolog", 10/05/1987, sta. 18, bot- tom trawl, 37-40 m, 11°56'N, 53°44'E: 2 females, ML 71, 74 mm; Gidrobiolog", 28/05/1987, sta. 18, bottom trawl, 40 m, I3°59'N, 47°57'E: 6 females, ML 37-63 mm, collections of VNIRO and ZMMU, Nos. 184, 197, 304, 310, 314, 316, 319, 324, 331. Remarks: Sepions of S. trygonina are very similar to those of . Imn^iipi, but the soft bodies of these species are distinctly different. InS. trygonina, tliearmsof males are not transformed as in S.burnupi and the lateral arms of females have two widely spaced rows of suckers in the distal portions. Distribution: Scpia trygonina was first described from the Red Sea and later reported from different areas of the northwestern Indian Ocean: from the Gulf of Aden and Kuria-Muria Islands to the Gulf of Mannar (Adam & Rees, 1966; Druzhinin & Filippova, 1971; Sarvesan, 1976). We found this species south of the ecjuator on the Saya-de-Maiha Bank, at 15°35'N to 1 1°30'S, at depths of 20 to 205 m. Mantle length reaches 110 mm in males, 85 mm in females, and is much larger than reported previously. 6. Sepia (Doratosepion) niascaren.sis Filippova and Khromov, 1991 (Fig. 21) Material examined: "Professor Mesyatzev", 04/04/1976, sta. 128, bottom trawl, Kargados-Karajos Islands: 1 male, ML 105 mm; "Professor Mesyatzev" 05/04/1976, sta. 129, bottom trawl, 316-325 m,'l5°45'S, 61°15'E: 1 male, ML 90 mm; "Professor Mesvatzev", 27/03/1983, sta. 14, bottom trawl, 200 m, 11°36'S, 60°49'E: 20 males ML ? (in poor state); "Professor Mesyatzev", 27/03/1983, sta. 15, bottom trawl, 195 m, 11°31'S, 6r00'E: 15 males, ML 67-124 mm; "Yunaya smena", 1985, Saya-de-Malha Bank: dry and broken sepions, collections of VNIRO and ZMMU, Nos. 233, 234, 349, 350. "Vitvaz", 07/01/1989, sta. 2803, bottom trawl, 87-110 m, 11°21'-11°22'S, 61°47'E: 7 females, ML 50-63 mm, lOAS. Description and remarks: This species is fully described from males (Filijipova & Kliromov, 1991) and females (Khromov et al.. 1991 ). Our specimens agree completely with these descriptions. The males of ii. mascarensis are characterized by their untransformed arms, the absence of a hectocotylus, and a narrow and relatively thick se- pion that has a rib on its ventral side and reversed, W-shaped, ventral striae. The females of this species have long second arms with two rows of suckers distally and narrow sepions with reversed, W-shaped ventral striae as well as a furrow (not a rib) on the ventral side. Distribution: Scpia mascarensis is reported from the Mascarenc Ridge, the Sa\a-de-Mailia Bank and shoals around the Kargados-Karajos Islands, at depths of 87- 325 m. 7. Sepia (Dorato.sepiim) arahica Ma.ssv, 1916 (Fig. 21) Material examined: "Skif", 01/1 1/1986, sta 10, bottom Yu. A. Filippova et al., 199.5 Page 73 trawl, 20 m, 15°.35'N, 52°32'E: 1 male, ML 88 mm. col- lection of VNIRO. Description and remarks: This specimen agrees with the description and illustrations of Adam and Rees (1966). This mature male with a developed hectocotylus is the largest recorded specimen of this species. Distribution: Red Sea, Gulf of Aden to southern India. 8. Sepia (Doratoscpion) sp. (Fig. 21) Material examined: "Vitvaz", 07/01/1989, sta. 2804, bottom trawl, 230-235 m, ll°06'-ir09'S, 62°14'-62°13'E: 1 female ML 65 mm, lOAS. Description and remarks: Mantle elongate oval, ante- rior margin somewhat produced dorsally, emarginate ventrally. Arms subequal in length, lateral arms slightly longer. Arms pairs 1, II, and III with well developed, spade-like protective membranes covering the suckers on the distal third of arms. The suckers are cjuadriserial proximally, becoming biserial and widely spaced distalK . The fourth arms have quadriserial suckers. Tentacular club with 5 rows of suckers, differing slightly in size, with the suckers of the central row the largest. Sepion is cream colored, narrow, elongate, with a rib, delimited by furrows dorsally and a median groove ventrally, and slightly W-shaped in cross-section. The striated zone oc- cupies the entire width of the ventral side, with the anterior striae semi-rounded, slightly emarginated in the ventral groove. The inner cone forms a ledge that is slightly raised posteriorly, but not expanded. The outer cone is cup-like and calcified, forming a toothed structure on the ventral margin similar to that of S. incerta (Roe- leveld, 1972, Fig.6,d). The spine is long, lacks a base and spindle-like laterally. This female is similar to one of S. conjusa. but the sepion differs distinctly by the semi-rounded anterior striae, the absence of an inverted, triangle-shaped ventral margin on the inner cone, and a posterior dorsal keel on the sepion. It also resembles S. joubini, but the arms of the latter species are markedly longer and attenuated at their ends. Moreover, the striae of S. joubini are reversed V-shaped, its sepion is semi-rounded in cross-section, its inner cone is slightly expanded posteriorly, and its outer cone is decalcified. The sepion of Sepia sp. can be easily confused with that of S. incerta, but it has no groove on the posterior portion of the inner cone. We could not identify Sepia sp. in spite of the good condition of the specimen. Possibly it is a new species, but the description of a new species in so complex a subgenus as Doratosepion would not be appropriate in the absence of a male specimen. 9. Sepia (Anomalosepia) omani Adam and Rees, 1966 (Figs. 8, 22) Material examined: "Akademik Knipovich", 19/01/ 1966, sta. 310, bottom trawl, 210 m, 23°42'N, 66°14'E: 2 males, ML 78, 80 mm, 1 female, ML 85 mm, 3 juv, ML 30, 31, 35 mm, ZMMU 214. Figure 21. Distribution of Sepia (Doratosepion) in the western Indian Ocean. Circles = S. {D.) trygonina, triangles = S. (D.j mascarensis, square = S. (D.) arabica, cross = S.(D.) sp. Description and remarks: Sepia omani is the only spe- cies with two pairs of outer cone wings on the sepion. Thus, the specimens referred to S. omani by Voss and Williamson, 1971 are certainly not correctly identified. The opinion (Khromov, 1988b) that these animals are S. rex is also erroneous. The Hong Kong specimens probably belong to a new species. Distribution: Sepia omani was known only from the Gulf of Oman, at a depth of 201 m. We found it in the northern part of the Arabian Sea, off Pakistan, at nearly the same depth, 210 m. 10. Sepia (Rhombosepion) acuminata Smith, 1916 (Figs. 9-11, 22) Material examined: "Professor Mesyatzev", 19/12/1975, sta. 1, bottom trawl, off North Kenya: 1 male, ML 93 mm, 1 female, ML 75 mm; "Professor Mesyatzev ', 21/ 12/1975, sta. 3, bottom trawl, 290-295 m, 3°02'S, 40°26'E: 1 male, ML 84 mm, 1 broken sepion; "Professor Mesy- atzev", 04/01/1976, sta. 23, bottom trawl, 44 m, 04°03'S, 40°00'E: 1 male, ML 73 mm, 1 female, ML 75 mm; "Professor Mesyatzev", 06/01/1976, sta. 25, bottom trawl, p.... . - 1 THE NAUTILUS, Vol. 108, No. 3 Figure 22. Distribution of Sepia (Anomalosepia) and Sepia (Rhombosepion) in the western Indian Ocean Triangle = Sepia (Anomalosepia) omani, circles = Sepia (Rhombosepion) acu- minata, squares = S.(R.) hieronis, cross = S. (R.) elegans. 335 m. 05°19'S, 39°09'E: 1 male, ML 102 mm, 3 females, ML 56, 100, 111 mm; "Professor Mesyatzev", 24/01/ 1976, sta. 64, bottom trawl, 170 m, 24°25'S, 35°29'E: 1 sepion 84 mm; "Zheleznyakov ', 07/03/1977, bottom trawl, 230-235 m, 0r31'N', 44°39'E: 1 male, ML 55 mm, 2 broken sepions, collections of VNIRO and ZMMU, Nos. 191, 192, 204, 225, 242. "Vityaz", sta. 2635, 25/11/1988, bottom trawl, 225-228 m, 25°05'- 25°06'S, 35°15'E: 11 females, ML 84-119 mm, 14 males, ML 52-91 mm, 1 juv ML 31 mm, collection of lOAS, 66 unsexed and unmeasured specimens of the same species; "Vityaz' , sta. 2644, 02/12/1988, bottom trawl, 325-332 m, 22°19'- 22''23'S, 43°06'E: 4 females ML 89-115 mm. I sepion 110 nmi, collection of lOAS. Description and remarks: We note an interesting vari- ability of sepions in this species. Typical sepions of S. acuminata caught in South African waters (Port Eliza- beth) show a distinct median rib and a faint indication of two lateral ribs on the dorsal surface (Adam & Bees, 1966). These authors also reported on three ailditional sepions from Mombasa that had a deep, narrow groove instead of a rib Rocleveld (1972) described specimens from Mozambic]ue that had a median ridge sometimes sunken below the level of rest of dorsal surface. Having examined our material, we noted a clear rib on the sepions of specimens from southern Mozambique, a flat dorsal side or one slightly sunken below the rib on sepions from northern Mozambique and Tanzania, and a groove on the sepions of animals from Kenya and southern Somalia. Females of S. acuminata from south- ern Madagascar have a flat dorsal surface on the sepion. Small males look alike, but the largest male has a sepion with a slightly marked groove. We therefore suggest that the presence of a rib or groove may be a geographical variation that is more evident in large animals. Distribution: Sepia acuminata has not been previously reported from off Madagascar, northern Kenya, or So- malia. This species has a wide range along all eastern Africa, from 0r30'N to 30°S, including Madagascar. It was recorded at depths of 44-369 ni (Roeleveld, 1972; our data). Maximum size is presently a mantle length of up to 100 mm in males and 120 mm in females. Egg size in a female of 110 mm ML was 5.0-5.2 x 4. 5-4. 7 mm, while in smaller females it ranged between 4.2-4.9 X 3.3-4.5 mm. Females mature at about 85-90 mm ML. 11. Sepia (Rhombosepion) hieronis Robson, 1924 (Fig. 12, 13, 22) Material examined: "Professor Mesyatzev", 05/01/1976, sta. 24, bottom trawl, 180-185 m, 04°47'S, 39°24'E: 1 male, ML 62 mm; "Professor Mesyatzev", 27/01/1976, sta. 73, bottom trawl, 430-440 m, 25°28'S, 33°32'E: 2 males, ML 42, 48 mm, ZMMU 227, 228. Description and remarks: Our specimens agree com- pletely with the descriptions of Adam and Rees (1966) and Roeleveld (1972). Distribution: Sepia hieronis has been reported in the south-eastern Atlantic, from northern Namibia to Slang- kop and in the western Indian Ocean off southern Mo- zambique and on the Agulhas Bank (Voss, 1962, 1967; Roeleveld, 1972; Okutani & Ilasegava, 1979; Sanchez & Moli, 1984; Sanchez, 1988; Sanchez & Villanueva, 1988). We collected it off Mozambique and Kenya, so S. hi- eronis is quite widely distributed south ot Angola and Zanzibar, but is not reported off South .\frica between the Cape of Good Hope and St. Francis Ba\. The depth range is 43-457 m (Roeleveld, 1972). 12. Sepia (Rhombosepion) elegans d'Orbigny, 1826 (Fig. 22) Material examined: "Ob", 18/03/1957, sta. 263, bot- tom trawl, 110 m, 37°12'S, 22°30'E: 4 males, ML 39-45 mm; "Gizhiga", 15/02/1975, sta. 266, bottom trawl, 180- 200 m, 21°10'S, 13°20'E: 1 female, ML59 mm; "Fiolent", .sta. 136, 12/03/1976, 250 m, 06''26'S, 1 1°36'E: 4 females, ML 73-89 mm; "Odissey" 06/12/1980, bottom trawl, 200 m, 31°25'N, 2riO'\V: 1 male, ML 55 nmi; "Aka- demik Knipovich", sta. 8, 05/01/1967, 180-210 m. West Africa: I male, ML 50 mm; "Akademik Knipnvich", 07/ Yu. A. Filippova et al., 1995 Page 75 06/1969, 80-95 m, 21°50'N, 17°20'W; 1 male, ML 50, 1 female, ML 53 mm; "Akademik Knipovich", sta. 1, 30/ 01/1984, 140-200 m, 23°46'N, 16°51'W; 3 females, ML 36, 39, 46 mm, collections of VNIRO and ZMMU, Nos. 186. 193 222, 226, 307. Description and remarks: Our specimens agree com- pletely with the description of Adam and Rees (1966). Distribution: Sepia clegans is a widely distributed east- ern Atlantic species, reported along the coasts of Europe and West Africa from Ireland to Namibia (Adam & Rees, 1966; Sanchez & Moli, 1984; Sanchez, 1988). We re- corded mature specimens from the Agulhas Bank, so the range of S. elegans extends into the southwestern Indian Ocean. 13. Sepia (Sepia) papillata Quoy and Gaimard, 1832 (Fig. 23) Material examined: "Professor Mesyatzev", 06/04/1976, sta. 131, bottom trawl, 180-210 m. 15°48'S, 59°58'E: 1 male, ML 123 mm, 1 female, ML 107 mm; "Odissey", 12/06/1984, sta. 14, bottom trawl, 35 m, 09°42'S, 61°08'E: 2 males, ML 50, 64 mm, ZMMU 254, 330; "Vitvaz", 08/ 01/1988, sta. 2808, bottom trawl, 58-61 m,' 11°05'S, 62°02'-62°04'E: 5 males ML 37-58 mm, 10 females ML 28-51 mm; "Vityaz", 08/01/1988, sta. 2810, bottom trawl, 57-70 m, 10°15'-10°16'S, 61°09'E: 2 males, ML 63, 65 mm, 1 female, ML 75 mm, lOAS. Description and remarks: This species differs from the previous ones in having a broadly oval sepion with a deep and wide median ventral groove and a considerably narrowed and flat inner cone According to Adam and Rees (1966), there is no spine, but only a blunt knob on the sepion posterior. Roeleveld (1972) described a short spine not exceeding the posterior margins of the shell. Our specimens from Saya-de-MaIha Bank are not typical, as some have a short spine visible from the ventral side of the sepion. Thus, the length of spine of S. papillata is variable. Living animals of S. papillata from "Vityaz" station 2808 were very brightly colored, scarlet and gold, with- out stripes or spots, while specimens from station 2810 were brown with zebra-like stripes and red spots. The color of the animals probably reflects the color of the substrate (crustose red algae at sta. 2808). Fixed animals are not brightly colored and can be easily confused with S. o. vermiculata and S. simoniana. Females mature at about 50 mm ML. Distribution: This species was previously reported along the coast of South Africa from Luederitz Ba\ (Atlantic) to the Tugela River (Indian Ocean), and especially near Cape Town (Roeleveld, 1972; Okutani & Hasegawa, 1979). As we have caught it far from this region, on the Mascarene Ridge, the range of S. papillata is greatly ex- tended. The depth range for our samples is 35-210 m, somewhat deeper than 26-127 m range recorded pre- viously by Roeleveld (1972). Figure 23. Distribution of Sepia (Sepia) in the western Indian Ocean. Triangles = Sepia (Sepia) papillata, squares = S. (S) simoniana, circles = S. (S.) officinalis vermiculata 14. Sepia (Sepia) airnoniana Thiele, 1921 (Fig. 23) Material examined: "Professor Mesyatzev", 05/01/1976, sta. 24, bottom trawl, 180-185 m, 04°47'S, 39°24'E: 1 male, ML 91 mm. 1 female, ML 95 mm; "Professor Mesyatzev". 24/01/1976, sta. 64, bottom trawl, 170 m, 24°23'S, 35°29'E: 5 females, ML 58-99 mm. ZMMU 200, 250. Description and remarks: Sepia simoniana is closely related to S. papillata, but differs in having equal-sized suckers on the long club, and a long spine on the sepion. Distribution: This species was first described from Si- mons Bay. South Africa, and repeatedly reported from the same region, near Cape Town and along the east coast of South Africa to the Tugela River (Roeleveld, 1972). Two specimens were caught on the Agulhas Bank (Adam, 1983). Our material extends the range of S. si- moniana to the area trom northern Kenya to southern Mozambique, at depths 170-185 m (14-134 m reported by Roeleveld, 1972). 15. Sepia (Sepia) officinalis vermiculata Quoy and Gai- mard, 1832 (Fig. 23) Material examined: sta. 46, bottom trawl "Professor Mesyatzev", 19/01/1976, 30-40 m, 19°17"S, 36°22'E: 2 males, ML 82, 87 mm, 1 female, ML 82 mm; "Professor Mes- yatzev", 20/01/1976, sta. 47, bottom trawl, 70-75 m, i9°45'S, 36''22'E: 1 male, ML 98 mm; "Professor Mes- yatzev", 20/01/1976, sta. 50, bottom trawl, 30 m, 19°37'S, 35°43'E: 4 females, ML 87-133 mm; "Professor Mesy- atzev", 26/01/1976, sta. 68, bottom trawl, 44-50 m, 24°54'S, 34°53'E: 1 sepion 93 mm; "Professor Mesy- Page 76 THE NAUTILUS, Vol. 108, No. 3 Figure 24. Distribution of Sepia (Acanthosepion) and Sepia (Hemisepius) in the western Indian Ocean. Circles = Sepia (Acanthosepion) zanzibarica, squares = S. (A.) prashadi, cross = S. (A.) savignyi. triangle = S. (H.) typica. atzev", 13/04/1976, sta. 148, bottom trawl, 25 m, 09°52'S, 60°57'E; 2 males, ML 64, 69 mm, collections of VNIRO and ZMMU, Nos. 224, 251, 252, 253, ■'Vityaz", 25/11/ 1988, sta. 2634a, bottom trawl, 90-92 m, 25°05'S, 34°50'- 34°44'E: 1 male, ML 100 mm, 1 female, ML 135 mm; "Vityaz", 25/11/1988, sta. 2634b, bottom trawl, 95-102 m, 25°05'- 25°06'S, 34°45'E: 1 female, ML 1 18, collection of lOAS. Description and remarks: Our specimens agree com- pletely with the description of Adam ands Rees (1966). Distribution: Sepia officinalis vcrmictilata has been found along southern Africa from 30°S in the southeast- ern Atlantic Ocean to southern Mozambique in the south- western Indian Ocean (Voss, 1962; Adam & Ree.s, 1966; Roeleveld, 1972) and reported off Namibia (Sanchez & Moll, 1981; Sanchez, 1988). According to our data, S. o. vermiculata ranges northward to central Mozambique, 19°S and on the Saya-de-Malha Bank at depths of 25- 102 m. The largest of our specimens are immature (male 100 mm ML and females 1 18 anil 135 mm ML were in the second stage of maturity). Figure 25. Distribution of Sepiella in the western Indian Ocean. Triangles = Sepiella cyanea. circles = S. inermis. 16. Sepia (Acanthosepion) zanzibarica Pfeffer, 1884 (Fig. 14, 24) Material examined: "Odissey", 27/05/1984, sta. 9, bot- tom trawl, 28 m, 12°19'N, 53°22'E: 1 female, ML 104 mm; "Odissey", 12/06/1984, sta. 14, bottom trawl, 35 m, 09°42'S, 61°08'E: 1 female, ML 58 mm; "Odissey", 17/04/1985, sta. 95, bottom trawl, 50 m, 12°22'N, 54°28'E: 9 males, ML 71-139 mm, 4 females, ML 88-115 mm; "Odissey", 19/04/1985, sta. 95, bottom trawl, 40 m, 12°16'N, 53°59'E: 1 male, ML 172 mm; "Odissey", 17/ 04/1985, sta. 94, Iwttom trawl, 40 m, 12°22'N, 54°21'E: 1 female, ML 73 mm, collections of VNIRO and ZMMU, Nos. 317, 326. "Vitvaz", 04/12/1988, sta. 2657, bottom trawl, 40 m, 22°10'- 22°15'S, 43°09'-4.3°10'E: 1 female, ML 204 mm; "Vityaz", 08/01/1988, sta. 2810, shrimp trawl, 57-70 m, l6°16'-10°15'S, 61°09'E: 6 males, ML 57-122 mm, 2 females, ML 108, 178 mm, 1 juv ML 41 mm; Toliara (Madagascar), beach: 1 sepion 118 mm, collection of lOAS. Description and remarks: Our specimens agree \\itli the description of Adam and Rees (1966). Distributinon\ni. The two species (Pseitdotaranis strongi and P. hyperia) live off- shore in moderately deep water (100-400 m). Diagnosis: Shell small (length to 17 nun), spire high, anterior canal short, whorls 6. Axial sculpture lacking, spiral sculpture of two cords emerging on teleoconch and three cords on base; anal sinus shallow, at periphery and coinciding with uppermost cord; lip not projecting. Pro- toconch paucispiral, of 1.2 low, rounded whorls. Radula of marginal teeth attached to membrane; teeth of long, flat type. Remarks: Authors prior to 1971 used Taranis Jeffreys, 1870 for the type species, but the presence of a radula precludes placement in that genus, in which the radula is lacking (see Powell, 1966:55). Earlier (McLean, 1971; 120, fig 40), I illustrated the radula of strongi and as- signed it to Antiplanes (Rectisulciis), a genus now in the subfamily Cochlespirinae, according to Ta\lor et al. (1993). Pseudotaranis differs in having the sinus more shallow, the lip less protracted, the anterior canal shorter, and the protoconch more compressed. The radula lacks the vestigial rachidian tooth and the marginals are not of the bifurcated type indicated for Antiplanes by Kan- tor and Sysoev (1991:122). The flat morphology of the lateral teeth agrees with the subfamily Crassispirinae, as defined by Taylor et al. (1993). Thus, the radular dif- ferences place Antiplanes and Pseudotaranis in different turrid subfamilies. However, on shell characters, Pseu- dotaranis is atypical of Crassispirinae in not having pro- nounced callus developed near the anal notch. NEW NAMES FOR SPECIES LEVEL HOMONYMS Family TROCHIDAE Rafinesque, 1815 Subfamily MARGARITINAE Stoliczka, 1868 Margatites (Costomargarites) baxteri McLean, new- name for Margarites (Pupillaria) rudis Dall, 1919b:364, not Margarita groenlandica var. rudis Morch, 1869:23. Remarks: Margarites (Costomargarites) baxteri was fig- ured by Dall (1921:179, pi. 18, fig. 13, 14) and by Kosuge (1972, pi. 2, fig. 5), but has otherwise been ignored in the literature. It will be treated by me in work in prep- aration as a geographic subspecies (in the Gulf of Alaska) of the boreal M. (Costomargarites) costalis (Gould, 1841). The new name honors the late Rae Baxter for his efforts at reviewing Alaskan mollusks, cut short by his untimely death in 1991. Family TURRIDAE Swainson, 1840 Subfamily COCHLESPIRINAE Powell, 1942 Leucosyrinx kantori McLean, new name for Antiplanes anujeus Dall, 1919a (p. 36, pi. 11, fig. 5), not Leucosyrinx amyctis Dall, 1919a (p. 5, pi. 3, fig. 7). Remarks: The proposal of Leucosyrinx kantori remedies a problem of secondary homonymy that was initiated when Dall (1919a) described a species in Antiplanes that I now consider to be a true member of Leucosyrinx Dall, 1889. Dall (1919a) also proposed a species with the name Leucosyrinx amycus. Note that both species were de- scribed in the same paper. Previously (McLean in Keen, 1971:713), I placed Leucosyrinx amycus Dall, 1919 in the synonymy of Aforia goodei (Dall, 1890). The new name honors Yuri Kantor for his recent work on turri- form gastropods. LITERATURE CITED Abbott, R. T. 1974. American Seashells. Second Edition. New York: Van Nostrand Reinhold, 663 pp. Dall, W. H. 1919a. Descriptions of new species of mollusks of the family Turritidae from the west coast of America and adjacent regions. Proceedings of the United States National Museum 56(2288): 1-86, pis. 1-24. Dall, W. H 1919b Description of new species of Mollusca from the North Pacific Ocean in the collection of the United States National Museum. Proceedings of the L'nited States National Museum 56(2295):29;3-371. Dall, W H, 1921. Summary of the marine shell-bearing mol- lusks of the northwest coast of America, from San Diego, California, to the Polar Sea, mostly contained in the col- lection of the United States National Museum, with illus- trations of hitherto unfigured species. United States Na- tional Museum, Bulletin 112, 217 pp., 22 pis. Grant, U. S, IV and H. R Gale 1931. Catalogue of the marine Pliocene and Pleistocene Mollusca of California and ad- jacent regions. Memoirs of the San Diego Society of Nat- ural History 1:1-1036, pls.1-32. Hoagland, K. E. 1977. Systematic review of the fossil and Recent Crepidula and discussion of evolution of the Ca- lyptraeidae. Malacologia 16(2):353-420. Kantor, Y. I, and A. V. Sysoev. 1991. Mollusks of the genus Antiplanes (Gastropoda: Turridae) of the northwestern Pacific Ocean. The Nautilus 105(4):1 19-146 Keen, A, M. 1971. Sea shells of tropical west America. Second Edition. Stanford Universitv Press, Stanford, 1064 pp., 22 pis. Kosuge, S. 1972. Illustrations of type specimens of Molluscs described by William Healey Dall. National Science Mu- seum, Tokyo, 29 pis. and captions. THE NAUTILUS 108(3):82, 1995 Page 82 McLean, J H. 1971. A revised classification of the family Turridae, wilh the proposal of new subfamilies, genera, and subgenera from the Eastern Pacific. The Veliger 14(1): 11-4-130. McLean, J. H. 1995. Four new genera for northeastern Pacific prosobranch gastropods. The Nautilus 108(2):39-41. Moody, C L. 1916. Fauna of the Fernando of Los Angeles. University of California Publications, Bulletin, Depart- ment of Geology 10(-4):39-62, pis. 1-2. Morch, 0. A. L. 1869 Catalogue des mollusqnes du Spitzberg recueillis par le Dr. H Kroyer pendant le voyage de la corvette La Recherche en juin 1838. Memoires de la So- ciete Malacologique de Belgique 4:7-32 Pilsbry, H. A. 1889-90. Manual of Conchology. Vol. 11 Tro- chidae. Philadelphia, 519 pp., 67 pis, Powell, A. W. B. 1966. The molluscan families Speightiidae and Turridae. Bulletin of the Auckland Institute and Mu- seum, no. 5, 184 pp., 23 pis. Taylor, J. D., Y. \. Kantor, and A. V'. Sysoev 1993. Foregut anatomy, feeding mechanisms, relationships and classifi- cation of the Conoidea ( = Toxoglossa) (Gastropoda). Bul- letin of the Natural History Museum London (Zoology) 59(2): 125- 170. INSTRUCTIONS TO AUTHORS THE NAUTILUS publishes papers on all aspects of the biology and systematics of mollusks. 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Box 7279, Silver Spring. MD 20907-7279, USA. Chdit'^c of (uUlre.ss: Please inform the publisher of your new address at least 6 weeks in advance. All communications should include both old and new addresses (with zip codes) and state the effective date. THE NAUTILUS (ISSN 0028-1.344) is published quarterly by Trophon Corporation, 891 1 Alton Parkway, Silver Spring, M 1)20910. Second Class postage paid at Silver Spring, MP) and additional mailing offices. POSTMASTER: Send address changi's to: THE NAUTILUS PO. Box 7279 Silver Spring, MD 20907-7279 TH EtyNAUTI LUS Volume 108, Number 4 May 11, 1995 ISSN 0028-1344 CONTENTS Bruce A. Marshall A Revision of the Recent Calliostoma Species of New Zealand (Mollusca: Gastropoda; Trochoidea) 83 THE NAUTILUS 108(4):83-127, 1995 Page 83 A Revision of the Recent Calliostoma Species of New Zealand (Mollusca: Gastropoda: Trochoidea) Bruce A. IVIarshall Museum of New Zealand Te Papa Tongarewa P.O Box 467 Wellington, New Zealand ABSTRACT Thirty-three Recent calliostomatids are recorded from New Zealand, ten of which are described as new Calliostoma (Otu- haia) blacki (Dell, 1956) is renamed due to homonymy, and the following taxa are newly synonymized: Venustas tigris chat liamensis Dell, 1950 with Calliostoma (Maurea) tigris (Gmelin, 1791); Maurea punctulata ampla Powell, 1939 and V. punctulata multigemmata Powell, 1952 with C. granti (Powell, 1931); V. cuuperi Vella, 1954 and Thoristclla chath- amensis profunda Dell, 1956 with C. blacki (Powell, 1950), which is resurrected from synonymy under C. joveauxanum (Dell, 1950); Zizijphinus hodgci Hutton, 1875, Z. ponderosus Hutton, 1885, and C carnicolor Preston, f907 with C. sctectum (Dillwyn, 1817); C. undulatum Finlay, 1923 and C. pellucidum spiratum Oliver, 1926 with C. pellucidum (Valenciennes, 1846). Lectotypes are designated for Trochus selectus Dillwyn, 1817, T. pellucidus Valenciennes, 1846, and Calliostoma onustum Odhner, 1924. Key words: Mollusca; prosobranch gastropods, Trochoidea; Calliostomatidae; Calliostoma, systematics. INTRODUCTION The family Calliostomatidae comprises about 250 living species. They occur in all oceans from the intertidal zone to about 3000 meters depth, mostly on rocky ground. All known species are carnivores, most feeding on cnidaria, and sometimes carion, though a few feed exclusively on sponges. The group is particidarK well represented in the New Zealand region, with 33 living endemic species, including some of the worlds largest, notably Callios- toma (Maurea) tigris (Gmelin, 1791), which may e.xceed 100mm in shell height. Besides the 33 Recent species recorded herein, at least double this number are known from the New Zealand Cenozoic (Early Eocene — Early Pleistocene), of which more than half are undescribed. Several of the living species have excellent fossil records in the extensive Plio- Pleistocene deposits of the southern North Island, and fossil material is recorded and discussed where appro- priate. Two of the recorded Recent species, both of which are type species of new genera, together with a new genus for Calliostoma onustum Odhner, 1924 are named elsewhere (Marshall, 1995). Five additional species are known from off the Kermadec Islands (Appendix). Attention is drawn to the fact that calliostoinatid shell morphology tends to become more variable with increas- ing size/age, so that species with dissimilar early teleo- conchs can be superficially similar at maturity and vice versa. For accurate discrimination of species (and for objective descriptions), it is thus essential to trace and compare the development of individual sculptural ele- ments from the earliest teleoconch whorls following the system used by Ikebe (1942) and Marshall (1988, 1995) (i^igure 28). ABBREVIATIONS AND TEXT CONVENTIONS AUG Geology Department, Auckland University BMNH The Natural History Museum, London MNHN Museum National d'Histoire Naturelle, Paris MNZ Museum of New Zealand, Wellington NZGS Institute of Geological and Nuclear Sciences, Lower Hutt NZOI National Institute of Water and Atmospheric Research, Wellington sa spire angle (see below) ZMA Zoological Museum, Amsterdam Spire angle measurements for individual shells were av- eraged (mean spire angle) to reduce bias induced by cyrtoeonoid spire profile and expanded or narrowed last adult whorl. In other words, for a shell with a cyrtoeonoid spire outline, which becomes more narrowly conical with increasing shell size, the given spire angle is the mean of the maximum and minimum spire angles measured from that specimen. Vice versa for a shell with an evenly conical or coeloconoid contour in which the last adult whorl expands more rapidly than the previous ones. In illustrations of shells, height dimension precetles diam- eter. Readers requiring more detailed information on New Zealand stratigraphy should refer to Fleming (1953), Beu and Maxwell (1990), and Abbott and Carter (1994). Page 84 THE NAUTILUS, Vol. 108, No. 4 SYSTEMATICS Order Vetigastropoda Salvini-Plawen, 1980 Siiperfamily Trochoidea Rafinesqiie, 1815 Family Calliosiomalidae Thiele, 1924 Genus Calliostoma Swainson, 1840 Calliostoma Swainson, 1840:218, 351. Type species (by sub- sequent designation of Herrmannsen, 1846:154): Trochus conulus Linnaeus, 1758; Recent, north-eastern Atlantic and Mediterranean For further discussion and synonymy, see Marshall (1995). Subgenus Maurea Oliver, 1926 Maiirea Oliver, 1926:108. Type species (by original designa- tion): Trochus tigris Gmelin, 1791; Recent, New Zealand [20th December 1926-see below]. Maitriella Oliver, 1926:109. Type species (by original desig- nation): Trochus punctulatus Martyn, 1784; Recent, New Zealand [20th December 1926]. Calliotropis Oliver, 1926:110 Type species (by original des- ignation): Trochus cunninghami Gray, 1834 = Trochus selectus Dillwyn, 1817; Recent, New Zealand Not Cal- liotropis Seguenza, 1903 [20th December 1926] Mucrinops Finlay, 1926:360. Type species (by original desig- nation): Ziziphinus spectabilis A. Adams, 1855); Recent, New Zealand [23 December 1926]. Venustas Finlay, 1927:360. Type species (by original desig- nation): Trochus tigris Gmelin, 1791; Recent, New Zea- land [10 March 1927]. Officially rejected name (ICZN Opinion 479). Not Venustas Allan, 1926. Calotropis Thiele, 1929:49. Replacement name for Calliotropis Oliver not Seguenza Remarks: As indicated by Ben et al. (1969), Venustas Allan, 1926 (Allan, 1926) was published on 7th December 1926 (ICZN opinion 479) and has priority over Maurea Oliver, 1926 (and the officially rejected name Venustas Finlay, 1927). Fortunately the type species of Venustas Allan (Calliostoma fragile Finlay, 1923; Early Miocene, New Zealand) seems unlikely to be consubgeneric with Calliostoma {Maurea) tigris (Gmelin, 1791), the type species of Maurea. The paper in which Oliver (1926) introduced Maurea was published in Parts II / III of Volume 17 of the Proceedings of the Malacological So- ciety of London, the date of publication of which was interpreted as 20th December 1926 by Dell (ICZN Opin- ion 479) on the basis of a letter from L. R. Cox to M. K Mestayer bound into the MNZ volume of the journal. This letter is dated 19th April 1927. The title page to Volume 17, issued on 30th December 1927, states that Parts II and III were issued on 30th December 1926. Whereas I have been unable to trace any firmer evidence that Parts II and 111 were published prior to 30th De- cember, for the sake of noinenclatural stability it is ap- propriate to follow ICZN acceptance (right or wrong) of 20th December 1926, otherwise Maurea would fall as a junior synonym of Mucrinops Finlay, 1926 (23 Decem- ber). Maurea is not endangered by Venusta.s Finlay, which has been deemed to have been published on lOth March 1927 and officially rejected (ICZN Opinion 479). The type species of Maurea, Calotropis, Mauriella, and Mucrinops are similar to the type species of Cal- liostoma in radular morphology and external anatomy. Although I am unable to justify genus-level status for Maurea. it would be inappropriate to treat it as a syn- onym of Calliostoma because all of the New Zealand species are strongly dissimilar to the type species of Cal- liostoma in shell morphology. On the other hand, there is strong mosaic overlap between the type species of Maurea. Calotropis, Mauriella, and Mucrinops via the other Recent species herein referred to Maurea, and I am unable to justify segregation of these genus-group taxa from each other. Accordingly, they are all inter- preted as synonyms of Maurea, which in turn is inter- preted as a subgenus of Calliostoma. Although the origins and relationships of the Recent species are obscure, it is nevertheless clear that Maurea as here limited is poly- phyletic. In whatever way the Recent species are ar- ranged in groups, most or all contain species of excep- tionally large size for the family. With the exception of "Trochus" mutus Finlay, 1924 and " Benthastelena" su- sanae Maxwell, 1992 (Late Eocene — Early Miocene), which are not closely related to any taxa living in the New Zealand region, none of the numerous pre-Pliocene species from New Zealand exceeds much more than about 20 mm in maximum shell dimension, so it would seem that gigantism occurred independently in several species groups after the Miocene. Many of the pre-Pliocene spe- cies are similar to the type species of Fautor Iredale, 1924 (Ziziphinus compttis A. Adams, 1854; Recent, southern Australia) and other species referred there (as a subgenus of Calliostoma) by Marshall (1995) from the New Caledonia area. Calliostoma regale new species and C. aupourianum new species from northern New Zea- land are similar to these small-shelled New Caledonian Fautor species and to many of the New Zealand pre- Pliocene taxa, and because (independently derived) gi- gantism alone is no criterion for genus-group discrimi- nation, it is thus difficult to justif\ segregation of Maurea even from the prior Fautor. EvidentK highlv conser- vative external anatomy and gross shell and radular mor- phology are inadequate to construct objectively definable supraspecific groupings, or rather, real phylogenetic groups are rendered nebulous and are obfuscated due to conservation, convergence, gigantism, and uncertain character-state polarity. Molecular cladistic techniques would seem to be a promising source of data for reso- lution of these problems. For more detailed discussion see Marshall (1995). Calliostoma (Maurea) tigris (Gmelin, 1791) (Figures 1-9, 110, 127) Chemnitz, 1781:100, pi 170, figs 1654, 16,55 Trochus tigris Martyn, 1784, fig 75, (Officially rejected name — ICZN Opinion 479). Trochus granatuvi Gmelin, 1791:3584 (refers to Chemnitz, 1781. pi. 170, figs. 1654, 1655); Lamarck, 1822:26; Fischer, 1875:69, pi 15, fig. 1. Trochus tigris Gmelin, 1791:3585 (refers to Martyn, 1784, fig. Bruce A. Marshall. 1995 Page 85 75) (Officially accepted name — ICZN Opinion 479); Plii- lippi, 1848;50, pi. 10, figs. 16, 17. Turbo granalum — Rixling, 1798:88. Ziziphmus tigris — Gray, 1843:237. Ziziiphmus ligris— Reeve, 1863, fig 4; Hutton, 1873:38. Zizyphimis graimtum—HiMon. 1880:98; Hutton, 1884:359. Cailiostoma gratxatum—?ihhry, 1888:313, pi. 41, fig. 30. Calliostoma ( igris— Pilsbry, 1889:333; Suter, 1897:280; Suter, 1913:148, pi. 40, fig. 6. Calliostoma (Maurea) (igrii— Oliver, 1926:108; Wenz, 1938: 282, fig. 600; Shikama, 1964:106, fig. 185. Venustas (Venustas) tigris — Finlay, 1926:360, 371. Maitrea tigris — Powell, 1937:64, pi. 1, fig. 12; Matsukunia, Okutani & Habe, 1991, pi 17, fig 11. Venustas tigris tigris— Dell. 1950:41, figs. 22, 23, 24. Venustas tigris chathamensis — Dell, 1950:43, figs. 26, 27. New synonym. Maurea tigris tigris — Powell, 1957:88, pl.l, fig. 12; Powell, 1979:60, pi. 10, fig 1. Maurea tigris chathamensis — Powell, 1957:88; Powell, 1979: 61, pi 19, fig 3 NOT Zizyphinus granatum — Reeve, 1863:pl.l, fig. 2 (C. pel- lucidum) Type Data: Trochus tigris: MarUn, 1784, fig. 75, "New Zealand"; Trochus granatum: Chemnitz, 1781, pi. 170, figs. 1654, 1655, "Neuseeland"; Venustas tigris chath- amensis : Holotype MNZ M.2128, Mangare Island, Chat- ham Islands. Other IVIaterial Examined: Fossil — Boulder at head of largest bend in Wainui Stock Road, Ohope (map ref. W15/638502), B.A. Marshall, 1965 (late Castlecliffian, Late Pleistocene) (1 MNZ); Banks of Ohinekoao Stream, coastal clifl's, Matata, B.A. Marshall, 1969 (late Castle- cliffian, Late Pleistocene) (1 MNZ). Recent — 478 speci- mens in 198 lots MNZ, 13 specimens in 8 lots NZOI. Distribution (figure 9): Mid-Pleistocene (late Castleclif- fian) to Recent, off Three Kings, North, South, Stewart and Chatham Islands, living at 0-211 m on rocky sub- strata. Diet: All guts examined contained thecate hydroids (Cni- daria) and indeterminate tissue. Remarks: This well-known species, the largest known calliostomatid, is characterised by its large size, relatively thin shell, coeloconoid spire, very narrowly conical and distinctively sculptured early teleoconch (figure 108), and by the (usual) colour pattern of yellowish or reddish brown wavy axial bands. Sculptural development in nor- mal specimens (see below) proceeds through a distinctive intermediate stage in which the spiral cords become weakly nodular or smooth and much broader than the interspaces, then reverts to strongly nodular with wider interspaces on the last adult whorl. Mature specimens have a gently sloping shoulder on the last part of the last whorl, a pronounced thickening within the outer lip, and typically a slight abapical descent of the apertural rim. Shells having adult characteristics may be as small as 27.5 mm in height (MNZ M. 84227, Cape Runaway), although most are about twice this size. The species at- tains exceptional size off the Three Kings Islands (height up to 100 mm, figure 3). Most specimens from Parengarenga Harbour (figure 6) and many specimens from off eastern Northland (fig- ure 2), including Whangamata and the Aldermen Islands in the northern Bay of Plenty, have appreciably narrower spire angles than specimens from north of Cape Reinga, from Hauraki Gulf, and from south of Mayor Island, Bay of Plenty (sa = 61-72°, mean 66°, S.D. 3.26, n = 20 as against sa = 68-87°, mean 77°, S.D. 3.91, n = 58). There is complete intergradation between narrowly and broad- ly conical forms, however, both within and between pop- ulations, and there are no other differences between them. Because narrowly conical forms occur only in the warm waters off the north-eastern North Island, it would seem that differences in spire angle are linked in some way to sea temperature (see C. hlacki below). The occurrence of broadly conical specimens from further north, off the Three kings Islands, presumably reflects local cooling due to upwelling (Marshall, 1981). Most specimens seen from off the C^hatham Islands, including the holotype of Venustas tigris chathamensis Dell, 1950, differ from mainland specimens in lacking discrete axial bands and are instead predominantly red- dish or yellowish brown with scattered, irregular white spots and streaks (figure 5). Some Chatham Islands shells (e.g., NZOI Q23), however, have axial bands and are essentially similar to specimens from off the South Island and Stewart Island, which commonly have broader bands than northern shells (figure 7). One specimen from the Three Kings Islands (figure 4) is indistinguishable from typical Chatham Islands shells in colour and in density of colour pattern. Although Dell (1950) considered that Chatham Islands specimens differed from mainland shells in having stronger, more persistent nodules, examination of much additional material reveals that sculpture is vari- able and that the Chatham Islands form cannot be dis- tinguished using this criterion. Accordingly I am unable to justify continued recognition of V . tigris chathamensis as a geographic subspecies. Unless C. tigris intermittently reaches the Chatham Islands as drifting eggs or larvae from the mainland, it may have reached there along the summit of the Chatham Rise during periods of low sea level, presumably during Pleistocene glaciations. Three specimens are known from the north-eastern North Island on which the spiral cords remain strong, widely spaced, and nodular throughout (figures 6, 8). The two from Paua, Parengarenga Harbour (figure 6) are also more deeply pigmented and more densely pat- terned than others living beside them. The third example, from off Cape Karikari (figure 8), differs from all other material examined in that the spiral cords are alternately spotted yellowish brown and white. To some extent the Parengarenga Harbour shells resemble C. pellucidum (Valenciennes) in adult fades, whereas that of the Cape Karikari specimen approaches those of both C. punctu- latum (Martyn) and C. osbornei Powell, w ith all of which they respectively occurred. Although we cannot entirely preclude the possibility, they seem unlikely to be hybrids Page 86 THE NAUTILUS, Vol. 108, No. 4 Bruce A. Marshall, 1995 Page 87 because all have the characteristic and highly stable early teleoconch morphology of C. tigri.s (figure 110). The epithet tigris is from the Latin tigris ("tiger", third declension feminine) and is a noun in apposition to the generic name Calliostoma, hence it is not declin- able. Calliostonia (Maurea) punctulatum (Mart\n, (Figures 10-18, 28, 32, 128) 784) Spengler, 1776: 152, pi. 5, figs. 2a, b. Chemnitz, 1781:26, pi. 161, figs. 1520, 1521. Trochus punctulatus Martvn, 1784, fig. 36; Philippi, 1855, pi. 15, fig. 7; Reeve, 1862, pi 16, figs. 95a-d. Trochus diaphanus Gmelin, 1791:3580 (refers to Spengler, 1776: 152, pi. 5, figs. 2a, b; and C:hemnitz, 1781, pi 161, figs. 1520, 1521); Wood, 1825, pi. 29, fig. 99; Quoy & Gaimard, 1834:254, pi. 64, figs. 1-5; Gray, 1842, pi. 40*, figs. 1, la; Philippi, 1846:8, pi, 2, figs, 5, 6; Fischer, 1873:43, Turbo pi. 10, fig 2. Turbo diaphaiius — Lamarck, 1822:45. Ziziphinu.s punctulatus — Gray 1843:237. Turbo grandineus Valenciennes, 1846, pi. 4, figs. 4, 4ab; Kiener, 1847, Turbo pi. 10, fig. 2. lizyphinuspiinctidatus^Huiion, 1873:39; Hiitton, 1880:98 (in part = C. granti): Hutton, 1882:165, pi 7, fig 11; Hutton, 1884:360. Calliostoma punctulatum — Pilsbry, 1889:334, pi. 65, fig. 75; Suter 1897:280; Suter, 1913:146, pi. 8, fig. 11 (in part = C. granti), Buclcnill, 1924:32, pi. 3, fig. 4 (in part = C. granti). Calliostoma (Mauriclla) punctulatum punctulatum — Oliver, 1926:109. Calliostoma (Mauriella) punctulatum stcwartianum Oliver, 1926:109, pi. 10, fig. 1. Calliostoma (Mauriella) wanganuicum Oliver, 1926:109, pi. 10, fig. 2. INew synonym. Venustas (Mucrinops) punctulata punciulata — Finiay. 1926: 361, 371. Venustas (Mucrinops) punctulata urbanior P'inlay, 1926:361, pi. 18, fig. 27; Cernohorsivy, 1972:243, fig. 17'. Maurea (Mucrinops) punctulata punctulata — Powell, 1937:64, pi. 13, fig. 3 (in part = C. granti)- Calliostoma (Mauriella) punctulatum — Wenz, 1938:282, fig, 601. Maurea (Mucrinops) punctulata urbanior — Powell, 1937:64 Venustas punctulata punctulata — Dell, 1950:46, fig. 19, 20, Venustas punctulata urbanior — Deii, 1950:47. Maurea punctulata punciulata — Powell, 1957:88, pi. 13, fig. 3 (in part = C. granti). Maurea punctulata stewartiana — Powell, 1957:88. Calliostoma (Maurea) punctulata stewartiana — Shikama, 1964, pi. 59, fig. 6. Figure 9. Map of New Zealand region showing distribution of Calliostoma (Maurea) tigris. 200 and 1000 meter contours in- dicated. Maurea (Mauriella) punctulata punctulata — Fleming, 1966: 38. Maurea (Mauriella) punctulata stewartiana — Fiemmg, 1966: 38. Maurea punctulata— PoweW. 1976:84, pi. 20, fig. 3; Powell, 1979:62, pi. 10, fig. 4 (both in part = C. granti). Calliostoma (sensulato) punctulatum — Beu & Maxwell, 1990: 404 Calliostoma (sensu lato) wanganuicum — Beu & Maxwell, 1990: 404. NOT Venustas punctulata urbanior — Powell, 1955:55 (C. granti). NOT Maurea punctulata— DeW, 1956:46 (C. granti). Figures 1-8. Calliostoma (Maurea) tigris (Gmelin, 1791). 1. Off Southwest Island, Three Kings Islands. 22-23 m, MNZ M. 75143 (65.5 X 65.0 mm). 2. Off Poor Knights Islands, 20-30 m, MNZ M. 75150 (56.2 x 45.7 mm). 3. Off Three Kings Islands, craypot, MNZ M. 75191, natural size (96.0 x 85.3 mm). 4. Reef between Great Island and Farmer Rocks, Three Kings Islands, 33 m, MNZ M. 84239 (54.0 x 52.3 mm). 5. Off Southeast Island, Chatham Islands, MNZ M. 118120 (54.0 x 51 7 mm) 6. Paua wharf, Parengarenga Harbour, low tide, possibly hybridized with C. (M.) pellucidum (Valenciennes, 1846), MNZ Ml 18337 (57.0 x 45.0 mm). 7. Off Ruggedy Island, Stewart Island, 37 m, MNZ M. 18996 (700 x 68 0 mm) 8. Off Cape Karikari, 80-120 m, possibly hybridised with C. (M.) punctidatum (Martyn, 1784) or C. (M.) osbornei Powell, 1926, MNZ M 87026 (.36.6 x ;52 8 mm). Page 88 THE NAUTILUS, Vol. 108, No. 4 Figures 10-18. Calliostoma (Maurea) punctulatum (Martyn. 1784). 10. Houhora Heads, low tide, MNZ M.2133 (34.6 x 33.6 mm). II. Paua, Parengarenga Harbour, low tide, MNZ M. 80482 (32.8 x 29.0 mm). 12. Northern Pania Reef, Napier, 20m, MNZ M.86724 (34 0 x 3,5.3 mm). 13, 14. Barretts Reef, Wellington Harbour, MNZ M 45062 (29.0 x ,30.4 mm, and 29.0 x 31.2 mm). 15. Simp.son's Rock, Hauraki Gulf, 15-18 m, MNZ M.89971 (31.0 x 29.0 mm). 16. Timaru breakwater, low tide, MNZ M 5324 (42.4 X 408 mm).17. Cook Strait, 256-274 m, MNZ M.54911 (32.6 x 38.6 nun). 18. Masons Bay, Stewart Island, beach, MNZ M.7200 (51.5 X 47.3 mm). Type Data: Trochufi puncttdatus: Martyn, 1784; fig. 36, "New Zealand"; Truchus diaphanus. Spongier, 1776: pi. 5, figs. 2a, b, "Siidsee"; Turbo grandineus. Lectotype (here selected) and 1 paralectotype MNHN, New Zea- land; Calliostoma {Matiriella) punctulatum stewartian- um. Holotype MNZ M.879, Stewart Island; Calliostoma (Mauriclla) tvanganuicum: Holotype NZGS TM 4999, mouth ot Okehu Stream, Nukumarii Beach, near Wang- Bruce A. Marshall, 1995 Page 89 anui (Castiecliffian, Middle Pleistocene); Venustas (Mu- crinops) punctulata urbanior. Holotype AIM 70824, Foveau.x Strait, 37 m. Other Material Examined: Fossil — 56 Early-Middle Pleistocene (Castlecliffian) specimens in 22 lots AUG, MNZ, NZGS; Recent— 1169 specimens in 268 lots MNZ. Distribution (figure 32): Early Pleistocene to Recent. North, South, and Stewart Islands, living intertidally to 274 m on hard substrata. Diet: The intestinal tracts of most of the 22 specimens examined (10 localities) contained thecate hydroids (Cni- daria), sand, and sometimes soft tissue of unknown origin (cnidarianP). Two of three specimens examined from Cornwallis, Manukau Harbour contained thecate hy- droids, soft tissue, and sand, whereas the intestinal tract of the third animal was packed with calcareous octocoral spicules. Remarks: This-well known, common species is extremely variable in shape, size, thickness, colour, colour pattern, and sculpture (figures 10-18). Recent specimens, how- ever, are readily distinguishable from closely related taxa (see below) by features of the early teleoconch, notably the pronounced whorl angulation at P2, the relatively weak axial costae, and the slow enlargement of S3 (figure 28). Shells range in shape from narrowly to broadly con- ical (height/diameter ratio 0.87-1.13; sa 65-96°, mean 84.7°, SD 5.83, n = 74). Specimens from Parengarenga Harbour are the most narrowly conical, with spire angles ranging from 65° to 72° (figure 11). Specimens taken alive from Cook Strait at 256-274 m (figure 17), the deepest record for the species, are the most broadly con- ical (sa 86-96°). The latter specimens, and some from off East Otago, Stewart Island, and Foveaux Strait, are un- usual in having a callous-filled umbilical depression, the umbilicus being completely invaded by the inner lip in other Recent material. Mature specimens are usualK characterised by contraction and descent of the last part of the last whorl and range from 18 to 51 (est.) mm in height, the smallest adult specimens examined occurring off Cape Maria van Diemen, and the largest at Stewart Island in shallow depths, mainly as beach shells (figure 18). Shell thickness varies by a factor of 2 or more. Thick- er specimens tend to predominate in exposed situations, and most are more darkly pigmented with fewer and coarser spiral cords than shells from deeper water. Colour ranges from dark reddish to pale yellowish brown, the spirals darker, the nodules either predominantly white or roughly alternating brown and white. Specimens from the northern North Island tend to be more darkh pig- mented than material from the southern South Island. Spiral cords are multiplied by repeated intercalation of secondaries and tertiaries at variable stages of growth and range in number from 6 to 25 on the second-to-last whorl of mature specimens There is a strong correlation between the strength of sculpture and the number of spiral cords on later whorls. Coarsely sculptured speci- mens with strong primary and secondary spirals tend to develop fewer secondary spirals, whereas more finely sculptured specimens tend to develop more numerous tertiary spirals that enlarge more rapidly to resemble the adjacent spirals Coarsely sculptured specimens tend to predominate in the northern North Island, finely sculp- tured forms in the southern South Island and in deep- living populations. A notable exception is the occurrence of a pale, thin, finely sculptured form at low tide in Manukau Harbour, presumably in response to some local environmental factor. In most specimens from north of Cape Egmont and north of East Cape, including Ma- nukau Harbour, S3 commences about midway between P3 and P4, whereas in most specimens from the south, especially the southern South Island and Stewart Island, S3 commences close beside P4. Although samples from the southern North Island and northern South Island often comprise one form or the other, both forms fre- quently occur together and completely intergrade in sev- eral large samples (e.g., MNZ M.32569, M.44051, M. 45062, M.54911). Southern forms have been treated as geographic subspecies [stewartiana Oliver, urbanior Finlay) of the coarser nominate northern forms, but in fact there is complete mosaic intergradation in all ex- tremes of shell morphology both within and between populations, and it is quite impossible to define regional subspecies. In the Wanganui basin, C. punctulatum first appears in the Butlers Shell Conglomerate (early Castlecliffian, Early Pleistocene) (AU1047, AUG; GS4l69, NZGS) (Tur- ner & Kamp, 1990). An imperfectly preserved shell (MNZ M. 95387) from an Early Pleistocene (late Nukumaruan) horizon exposed in a cutting on White Rock Road, south of Hautotara Bridge, Martinborough appears to repre- sent this species. A worn, incomplete specimen from a Middle Pliocene (Waipipian) horizon in the Waipara Gorge (GS4946, NZGS) is also similar but cannot be iden- tified with certainty. All specimens from the Butlers Shell Conglomerate and from between it and the younger Kaikokopu Shell Grit (GS4064, 4075. 4134, 4163, 4168, NZGS) (late Castlecliffian, Early Pleistocene), including the holotype of C. wanganuicum, differ from Recent specimens in having stronger axial costae on the early teleoconch and in the earlier appearance and more rapid enlargement of S3 (appearing at shell diameter of 1.9- 3.4 mm, mean 2.6 mm, n = 11 as against 3.1-8.0 mm, mean 4.72 mm, n = 24), which is highly conspicuous on early whorls as a suprasutural cord close beside P4. In this character, these specimens bear some resemblance to the Waipipian-Recent species C. granti (Powell) but differ in being more broadly conical and by having more strongly convex whorls as in Recent specimens of C. punctulatum. S3 appears midway between P3 and P4 in most Recent specimens of C. punctulatum, though as already stated, its origination position may vary from median to submedian in some populations. Specimens from the Kupe Formation (late Castlecliffian, Middle Pleistocene) and overlying formations (GS4041, 4052, 4120, 4121, 4122, 4175, 4186, NZGS) are morphologi- cally intermediate between earlier and Recent forms. Page 90 THE NAUTILUS, Vol. 108, No. 4 Moreover, the fossils are as variable in shape, thickness, nodule size, and strength and luiinber of spiral cords as in Recent material, and there is intergradation between extremes within horizons. Accordingly it seems impos- sible to justify recognition of C. wanganuicutn as a dis- tinct species or chronosubspecies. Three specimens from the Butlers Shell Conglomerate (GS4109, NZGS; AU1047, AUG) are unusual in having the umbilicus wide open instead of fully closed through invasion by the inner lip. One of the two umbilicate specimens from GS4109 has the umbilicus considerably narrower than in the other and is thus intermediate between umbilicate and an- omphalous forms from the same sample, which are oth- erwise indistinguishable. Calliostonia punctulatiim is notable for its absence from the Chatham Islands and the subantarctic islands, including The Snares, and all specimens hitherto so iden- tified from there are C. granti (see below). Of yet greater interest is its absence from the Three Kings Islands, some 60 km north of Cape Maria van Diemen, where there are two superficially similar species with closer relation- ship to C. granti (see below). Its absence from this island group may be due to a locally unfavourable environment, either at present or when sea levels were low enough for the species potentially to have reached them, for example during Pleistocene glaciations. As with other local cal- liostomatids that show pronounced clinal variation, it is likely that C. punctulatuni has a drifting larval stage of short duration. Type Data: Maurea granti: Holotype AIM 70449, Waihi Beach, Hawera (Waipipian, Middle Pliocene); Maurea punctulata ampla. Holotype AIM 70450, Masons Bay, Stewart Island; Vcnustas punctulata multigemmata: Holotype AIM 71 185, off East Otago, 91-128 m. Other Material Examined: Fossil — GS4253, Upper Wai- pipi Shellbed, Waverley Beach, S. Taranaki (Waipipian, Middle Pliocene) (3 NZGS); GS4949, c. 200 m SE of north end of Greenwood's Bridge, left bank, lower Wai- para Gorge, map ref. N34/922867 (f6180) (Waipipian) (1 NZGS); GS5237, 724 m N of Trig Turangatairoa, on main Taihape/Waiouru Road, map ref. T21/429771 (f8503) (probably Waipipian; A.G. Beu, per.comm); GS40I3, Tainui Shellbed, Wanganui (late Castlecliffian, Late Pleistocene) (1 AUG); GS4025, Pinnacle Sand, Wanganui (late Castlecliffian) (1 NZGS). Recent— 508 specimens in 124 lots MNZ, and 4 specimens in 2 lots NZOI. Distribution (figure 33): Middle Pliocene (Waipipian) to Recent. North Island south of East Cape and Cape Eg- mont, and South, Stewart, Snares, Auckland, Campbell, and Chatham islands, living from low-tide level to 220 m on hard substrata. Diet: From gut contents (pers.obs. ) and field observations (M.H.B. O'Neill, pers. comm.), C. granti feeds princi- pally on Cnidaria, including thecate hydroids and anem- ones, as well as ascidians. Calliostonia (Maurea) granti (Powell, 1931) (Figures 19-27, 29, 33, 129) Zizyphinus punrtulatus — Hutton, 1880:98 (in part not Martyn, 1784). Calliostoma puncttilatum — Suter, 1913:146 (in part). Maurea (Mucrinops) granti Powell, 1931:97, pi. 13, figs. 34,35. Maurea (Mucrinops) punctulata punctulata — Powell, 1937:64 (in part) Maurea (Mucrinops) punctulata ampla Powell, 1939:229, pi. 50, fig. 6. New §ynonyni Venustas punctulata ampla — Dell, 1950:47 Venustas punctulata (?) n.subsp. Dell, 1950:47. Venustas punctulata multigemmata Powell, 1952:173, pi. 35, figs 2,3. New synonym Venustas punctulata urhanior — Powell, 1955:55 (not Finlay, 1926). Maurea punctulata — Dell, 1956:46 (not Martyn, 1784). Maurea punctulata punctulata — Powell, 1957:88 (in part). Maurea punctulata ampla — Powell, 1957:88. Maurea punctulata multigemmata — Powell, 1957:88. Maurea (Mauriella) granti — Fleming, 1966:38. Maurea (Mauriella) osbornei — Fleming. 1966:38 (not Powell, 1926). Maurea punctulata — Powell, 1976:84; Powell, 197962 (in part) Maurea nmlligemmata—WmeW. 197681; Powell, 197962, pi 10, fig. 6, pi. 19, fig. 8. Maurea hlackii [sic] — Horikoshi, 1989, pi. 4, fig. 12 (not Powell, 1950). Calliostoma (sensu lato) granti^ncn & Maxwell, 1990:404 Remarks: I am unable to detect any constant differences between Waipipian (Middle Pliocene) specimens of Cal- liostoma granti and the Recent forms named Maurea punctulata ampla and Venustas punctulata multigem- mata. Maurea punctulata ampla was originally sepa- rated from the sympatric(!) "subspecies " M. punctulata urhanior Finlay, 1926 [i.e., C. punctulatiim Martyn, 1784) on the basis of the larger, more elevated shell, that has 9-12 instead of 12 equally developed spiral cords. Venustus punctulata multigemmata was separated from M. punctulata ampla because of its more lightly built shell and finer, more numerous spiral cords. Study of many times the number of specimens available when these taxa were proposed, however, reveals that M. punc- tulata ampla and V. punctulata multigemmata are based on forms of a single polymorphic species (i.e., C. granti) that occurs sympatricalK with C. punctulatum from East Cape southward to Stewart Island. It transpires that neither M. punctulata ampla nor V. punctulata mul- tigemmata can be distinguished from C. punctulatum using the characters cited in the original descriptions. Like C. punctulatum. C. granti has considerable vari- ation in shell size and shape, both within populations and clinally, and in the strongly correlated number and rel- ative sizes of the spiral cords on the last two adult whorls. Beach shells and specimens collected intertidally (figure 22) tend to be thick and darkly pigmented with few, strong, strongly nodular spiral cords, whereas specimens from deeper water (figures 20,23) tend to be thin and Bruce A. Marshall, 1995 Page 91 Figures 19-27. Calhoatuina (Maurca) granti (Powell, 1931). 19. Boil Reel, Napier, 10-20 m, MNZ M. 84224 ( 32.0 x 28.5 mm). 20. Off Wanganui, 82 m, MNZ M. 50181 (21.4 x 22.0 mm). 21. Westhaven Inlet, low tide, MNZ M. 81,531 (38.4 x ,35.1 mm). 22. Kaliurangi Point, north-west Nelson, beach, MNZ M. 23101 (28.0 x 27.4 mm). 23. Off Oamaru, c. 100 m, MNZ M. 102602 (44.6 X 41.4 mm), 24. Off Se>mour Island, Dusky Sound, 24 m, MNZ M. 80476 (36.0 x 34.8 mm). 25. Paterson Inlet, Stewart Island, beach, MNZ M. 19119 (45.6 x 42.0 mm) 26. Point Munning, Chatham Islands, 12 m, MNZ M.111927 (41 0 x ,38.1 mm) 27. Off Auckland Islands, 104 m, NZOI D80 (24 0 x 2.3 8 mm). Page 92 THE NAUTILUS, Vol. 108, No. 4 lightK pigmented (or white), with finer, more finely nod- ular, more numerous spiral cords: the former include the type material of M. puuciulata atupla, the latter V. punctulata rnultigemrnata and C granti. There is com- plete mosaic intergradation between these extremes in the Recent material. Although C. granti and C. punctulatum are similar in gross facies, there are nevertheless marked and con- stant differences in the shape and sculpture of the early teleoconch whorls (figures 28, 29). Calliostoma granti differs from C. punctulatum in the following features: (1) the more rapid development of S3, which enlarges to resemble the adjacent primaries at least a full whorl earlier; (2) the stronger a.xial riblets, which persist over one or two additional whorls; (3) the more weakly convex and more slowly expanding early teleoconch whorls {i.e., narrower early spire angle); and (4) by comparison with C. punctulata from within its range, the early teleoconch is evenly conical in shape instead of coeloconoid, al- though in allopatric C. punctulatum from the far north of its range, the spire is typically evenly but yet more broadly conical. Adult teleoconch whorls are typically more weakly convex than in C. punctulatum, although they tend to become as strongly convex in deep-living populations (figures 20, 23). The sides of the foot are considerably more finely pustulose in C. granti than in C. punctulatum. The holotype of C. granti (Powell, 1931, pi. 13, figs. 34, 35) is more narrowly and evenly conical than any known Recent specimen (sa 67°, instead of 70-87°, mean 78°, n = 31), though all other fossils resemble Recent shells in shape. A coeval specimen (GS4949, NZGS) is substantially larger than the largest known Recent shell (diameter 61 mm, as against 49 mm). All of the fossils are indistinguishable from Recent specimens in sculp- ture. Because I am unable to detect any character or character state, single or combined, that would enable separation of Recent and fossil specimens, they are all interpreted as forms of a single polymorphic species. Should the Recent form prove to be specifically distinct when additional, better-preserved Waipipian material is available, Powell's amplum will be available for the Re- cent form. All specimens seen from the Chatham Islands (figure 26) and beach shells from the Auckland Islands and Campbell Island are particularly thick and heavily sculp- tured. Two specimens from 104 m depth off the Auck- land Islands (NZOI D80, figure 27) are thin and finely sculptured and resemble topotypes of the multigemmata form. Unless the species has a planktonic larval stage of sufficient duration for transportation to these islands from the mainland by ocean currents, it probably had a more or less continuous distribution between them during pe- riods of lowered sea level, presumably during the Plio- cene or Pleistocene. That sea temperature may be a fac- tor restricting its northernmost limit to East Cape is sug- gested by the occurrence of two closely related species (C. jamic.soni new species and C. gihhsorum new species) off the Three Kings Islands where sea temperature is cooler than off adjacent Northland due to local upwelling (see C. tigris). Calliostoma granti resembles C. osbornei Powell, 1926 in the rapid enlargement of S3 relative to SI and S2, but the latter two spirals enlarge more slowly in C. osbornei, and C. granti has a weak but distinct angulation at P2 on the early teleoconch (figures 29, 31). By comparison with specimens of C. osbornei from north of Cape Eg- mont, the spiral cords on the early teleoconch whorls in C. granti are yellowish brown with white nodules instead of being predominantly uniform white, whereas the spi- ral interspaces are pale butt instead of orange or yellow- ish to reddish brown Calliostoma granti differs further from C osbornei by having stronger, more numerous axial costae on the early teleoconch whorls, especially on the 4th and early 5th whorls, which are also more strongly convex. Specimens of C. osbornei from off the south-western North Island (figure 41) differ from most northern shells in being as broadly conical as C. granti (sa 75-82°, mean 78°), and in that the spiral cords on the early teleoconch whorls are reddish brown between white nodules. Spec- imens of C. osbornei from these southern populations differ from coexisting C. granti by having more weakly convex 4th and 5th teleoconch whorls and b\ having much more deeply and evenly pigmented spiral cords on the early teleoconch whorls. Calliostoma (Maurea) benthicola (Dell, 1950) (Figures 33, 34) Venmtas benthicola Dell, 1950:47, fig 21. Maurea benthicola— PoweW, 1957:88; Powell, 1979:62. Type Daia: Holotype MNZ M.4728, 41°21'S, 175°00'E, Mernoo Bank, western Chatham Rise, 95 m. Other Material Examined: 22 specimens in 5 lots MNZ. Distribution (figure 33): Endemic to Mernoo Bank, west- ern Chatham Rise, 75-129 m, taken alive at 95-129 m on shell substratum. Remarks: Calliostoma benthicola differs from C granti by having weaker nodules, aiul by being white with broad, reddish brown bands on the spiral cords after the fifth teleoconch whorl. It is otherwise extremely similar, particularly to specimens from equivalent depths off Ota- go, and there can be little doubt that the two forms are very closely related. Calliostoma benthicola appears to be endemic to the Mernoo Bank and is absent from hundreds of dredge and trawl stations from the adjacent (Chatham Rise and off Banks Peninsula and Kaikoura. Mernoo Bank rises Irom depths exceeding 450 m, which are considerably greater than the known lower limit for living C. granti (220 m). It seems likely that C. benthicola diverged from C. granti stock that was isolated on the Mernoo Bank (and the Chatham Islands), perhaps following a period of lowered sea level during the Pleistocene when Mernoo Bank was an island and the Chatham Rise was sufficiently Bruce A. Marshall, 1995 Page 93 close to the surface for icebergs to strand on its summit (Culien, 1962). Calliostoma bcnthicola appears to be an example of a species that arose from a small, isolated population that carried only a fraction of the total genetic material of the parent population (founder effect of Mayr, 1963). This explanation probably accounts for the origin of the distinctive buccinid Cominella olsoni (Dell, 1956), which is also endemic to the Mernoo Bank and is prob- ably derived from Cominella nassoides (Reeve, 1846), forms of which occur to the east off Kaikoura and to the west off the Chatham Islands. Calliostoma (Maurea) jamiesoni Marshall, new species (Figures 30, 33, 36) Description: Shell up to 33 mm high, higher than broad, spire gently cyrtoconoid, spire angle 69-78° (mean 73°, n = 20), stout, glossy, anomphalous. Colour of protoconch and 1st 4 teleoconch whorls uniform yellowish to orange brown. Subsequent whorls yellowish to orange brown, spiral cords reddish brown, nodules white or buff white. Protoconch 400 ^m wide, sculptured with fine threads that enclose hexagonal spaces. Teleoconch of up to 8 whorls, last adult whorl contracted at maturity. Spire whorls convex, periphery rounded, base weakly convex. First 0.25 whorl delineated by a growth scar, with 2 axial riblets and fine spiral threads. Subsequent whorls sculp- tured with spiral cords, rounded nodules, and axial rib- lets; axials strong on 1st 3 whorls, weakening on 4th whorl, becoming obsolete on 5th whorl. Spiral cords rounded, narrow, with broad interspaces on 1st 3 whorls, cords broadening and interspaces narrowing on 4th whorl Spiral cords numbering 7 or 8 on penultimate whorl and 8 or 9 on last adult whorl. Pl-4 commencing immedi- ately, P4 partly covered by succeeding whorls, becoming fully exposed on penultimate whorl by descent of last adult whorl. Sl-3 appearing late on 2nd or on 3rd whorl, gradually enlarging to resemble adjacent primaries (S3 absent in 1 specimen). A subsutural spiral appears on penultimate or last whorl in most mature specimens and gradually enlarges to resemble primaries; 1 or more ad- ditional spirals intercalate on last adult whorl. Spiral in- terspaces with collabral growth lines, most interspaces on later spire whorls with up to 4 fine spiral threads. Columella thick. Aperture subcircular. Outer lip thin at rim, thickened within, inner lip spreading onto colu- mella, parietal glaze very thin. Type Data: Holotype MNZ M. 75141 (height 28.5 mm, width 26.0 mm), off Prince s Rocks, Three Kings Is., alive, 15 m, 2 December 1983, scuba, G.S. Hardy and A.L. Stewart. Paratypes (26 MNZ), all from off Three Kings Is.: Off Three Kings Is., alive, craypot, AD. Howell (1); Three Kings Is., alive, 5 m, March 1982, scuba, K. Burch (2); Tasman Bay, Great King I., alive, 9 m, 19 February 1974, scuba, A.N. Baker & J. Moreland (1); South West I., alive, 22-23 m, 2 December 1983, scuba, G.S. Hardy & A.L. Stewart (3); S side South West I., alive, 27 m, 12 February 1986, scuba, G.S. Hardy (1); reef between Great King 1. and Farmer Rocks, dead, 33 m, 17 February 1986, scuba, G.S. Hardy (15); North West Bay, Great King I., alive, 30 m, 14 February 1986, scuba, G.S. Hardy (1); N. face Hinemoa I., 24 m, 11 February 1986, alive, 3-5 m, 12 February 1986, scuba, G.S. Hardy (2). Other Material Examined: 129 specimens from 6 stations off Three Kings Is., MNZ, mostly immature. Distribution (figure 33): Endemic to Three Kings Islands, 5-128 m, taken alive at 5-55 m on rock. Diet: Intestinal tracts contained thecate hydroids (Cni- daria) and much indeterminate organic matter. Remarks: CMlliostonia jamiesoni is superficially similar to C. punctulatum, C. granii, C. benthicola, and broad forms of C. osbornei, but most closely resembles C. granti and C. benthicola in development of the teleoconch sculpture. It differs from all of them, however, in the uniform coloration of the protoconch and early teleo- conch whorls and in the low relief of the spiral cords and nodules on the fourth to sixth teleoconch whorls (figures 28-31). The spire tends to be more strongly cyr- toconoid and the body whorl more contracted at ma- turity. It differs further from C. punctulatum, C. granti, and C. bcnthicola in the early appearance and more rapid enlargement of the secondary spirals, especially S3, and from C. osbornei in the roughly simultaneous ap- pearance and rather even rate of enlargement of S2 and S3 and the more strongly convex whorls. Judging from development of the teleoconch sculp- ture, C. jamiesoni is more closely related to C. granti, C. benthicola, and C. gibbsorum new species (see below) than to the superficially similar species C. punctulatum and C. osbornei. Etymology: After Peter Jamieson (Wellington), who sought and provided much material for this revision, and as an appreciation for his fostering of New Zealand mal- acology in general, both amateur and professional. Calliostoma (Maurea) gibbsorum Marshall, new species (Figures 33, 35, 109, 130) Description: Shell up to 28.5 mm (est.) high, higher than broad, spire angle 53-76°, stout, glossy, anomphalous. Colour of protoconch and 1st teleoconch whorl buff or white. Next 2 whorls buff with yellowish brown spirals and white nodules and axials. Subsequent whorls yellow- ish to orange brown, nodules more lightly pigmented or white, spirals more darkly pigmented. Protoconch 370- 400 ^im wide, sculptured with fine threads that enclose hexagonal spaces. Teleoconch of up to 8.2 whorls. Spire whorls weakly convex or almost flat, periphery rounded or subangulate, base weakly convex. First 0.25 whorl delineated by a growth scar, with 2 axial riblets and fine spiral threads. Subsequent whorls sculptured with spiral cords, rounded nodules, and axial riblets, the axials strong on 1st 4 whorls, weakening and becoming obsolete on 5th whorl. Spiral cords rounded, narrow with broader Page 94 THE NAUTILUS, Vol. 108, No. 4 Bruce A. Marshall, 1995 Page 95 Figure 32. Map of New Zealand region showing distribution of Calliostoma (Maurca) ptinctulatum 200 and 1000 meter contours indicated interspaces on 1st 4 whorls, spirals broadening and in- terspaces narrowing on 5th whorl. Spiral cords number- ing 7 on adult penultimate whorl and 11 on base. PI- 4 commencing immediately, P4 almost or entirely covered by succeeding whorls throughout or (as in holotype) be- coming entirely exposed at suture from as early as 5th whorl. SI -S3 gradually enlarging to resemble adjacent primaries. Additional spirals intercalated on penultimate and last adult whorls of an exceptionally large specimen. Spiral interspaces with collabral growth lines, a few spiral threads in some specimens. Aperture subquadrate to sub- Figure 33. Map of New Zealand region showing distribution of Calliostoma (Maurea) granti ( circle), C. (M.) benthicola (star), and collectively C. (M.) gibbsorum, C. (M.) jamiesoni, and C. (M.) regale (triangle) 200 and 1000 meter contours indicated. circular. Outer lip thin at rim, thickened within, inner lip spreading onto thick columella, parietal glaze thin. Type Data: Holotype MNZ M.35456 (height 16.8 mm, width 12.5 mm), BS 389, between Palmer Rocks and South East I., Three Kings Is., dead, 82 m, 18 Februar\ 1974, r.v. Acheron. Paratvpes (3 MNZ): BS 893 (0639), 33°59.9'S, 171°45.3'E, Middlesex Bank, NW of Three Kings Is., dead. 186-196 m, 31 January 1981, r.v. Tan- Figures 28-31. Early teleoconch whorls. Primary (?) and secondary (S) spiral cords indicated. Figure 28. Calliostorrm (Maurea) puiictulatum (Martyn, 1784). Cook Strait, 256-274 m, MNZ M.54911. Note late appearance and slow enlargement of S3, weak axial sculpture, and broad, strongly convex .5th teleoconch whorl. Figure 29. Calliostoma (Maurea) granti (Powell, 1931). Off Stephens Island, Cook Strait, 183-187 m, MNZ M. 50266. Note early appearance and rapid enlargement of S3 (arrowed), strong axial sculpture, and narrow, weakly convex 5th teleoconch whorl. Figure 30. Calliostoma (Maurea) jamiesoni Marshall, new- species. Off West Island, Three Kings Islands, 37 m, MNZ M. 80656. Note closer similarit> of sculpture to that of C, (M.) granti (29) than to C. (M.) punclulatum (28), S3 arrowed. Figure 31. Calliostoma (Maurea) osbornei Powell, 1926, Off Cape Maria Van Diemen, 38-43 m, MNZ M. 74665 Note late appearance of SI and S2, early appearance and very rapid enlargement of S3 (arrowed), evenly conical outline, and flattened whorls. Scale bars = 1 mm. Page 96 THE NAUTILUS, Vol. 108, No. 4 garoa (1); RS 921, Elingamite wreck, off West Island, Three Kings Is., dead, 37 m, 16 March 1981, suction dredge, K. Tarlton (1); off Three Kings Is., aUve, craypot, AD. Howell (1). Other Material Examined: 60 specimens from 13 stations off Three Kings Islands, MNZ, mostly juveniles. Distribution (figure 33): Off Three Kings Islands, 33-805 m, taken alive at 102 m and probably living as shallow as about 30 m on rugged bryozoan/shell substratum with corals, sponges, and gorgonians. Probably endemic. Diet: Unknown. Remarks: Calliostoma gibbsoriim is superficially similar to C. jamicsoni, C. granti, C. benthicola, and to a lesser degree, C. punctiilatxim, and C. osbornei. It closely re- sembles C. jarniesoni. C. granti, and C. benthicola in development of the spiral teleoconch sculpture, specifi- cally in the more or less simultaneous origin and even rate of development of Sl-3. Compared with the sym- patric species C. jarniesoni, it differs constantly in colour and colour pattern (especially when immature — see de- scriptions); in having stronger, more crisply defined spi- rals and nodules after the third teleoconch whorl; in having narrower, more numerous spiral cords on the base (11 instead of 8 or 9); and in the shape of teleoconch whorls, which expand more slow ly and are more weakly conve.x after the fourth whorl. Compared with the al- lopatric species C. granti and C. benthicola, develop- ment of teleoconch sculpture is entirely more rapid, S2 and S3 appearing and enlarging to resemble the pri- maries a half to a full whorl earlier. Moreover, the nod- ules are more rounded, and the axial sculpture persists for longer and is stronger, especially on the fourth and early fifth whorls. It differs from the allopatric species C. punctulatum in the earlier appearance and more rap- id enlargement of the secondary spirals (especially S3), the stronger, more persistent axial costae, and the more weakly convex whorls. From the sympatric species C. osbornei it differs in numerous details of colour, colour pattern, and sculpture, most obviously in the more or less simultaneous appearance of S2 and S3. For further remarks see C. jarniesoni (above). Etymology: After David and Sharon Gibbs (Auckland) who provided much valuable material and as an appre- ciation for their fostering of amateur and professional malacology in New Zealand. Calliostoma (Maurea) osbornei Powell, 1926 (Figures 31, 37-39, 41, 43, 131) Calliostoma osbornei Powell, 1920 591, pi 102, fig I, 2. Calliostoma (Mauriclla) osbornei — Oliver, I92(j:l 10. Venustas (Murrinops) osbornei — Firilay, 1926;361, .371. Maurea (Mucrinops) osiwrnei — Powell, 19.37:64, pi. 13. fig. 4. yentistas osbornei — Dell, 1950:47. Maurea osbornei— ?ov,e\\, 1957:88, pi. 13, fig. 4; Powell, 1979: 62, pi. 10, fig. 5. Maurea (Mauriclla) osbornei — Fleming, 196():38 Type Data: Holotype AIM 72037, off Cape Barrier, Great Barrier I., c. 49 m, from fish stomach. Other Material Examined: Fossil — GS4253, Upper Wai- pipi Shellbed, Waverley Beach, south Taranaki (Wai- pipian. Middle Pliocene) (1 NZGS). Recetit — 167 spec- imens in 45 lots MNZ. Distribution (figure 43): Middle Pliocene (Waipipian) to Recent, Three Kings Islands southward to off Kapiti Is- land, southern North Island (34°08.5'S-40°50'S), 0-102 m, living at 12-93 m on the sponge Ancorina alata Dendy, 1924 on rocky ground. Diet: Ancorina alata Dendy, 1924 (Porifera : Stelletti- dae). Remarks: Calliostoma osbornei bears a strong superficial resemblance to C. punctulatum and C. granti. From C. punctulatum it differs in numerous details, most notably the rapid enlargement of S3 and the relatively slow en- largement of SI and S2 (figure 31). Other differences include the more weakly convex whorls, stronger nodules on the early teleoconch, and the lack of a prominent angulation at P2 on the third-fifth teleoconch whorls. Among differences in colour and colour pattern, the most notable are on the first four whorls, which are typically orange with white spiral cords and nodules instead of being predominantly white with reddish or yellowish brown spiral cords and white or alternately spotted nod- ules. The only known exceptions are specimens from oft New Plymouth (figure 41), Wanganui and Kapiti Island at the extreme southern limit of its range, which resemble C. punctulatum in colour pattern. Most specimens of C. osbornei are more narrowly conical than C. punctula- tum. though some shells may be as broadly conical, es- pecially those from the extreme south of its range (sa in southern material 50-82°, mean 66°, S.D. 7.29, n = 52, as against 65-96°, mean 85°, S.D. 5.83, n = 64). Despite the similarity in shape, broad specimens of C. osbornei are readily separable from C. punctulatum by the char- acteristic teleoconch sculpture. The Middle Pliocene (Waipipian) specimen lacks the early spire whorls but is otherwise well preserved and indistinguishable from Re- cent material. Judging from its present restricted north- ern distribution (figure 43) and rarity south of Cape Egmont, the lack of fossils from overlying horizons in the Wanganui section may be related to sea tempera- tures, which vverecooler after the Waipipian (Beu, 1966). The present distribution of C. osbornei is not entirely determined by that of its food, the grey sponge, An- chorina alata, which ranges at least as far south as Banks Peninsula (M.H.B. O'Neill, pers. comm). Calliostoma (Maurea) regale Marshall, new species (Figures 33, 40, 111, 132) Description: Shell up to 12.6 mm high, glossy, of mod- erate thickness; spire narrowly and rather evenly conical, 1.50-1.87 X higher than aperture in adults; spire angle 54-63°, anomphalous. Colour of tip of apical fold yel- Bruce A. Marshall, 1995 Page 97 lowish brown, rest of protoconch white Most fresh spec- imens pale yellowish brown between nodules on P2 and P3 on 2nd and 3rd teleoconch whorls, some specimens with addition of a sub- and/or suprasutural band of dull olive that persists onto 4th whorl First 2 \\ horls t\ picalK with a pinkish flush. Fresh juveniles with pale vellowish brown spots on spiral cords on innermost third of base. Shell elsewhere rather uniform pale buff white. Proto- conch 370 /um wide, sculptured with a network of fine threads that enclose hexagonal spaces, terminal vari.x strong, rounded. Teleoconch of up to 7.25 whorls. First few whorls rather strongly convex, angulated at P2 and P3, subsequent whorls very weakly convex. Periphery rounded at maturity, base more or less flat First quarter whorl delineated by a growth scar, with a strong, rounded axial varix and fine spiral threads. Subsequent spire whorls encircled by prominent spiral cords with rounded nod- ules, the spirals multiplying by intercalation of second- aries and tertiaries that enlarge to resemble primaries. Nodules evenly developed on each spiral, occasionally becoming very weak after 5th whorl on spirals other than PI. PI commencing late on 2nd or early on 3rd whorl, gradually enlarging to resemble P2 and P3, which commence immediately after growth scar on 1st whorl. P4 covered by succeeding whorls, becoming fully ex- posed on penultimate whorl by descent of last adult whorl, nodular on last adult whorl, smooth before it SI com- mencing from early on 3rd whorl to start of 6th whorl, S2 commencing from late on 2nd whorl to midway through 3rd whorl, S3 commencing from late on 3rd whorl to midway through 4th whorl In adults a tertiary spiral intercalates on penultimate whorl, where the total of 8 spirals includes emergent P4. Additional tertiaries intercalate on last adult whorl. Axial riblets strong on 1st 3 whorls, weakening on 4th and becoming obsolete on 5th whorl. Interspace of P3 and P4 on 1st 4 whorls with numerous fine, close, crisp, secondary axial riblets that become obsolete on succeeding whorl. Basal spirals mul- tiplying by intercalation of a few secondaries to number about 10 in adults; interspaces about as wide as each spiral or narrower, sculptured with fine collabral growth lines; spirals weak and smooth in immature specimens, becoming stronger and nodular with increasing size, rarely more or less smooth in adults. Outer basal spirals resem- bling spire spirals, inner spirals stronger and more strong- ly nodular. Aperture subquadrate, strongly thickened within and immediately behind apertural rim in adults, especially at base and columella. Type Data: Holotype M.86730 (height 12.5 mm, width 9.30 mm) and paratype MNZ, BS 902 (0648), 34°10.5'S, 172°I1 4'E, off Three Kings Is., dead, 153 m, 1 February 1981, r.v. Tangaroa. Paratypes (14, all from off Three Kings Is): BS 900 (0646), 33°57.0'S, 171°45.4'E, alive, 98-103 m, 31 January 1981, r.v. Tangaroa (3 MNZ); BS 905 (0651), 33°57.4'S, 172°19.4'E, alive, 128-123 m, 1 February 1981, r.v. Tangaroa (1 MNZ); BS 894 (0640), 34°00.9'S, 171°44.7'E, alive, 201-216 m, 31 January 1981, r.v. Tangaroa (1 MNZ); BS 898 (0644), 34°01.2'S, 17r45.8'E, alive, 221-206 m, 31 January 1981, r.v. Tan- garoa (4 MNZ); E 846, 34°07.5'S, 17r57.5'E, dead, 417 m, 16 March 1968, m.v. \'iti (1 NZOI); BS 902 (0648), 34°10 5'S. 172°11.4'E, dead, 1.53 m, 1 February 1981, r.v. Tangaroa (2 MNZ); BS 906 (0652), 34°14.8'S, 172°13.6'E, dead, 173-178 m, 2 February 1981, r.v. Tan- garoa (1 MNZ). Other IHaterial Examined: 189 specimens from 21 sta- tions oH Three Kings Is., 53-805 m, MNZ, mostly im- mature. Distribution (figure 33): Off Three Kings Islands (33°57.0'S-34°22.8'S), 53-805 m, taken alive at 98-221 m on rugged, comminuted bryozoan/shell substratum with sponges, hydroids, gorgonians, corals, etc. Diet: l^nknown Remarks: Calliostoma regale resembles the sympatric species C. osbornei in shape but differs in details of colour and colour pattern and in development of teleoconch sculpture, including appearance of S2 before S3 and slow enlargement of PI on the first two whorls. Among pre- viously described New Zealand Recent calliostomatids, it is rendered highly distinctive by the presence of fine axial riblets between P3 and P4 on the early teleoconch whorls, which facilitates recognition of even very im- mature specimens (figure 109). Calliostoma regale seems to be closely related to C. simplex Schepman, 1908 from the Banda Sea, from which it differs principally in having more markedly convex spire whorls and stronger nodules on the base. Moreover, PI develops later, and P2 and P3 are relatively larger on the early teleoconch whorls. Al- though the protoconch and first teleoconch whorl of the holotype of C. simplex (ZMA) are eroded, enough re- mains to show that PI is present almost immediately after the protoconch and that it is as large as P2 midway through the second whorl. In C. regale, PI commences late on the second or early on the third whorl and does not rival P2 in size until midway through the third or fourth whorl Calliostoma simplex is otherwise similar in the order of appearance of the secondary spirals and in having axial riblets between P3 and P4. Etymology: Royal (Latin). Alluding to the Three Kings Islands. Calliostoma (Maurea) aupourianum Marshall, new species (Figures 42, ,50, 112, 133) Description: Shell up to 8 60 mm high, glossy, of mod- erate thickness; spire narrowly and rather evenly conical, up to 1.65 X higher than aperture; mean spire angle 58- 65°, anomphalous Colour of extreme tip of protoconch yellowish brown, elsewhere white. Subsequent whorls yellowish or pale yellowish brown, spire irregularly ax- ially mottled in a darker shade, each dark band followed by a narrow white band, base in adults irregularly axially mottled in yet darker shades Protoconch 370-380 ^lm Page 98 THE NAUTILUS, Vol. 108, No. 4 Menioo Bank, Clhathaiii Hise, 9.5 m, MNZ M. 23626 29.0 mm). VifHirp :i4: Culliostoma {Maurea) hcnthicola (DeW, 19.56). Mernoo Hank, (..Mat .-x,-, .. or i Pantin 1957), Beu(1969, 1974, 1977), and Beu ct al. (1977). Past northward range ex- tensions of C. blacki also appear to be associated with low temperatures, and the narrowly conical, flat-sided, strongly sculptured, subrecent northern shells are con- sidered to have lived when sea temperatures were cooler than at present. This explanation could account for the occurrence of old-looking shells from submarine canyons off East Otago, notably the Papanui Canyon (NZGS RM4721), which have flatter-sided whorls than any known Recent specimens from the vicinity. The holotype of the Late Pliocene (Nukumaruan) Venustas couperi Vella, 1954 (figure 59) is essentially similar to Recent specimens of C. blacki from off Timaru and East Otago, and it is considered to be conspecific. Thoristclla chathamensifi profunda Dell, 1956 is based on juveniles from the vi- cinity of the type locality. The Late Pliocene (Nuku- maruan) specimen from Oaro recorded by Beu (1979) resembles Recent specimens from the vicinity (off Kai- koura). Calliostonm blacki may be distinguished from C. joveauxanum by the following characteristics: (1) spi- ral interspaces translucent white or pinkish white instead of yellowish brown; (2) spiral cords reddish brown with white nodules after first 1.5 whorls instead of uniform reddish brown or at least unicoloured (including nodules) on first 4 teleoconch whorls; (3) P4 on first 5 teleoconch whorls almost entirely covered by succeeding whorls and alternately spotted reddish brown and white instead of almost entirely exposed and uniform reddish brown; (4) 5th and 6th teleoconch whorls more strongly convex in C. foveauxanuni; and (5) C. blacki attains smaller shell size than C. joveauxanum w itiiin the geographic range of the latter (height up to 50 mm as against 62 mm or more). Unlike C. blacki. shell morphology in C. fov- eauxanuin is extremelv stable. Calliostoma (Maurea) megaloprepes (Tomlin, 1948) (Figures 63, 64) Maurea (Mucrinops) megaloprepes Tomlin, 1948:225, pi. 2, fig. 1. Venustas megaloprepes — Powell, 1955:55. Maurea megaloprepes — Powell, 1957:88; Powell, 1979:63, pi. 19:2, Type Data: Holotype BMNH 1951.6.13.38, BANZARE stn 80, off north-eastern corner of Macquarie Island, 120- 80 m. Other IMalerial Examined: 15 live-taken specimens: 54°24'S, 159°01'E, 79-93 m, 10 February 1965, USNS Eltanin stn 1417 (6 USNM, 3 MNZ); C733, 54°25'S, 159°02'E, 104 m, 25 November 1961, m.v. Viti (2 NZOI); 54°32'S, 159°02'E, 86-101 m, 10 February 1966, USNS Eltanin stn 1418 (1 USNM); D9, 54°52'S, 158°50'E, 113 m, 20 April 1963, m.v. Viti (2 NZOI); C730, 54°55'S. 158°47'E, 110 m, 24 November 1961, m.v. Viti {I NZOI). Distribution (figure 63): Off Macquarie Island, living at 79-113 m on bryozoan/shell substrata. Endemic. Diet: Gut contents examined comprised mostly forami- niferal sand with fragments of thecate hydroids (Cni- daria) and indeterminate organic matter. Remarks: This species is rendered highly distinctive by its rich chestnut-brown shell coloration and by the virtual obsolescence of all of the spiral cords other than (rarely including) PI on the sides and base of the last two adult whorls. Calliostoma megaloprepes is otherwise similar to C. blacki and the two species are evidently closely related. Calliostoma (Maurea) simulans Marshall, 1994 (Figures 63, 65-68, 116, 137) Calliostoma (Maurea) simulans Marshall, 1994: 68, pl.l, figs. 1-3. Type Data: Holotype MNZ M.87450 (height 30.5 mm, width 29.0 mm, 8.3 teleoconch whorls) and 33 paratvpes MNZ: BS 558, 43°30'S 173°31.3'E, head of Pegasus Can- yon, NE of Banks Peninsula, alive, 446 m, 27 September 1976, r.v, Acheron. Paratypes (158 MNZ): BS 786, 43°25'S, 173°26'E, wall of Pegasus Canyon, alive, 329-183 m, 21 February 1979, r.v. Acheron (35); BS 785, 43°25'S, 173°26'E, Pegasus Canvon, alive, 485- 476 m, 21 Feb- ruary 1979, r.v. Acheron (59); BS 784, 43°29.5'S, 173°30.5'E, Pegasus Canyon, alive, 402-338 m, 21 Feb- ruary 1979, r.v. Acheron (12); BS 783, 43°31'S, 173°30.5'E, Pegasus Canyon, alive, 256-293 m, 21 February 1979, r.v. Acheron (52). Other Material Examined: 3 specimens in 3 lots MNZ, 28 specimens in 13 lots NZOI, 3 specimens in 1 lot USNM. Page 104 THE NAUTILUS, Vol. 108, No. 4 Bruce A. Marshall, 1995 Page 105 Distribution (figure 63): Recent, Challenger Plateau, off New Plymouth, Cxxik Strait to SE of Banks Peninsula, Chatham Rise, and off Bounty and Campbell islands, 183-1006 m, taken alive at 256-410 m from soft substrata with shells. Diet: Intestinal tracts of all specimens examined con- tained fragmented chitinous polychaete tubes (Seden- taria, probably Chaetopteridae) and indeterminate or- ganic matter. Remarks: Calliostoma simulans is characterised by a rather large shell with strong spiral cords on the spire and convex, evenly expanding whorls. Shell characters are rather stable throughout the geographic and bathy- metric range, though specimens from the west coast (fig- ure 67) and from Cook Strait southward to Banks Pen- insula, including the type material (figure 65), are more lightly built and attain smaller size than specimens from elsewhere (figures 66, 68). Occurring throughout most of the range of the highly polymorphic species C. hiacki, and locally in sympatry, most specimens of C. simulans are strongly differentiated from C. blacki. but some forms of C. blacki from the Chatham Rise and off the south- eastern South Island (figures 54, 57) approach C. simu- lans in convexity of the late teleoconch whorls. Callios- toma simulans differs from all forms of C. blacki in having more strongly convex early teleoconch whorls. It differs further from C. blacki from localities other than the eastern Chatham Rise in that SI becomes as large as PI or (in most specimens) larger, and the nodules on SI become correspondingly as large or larger. By contrast, in C. blacki from other than the eastern Chatham Rise, SI only occasionally becomes as large as PI and the nodules on PI are consistently larger than those on SI before the last adult whorl. Although specimens of C. blacki from the Chatham Rise east of Veryan Bank (fig- ure 53) have PI as finely nodular as in C. simulans, all pass through a stage on the early teleoconch when PI is by far the most strongly nodular spiral cord as in other forms of the species. Despite the similarity of some Chat- ham Rise specimens to C. blacki. the two species are strongly differentiated where they occur in strict sym- patry, such as off Banks Peninsula (figures 52, 65) and particularly off the Bounty Islands and off the Auckland Islands and Campbell Island (figures 62, 68). Calliostoma (Maurea) antipodense Marshall, new spe- cies (Figures 63, 69, 117, 138) Figure 63. Map of New Zealand region showing distribution of Calliostoma (Maurea) blacki (solid circle), C. (M.) maiii (open circle), C. (M.) simulans (star, solid star = sympatric with C. (M.) blacki), C. (M.) antipodense (inverted triangle), and C. (M:) megaloprepes (upright triangle) 200 and 1000 meter contours indicated. Description: Shell up to 37 mm high, slightly higher than broad, rather thin, anomphalous, glossy; spire up to 1.77 X as high as aperture, rather evenly conical, spire angle 71-74° Colour orange buff, spiral cords reddish brown between paler nodules. Protoconch 400 ^m wide, sculptured with network of fine threads that enclose hex- agonal spaces, terminal varix strong. Teleoconch of up to 8.5 strongly convex whorls, suture becoming deeply channeled after 4th whorl, periphery rounded, base Figures 51-62. Calliostoma (Maurea) blacki (Powell, 1950). 51. Mernoo Bank, Chatham Rise. 95 m, MNZ M.11837S (37,5 x 40.0 mm). 52. Pegasus Canyon, off Banks Peninsula. 256-293 m, MNZ M. 64651 (41.7 x 40.4 mm), 53. Off Chatham Islands, 315-279 m, MNZ M.90037 (40.3 x 43.4 mm) 54. Off Oamaru, c 90 m, MNZ M. 102603 (44.3 x 42.8 mm) 55, 56. Off Timaru, 293 m, NZOI E424 (55 0 x .55,3 mm, and 51 0 x 49.4 mm). 57. Continental Shelf off Dunedin, MNZ M 7209 (51.0 x .500 mm) 58. Off The Snares, 154-168 m, MNZ M, 92443 (42,3 x 41.0 mm). 59. Holotype of Venustas couperi Velia, 1954, Late Pliocene-Early Pleistocene (Nukumaruan), Makara Stream, Wairarapa, NZGS TM4995 (34 2 x ,35.8 mm) 60. Off Bounty Islands, 155 m, NZOI 1708 (42.8 X ,37.5 mm), 61. Subrecent (probably Late Pleistocene), Mernoo Bank, Chatham Rise, 148-150 m, MNZ M. 61185 (41.0 X 39 0 mm) 62. Off Auckland Islands, 113 m, NZOI D200 (45.4 x 40.0 mm). Page 106 THE NAUTILUS, Vol. 108, No. 4 weakly convex. First 0.2 whorls delineated by a growth scar, with 2 axial costae and 4 spiral threads. Subsequent spire whorls sculptured with spiral cords that multipK by intercalation from 4 (P1-P4) to 9 (P1-P4, S1-S3 + 2 suprasutural spirals); summit of P4 partly covered by succeeding whorls on 1st 4 whorls, after which insertion point progressively descends to fully expose P4 and then 2 additional spirals, the lower of which is either partly covered by succeeding whorls or fully exposed; nodules rounded on PI, P2, P3, SI, and S2, other spirals smooth, spiral interspaces considerably wider than each spiral. PI at first much weaker than P2 and P3, which are strong and similar throughout, gradually enlarging over 1st 5 whorls then weakening and becoming obsolete. Second- ary spirals enlarging to resemble adjacent primaries. SI and S2 commencing on mid 4th to early 5th whorl, S3 commencing mid 4th to mid 5th whorl. Base with 6 spiral cords of similar size, weaker than spire spirals, inner 3 smooth or with rounded nodules, others smooth. Axial costae strong on 1st 3 whorls, weakening and vanishing on 4th whorl. Fine spiral lirae on 1st 3 or 4 whorls, fine collabral growth lines and obscure spiral lines through- out; fine spiral threads in basal interspaces and interca- lating in spiral interspaces on spire on last 1 or 2 adult whorls. .Aperture ovate, outer lip simple, inner lip thick- ened, parietal glaze extremely thin and transparent. Type Data: Holotype MNZ M.80434 (height 31.8 mm, width 29.2 mm, 8.3 teleoconch whorls) and paratype, off Leeward L, Antipodes Is., alive, 18-73 m, 21 November 1972, r.v. Acheron. Paratypes (31): Eltanin stn 2141, 49°40'S, 178°52'E, off Antipodes Is., alive, 86-95 m (2 MNZ, 4 USNM); Eltanin stn 27/1850, 49°40'S, 178°53'E, off Antipodes Is., alive, 103 m, 3 January 1967 (4 MNZ, 20 USNM). Distribution (figure 63): Off Antipodes Islands, taken alive at 18-103 m (minimum limit uncertain) from hard substrata. Probably endemic. Diet: The intestinal tract of the specimen examined con- tained many calcareous octocoral scales (Cnidaria, Prim- noidae), a few thecate hydroid fragments, and sand. Remarks: Compared with C. simulans, to which it is most closely related, C. antipodcnse differs in its deeper shell pigmentation, more strongly convex whorls, deepK' channeled suture, and obsolescence of PI after the fifth teleoconch whorl. Interestingly, C. eminens, the other Antipodes Islands endemic, also has an exceptionally deep suture. Calliostoma (Maurea) maui Marshall, new species (figures 63, 70-72) Description: Shell up to 44 mm high, broader than high, thin, anomphalous or with a shallow umbilical depres- sion, glossy; spire 1.24-1.59 X higher than aperture, even- ly conical, spire angle 73-91° Protocoiuh and earliest teleoconcli vvliorls translucent white. Suhseciuent whorls either pale pink through unilorin translucent while outer shell layer or pale buff or pinkish buff with yellowish or reddish brown spiral cords and predominantly white nodules. Protoconch ca. 400 ^m wide, surface sculpture worn away in all available specimens. Teleoconch of up to 9 strongly and rather evenly convex whorls, suture well impressed, periphery rounded, base weakly convex. Primary sculpture on spire consisting of spiral cords that multiply by intercalation from 4 (P1-P4) to 7 (P1-P4, Sl- S3) or occasionally 6 (S3 absent), up to 3 tertiary spirals occasionally arise on penultimate and last adult whorl, summit of P4 partly covered by succeeding whorls and locally fully exposed. Nodules roundly conical; spiral in- terspaces considerably broader than each spiral, becom- ing finely spirally lirate. P1-P4 commencing immediate- ly. PI at first much weaker than P2 and P3, which are similar throughout, gradually enlarging to resemble them, occasionally weakening and becoming obsolete on last adult whorl. Secondary spirals gradually enlarging to resemble primaries, SI commencing early to late on 4th whorl, S2 on late 3rd to mid 4th whorl, S3 on mid 4th to early 5th whorl. Base covered with spiral lirae, 2 or 3 nodular spiral cords on innermost part, other basal spiral cords almost or entirely obsolete. Axial costae strong on 1st 3 whorls, obsolete thereafter. Aperture ovate, inner lip thickened, parietal lip ver\ thin, outer lip thin and simple. Type Data: Holotype MNZ M. 87449 (height 27.3 mm, width 31.3 mm, 7.1 teleoconch whorls); BS 561, 41°24'S, 174°33'E, Cook Strait, alive, 256-274 m, 29 September 1976, r.v. Acheron. Paratypes (6) : 41°.35'S, 175°00'E, off Palliser Bay, alive, 256-490 m, 19-20 December 1966, USNS Eltanin stn 1848 (1 USNM); C 703, 42°42'S, 173''37.8'E, off Kaikoura, alive, 180-140 m, 19 June 1961, m.v. Viti (1 NZOI); E 759. 42°45'S, 17.3°40'E, off Kai- koura, alive, 195-213 m, 31 March 1967, m.v. Viti (1 NZOI); BS 783, 43°31'S, 173°30.5'E, Pegasus Canyon, NE of Banks Peninsula, alive, 256-293 m, 21 February 1979, r.v. Acheron (1 MNZ); 43°56.4'S, 179°25.1'W, W of Chatham Is., alive, 303-296 m, 15 September 1987, f.v. Chitjo Mart! 5(1 MNZ); off North Canterbury, alive, ca 100 m, J. Sutherland (1 MNZ). Other Material Examined: 10 specimens MNZ: BS 542, 41°08'S, 174°35.5'E, Cook Strait, subfossil, 282-293 m, 12 March 1976, r.v. Acheron (5); BS 561, 4r24'S, 174°33'E, Cook Strait, subfossil, 256-274 m, 29 September 1976, r.v. Acheron (4); VUZ 99, 41°34.3'S, 174°43.3'E, old shell, 274 m, 29 August 1957. Distribution (figure 63): Cook Strait. Chatham Rise, and off north-eastern South Island. li\ing at 140-490 m on rugged substrata. Diet: The intestinal tract of the holotype contained most- ly fragments of thecate hydroids (Cnidaria), with some indeterminate organic matter. Remarks: C.allioslaina ntatii clo.seK resembles lightly built forms of C. simulans and depressed forms of C. blacki from ihe eastern C;hathain Rise in general facies. Bruce A. Marshall, 1995 Page 107 Figure 64. Calliostoma (Maurca) mcgaloprepci: (Tomlin, 1948)^ Off Macquarie Island, 79-93 m, MNZ M. 23623 {32.9 x 34,2 mm). Figures 65-68. Calliostoma (Maurea) simularis Marshall, 1994 65. Holotype, Pegasus Canyon, off Banks Peninsula, 446 m, MNZ M, 87450 (30,5 x 29.0 mm). 66. Central Chatham Rise, 410 m, NZOI G259a (51,0 x 48,5 mm), 67. Challenger Plateau, 337 m, NZOI D242 (27.0 x 27,8 mm). 68. Off Campbell Island, 188 m, NZOI D35 (52 0 x 48,0 mm). Figure 69. Calliostoma (Maurea) antipodense Marshall, new species. Holotype, off Leeward Island, Antipodes Islands, 18-73 m, MNZ M 80434 (31.8 x 29 2 mm). Figures 70-72. Calliostoma (Maurca) maui Marshall, new species. 70. Holotvpe, Cook Strait, 256-274 m, MNZ M. 87449 (27.3 x 31.3 mm), 71. Off North Canterbury, ca.lOO m, MNZ M. 74647 (42.4 x 45.4 mm). 72. Off Kaikoura, 139 m, MNZ M.102606 (45.0 X 48,7 mm). Page 108 THE NAUTILUS, Vol. 108, No. 4 all three having similar development of teleoconch sculp- ture, numerous fine spiral lirae on adult teleoconch whorls, and a tendency toward obsolescence of all but the in- nermost few spiral cords on the base. Calliostoma maui closely resembles C. simulans in having all teleoconch whorls markedly convex and in that PI at no stage be- comes larger or more heavily nodular than the other primary spire spirals as it does in C. blacki. It differs from C. simulans in having considerably finer spiral cords on the spire with finer, mostly more sharply pointed nodules, and in being usually more broadly conical (sa 73°-91°, mean 81°, n = 7; instead of 66°-8r, mean 72°, n = 54). During early stages of the present study I sus- pected that C. maui might be an extreme phenotypic variant of either C. blacki or C. simularis. Subsequent recognition of a specimen living together with C. blacki and C. simulans in a single sample from off Banks Pen- insula (MNZ M.90065, 64651, 64650), however, suggests that another closely related species is involved. Further evidence is suggested by the occurrence in Cook Strait of populations of C. maui that are geographically inter- mediate between populations of C. simulans (figure 63). Living specimens from Cook Strait, including the holo- type (figure 72), are distinctive in having a shallow um- bilical depression. Broken, worn, rust-stained specimens from the floor of the Cook Strait Canyon (MNZ M. 52531, 54912) are evidently Pleistocene fossils that have been reworked by current scour of soft sediment following breaching of a Cook Strait land bridge, probably after the last glaciation (Fleming, 1951, 1963; Pantin, 1957; Marshall, 1978). They have considerably thicker shells than Recent specimens from Cook Strait but are other- wise identical. Etymology: After Maui, the legendary Maori fisherman. Calliostoma (Maurea) selectum (Dillwyn, 1817) (Figures 73-77, 85, 118) Chemnitz, 1795:168, pi. 196, figs 1896-97. Trochus selectus Dillwyn, 1817:801, refers to Chemnitz, 1795: 168, pi. 196, figs. 1896-97; Wood, 1825:140, pi. 29, fig. 101a. Trochus cunninghami Gray, 1834:600, pi. 1. fig 7; Fischer, 1876:119, pi. 39, fig. 1. Ziziphinus cunninghami — Gray, 1843:237; Trochus cuninghami [sic] — Philippi, 1855:281, pi. 41, fig 7 Zizyphinus cunningharnii — Reeve, 1863:pi. I, fig. 6 Zizyphinus cunninghami— HuHon. 1873:38; Huttoii 188098 Zizijphinus hodgei Hutton, 1875:458, pi. 21. New synonym Zizyphinus decarinatus—HuHun. 1884:359 (not Ferry, 1811). Zizyphintis ponderosus Hiilloii, 1885:322. New synonym. Calliostoma selectum— Vihhry, 1889:335, pi. 65, figs. 73, 74 (not pi. 65, fig. 78 = C. pellucidum); Suter, 1897:281 ; Suter, 1913:146, pi. 40, fig. 4. Calliostoma ponderosa — Hutton. 1893:69, pi. 8, fig. 75; Suter, 191.5:2. Calliostoma /lOfigfi— Hutton, 1893:70; Suter, 1915:2. Calliostoma carnicolor Preston, 1907: 140, pi. 8, figs. 6, 7 New synonym Calliostoma (Catliotro}}is) pagoda Oliver, 1926:1 12, pi 10, fig. 4. Calliostoma (Calliotropis) cunningharnii — Oliver, 1926:112. Calliostoma (Calliotropis) hodgei — Oliver, 1926:114. Venustas (Venustas) cunninghami — Finlay, 1926:360, 371. Venustas (Ver^uslas) ponderosa — Finlay, 1926:360, 371. Venustas (Venustas) hodgei — Finlay, 1926:.360, 371. Venustas cunninghami regifica Finla>, 1927:485, pi. 24, figs. 9, 10; Cernohorsky, 1972244. Calliostoma (Calotropis) cunninghamii — Thiele, 1929:49. Maurea (Maurea) cunninghami cunninghami — Powell, 1937: 64, pi. 13, fig. 1. Maurea (Maurea) cunninghami pagoda — Powell, 1937:64. Calliostoma (Calotropis) cuminghami [sic] — Wenz, 1938:282. Maurea (Calotropis) cunninghami cunninghami — Powell, 1946:66, pi. 13, fig. 1. Maurea (Calotropis) cunninghami pagoda — Powell, 1946:66. Venustas cunninghami cunninghami — Dell, 1950:53. Venustas cunninghami pagoda — Dell, 1950:53. Maurea cunninghami cunninghami — Powell, 1957:88, pi. 13, fig. 1. Maurea cunninghami pagoda — Powell, 1957:88. Calliostoma (Maurea) cunninghami — Sliikama & Horikoshi, 1963, pi 8, fig. 15. Maurea (Calotropis) cttnninghami — Fleming, 1966:38. Maurea hodgei — Fleming, 1966:38. Maurea ponderosa — Fleming, 1966:38. Calliostoma (Maurea) selectum — Cernohorsky, 1974:149, fig. 7. Maurea selecta—PoweW, 1979:61, pi. 10, fig. 2; Abbott & Dance, 1982:40; Matsukuma, Okutani & Habe, 1991, pi 17, fig. 10. Maurea pellucida — Abbott & Dance, 198240 (not Valenci- ennes, 1846). Calliostoma (sensu lata) selectum — Beu & Maxwell, 1990:404. NOT Trochus selectus—Phi\ipp\, 1855 (C. pellucidum) NOT Zizyphinus selectus— Reeve, 1863; Hutton, 1873, 1880. 1884 (C. pellucidum). NOT Calliostoma selectum— Pi\shr\ , 1889: pi 65, fig. 78 (C. pellucidum). NOT Calliostoma (Calotropis) selectum— Wenz, 1938 (C. pel- lucidum). Type Data: Trochu-f sclectus: Lectotype (here selected) University Zoological Museum, Copenhagen (Cernohor- sky, 1974, pi. 149, fig. 7), "coasts of New Zealand"; Tro- chus citnninghami: Holotype BMNH 1987047, ex J.E. Gray collection, no locality data, = New Zealand (prob- ably a Wellington west coast beach); Zizyphinus hodgei: Holotype Otago Museum, Dunedin C, 54.50, " Wanganui, in blue clay", probably Landguard Bluff (Late Pleisto- cene, early Haweran); Zizyphinus ponderosa: Holotype Canterbury Museum, Christchurch M.2766, "Wanga- nui", horizon unspecified but probably Butlers Shell Con- glomerate (Middle Pleistocene, early Castlecliffian); Cal- liostoma carnicolor: Repository unknown, "C>lebes(.'*) = New Zealand; Calliostoma (Calliotropis) pagoda: Ho- lotype MNZ M.1602, oft Cape Campbell, c.37 m; Ven- ustas cunninghami regifica: Holotype AIM 70823, off Otago Heads, 55m. Other Material Examined: Fossil — 20 Late Pliocene — Early Pleistocene (Nukuniaruan) specimens in 8 lots AUG, NZC;S; 21 Middle Pleistocene (Castlecliffian) specimens in 18 lots AUG, MNZ, NZGS; Recent— 213 specimens in 86 lots MNZ. Distribution (figure 85): Late Pliocene (Nukumaruan) to Bruce A. Marshall, 1995 Page 109 Recent; North, South, Stewart and Chatham islands, 0-293 m, living at 27-274 m on sandy or muddy substrata with shell or stones. Diet: Gut contents of all specimens examined contained mostly sand with fragments of either agglutinated poK - chaete tubes or thecate hydroids (C^nidaria), together with indeterminate organic matter. One specimen con- tained some crustacean fragments. Remarks: Recent specimens of this v\ell-known species are characterised by large size (diameter up to 70 mm); low, broadly conical spire (sa 73-94°); weakly and evenly convex whorls, angulate or narrowly rounded periphery, weakly to rather strongly convex base, weak axial costae on early teleoconch whorls, similarity of P1-P3 through- out, and the late appearance of the secondary spirals, especially S3. Recent specimens from the South Island east coast from Banks Peninsula southwards tend to have more narrowly conical spires (sa 73-80°, mean 76.8°, SD 1.76, n = 20) than specimens from the North Island (sa 74-94°, mean 84.2°, SD 4.49, n = 40), but there is com- plete intergradation between high- and low-spired forms off the northern South Island (sa 77-90°, mean 82°, SD 4.22, n = 20) and the southern North Island. Two spec- imens from off Jackson Bay, Westland (MNZ M, 17669) are as broadly conical as material from the northern South Island (sa 81° and 85°). North-south clinal grada- tion in spire height is independent of depth. There is considerable variation in the strength of nodules on the spiral cords, and many specimens from Cook Strait tend to be particularly weakly nodular (figure 74). Specimens from the Chatham Islands (figure 77) are indistinguish- able from mainland material (sa 76-81°, mean 79°, SD 1.87, n = 5). The fragmentary holotype of Zizyphinus hodgei re- sembles specimens from Landguard Bluff (early Haw- eran. Middle Pleistocene) in shell morphology and pres- ervation, suggesting that it may have come from that horizon. Moreover, it is indistinguishable from Recent specimens from off the east coast of the South Island. The holotype of Zizyphinus ponderosa is a heavily abraded adult shell from an unspecified horizon in the Wanganui sequence. The shell morphology and state of preservation closely match specimens from the Butlers Shell Conglomerate (early Castlecliffian, Early Pleisto- cene) (figure 76), suggesting that it originated from this horizon. All but one of these specimens (see below) and another from the Nukumaru Brown Sand (GS41 15, NZGS; Nukumaruan, Late Pliocene) differ from all specimens from later horizons in having distinctly cyrtoconoid in- stead of shallovvly coeloconoid spires. They also have considerably thicker shells than most Recent specimens, as do most adult Middle Pleistocene specimens from the overlying Pinnacle Sand (GS4022, NZGS) and Shake- speare Cliff Sand formations (AUG 989), although the few Recent specimens known from Stewart Island (MNZ) are as heavily built. However, Early Pleistocene speci- mens from the Lower Kai Iwi Siltstone (AUG 1028, 1034), which underlies the Pinnacle Sand, are as thin as the most lightly built Recent specimens. Specimens from the Shakespeare Cliff Sand and earlier formations (notably GS4022, NZGS) commonly show much later appearance of some or all of the secondary spirals (especially S3) than any known Recent specimens, often as late as the last or second- to-last adult whorl. Fossils from these early horizons are unlike Recent shells in that the spiral cords frequently become smooth and sometimes very weak after an initial nodular stage, becoming stronger and nodular again on the last or second-to-last whorl or re- maining weak and smooth throughout. A single specimen (GS10964, NZGS) collected by A.G. Beu from the Butlers Shell Conglomerate (Early Pleistocene) differs from oth- ers from this horizon in being considerably thinner and having a more broadly conical spire that is distinctly coeloconoid rather than cyrtoconoid in outline. As in some other specimens from this horizon, the shell is en- tirely devoid of nodules on the remaining three whorls, and the spiral cords are almost obsolete on the last adult whorl. It is thus similar to some Middle Pleistocene shells from the Kupe Formation (GS4041, 4121, NZGS) and the thin, weakly sculptured specimens from the Lower Kai Iwi Siltstone (AU1028, 1034, AUG). It is possible that the more lightly built form lived in situ in a low-energy environment, whereas the heavier shells may have washed in from a shallower, high-energy environment or were perhaps reworked from a slightly older horizon (presum- ably late Nukumaruan, Early Pleistocene). This conten- tion is supported by the fact that the heavier shells are all worn to some extent and many are bored, whereas the lightly built (fragile) form was in fresh condition. As discussed by Fleming (1953:175, 176), many shells from the Butlers Shell Conglomerate have undoubtedly been transported, whereas others are clearly reworked from older beds. All of the fossils otherwise resemble Recent specimens in early teleoconch morphology. There is in- tergradation between specimens with early- and late- developing secondary spirals and between specimens in which the spiral cords are weak or strong and nodular or smooth, both within and between samples from the Shakespeare Cliff Sand and older formations. Accord- ingly, Z. ponderosa and C. selectum are interpreted as forms of a single polymorphic species. Great intraspecific variation during the Pleistocene was probably at least partly a response to oscillating glacial-interglacial tem- peratures and sea levels, with associated variations in sediment type and habitat stability. Interpretation of the fossils is potentially complicated by the likelihood of reworking from older horizons during interglacial ma- rine transgressions. Calliostoma carnicolor Preston, 1907 was based on a specimen reputedly from the Celebes, though from the description and illustration (Preston, 1907, p. 140, pi. 8, figs. 6, 7), it appears to be a perfectly typical, northern specimen of C. selectum. Nothing similar has been re- corded subsequently from the Celebes (or from else- where for that matter), and it almost certainly represents a mislocalised specimen of C. selectum. The original specimen could not be traced despite extensive enquiries. Page 110 THE NAUTILUS, Vol. 108, No. 4 Bruce A. Marshall. 1995 Page 1 1 1 Calliostoma (Maurea) pellucidum (N'aleiK'ieniies, 1846) (Figures 78-84, 86, 119) Trochus pellucidtis Valenciennes, 1846;pl 4, fig 2; Fischer, 1875:70, pi. 15, fig. 2. Trochus torquatus Anton ;>i Pliilippi, 1848:126; Pliilippi, 1855: 261, pi. 38, fig. 13 (not H.C. Lea, 1846). Trochus punciulatus — Kiener, 1847:pl. 15, fig. 2 (not Martvn, 1784). Trochus selcclus — Philippi, 1855:261, pi 1, figs. 6, 7, pi. 38, fig. 12 (not Dillwyn, 1817). Zizyphinus selectiis — Reeve, 1863:pl 1, fig. 1; Hiitton, 1873: 38; Hutton, 1880:98; Huttt)n. 1884:359 (not Dillwyn, 1817). Zizyphinus granatus — Reeve, 1863:pl. 1, fig. 2 (not Gnielin, 1791). Calliostoma selectum — Pilsbry, 1889:pl. 65, fig. 78 only (not Dillwyn, 1817). Calliostoma pellucidum— Suler. 1897:281; Suter, 1913:145, pi 40, fig 3. Calliostoma undulalum Finlay, 1923: 104, pi 10. figs 5a, b. New synonym Calliostoma (Calliotropis) pellucidum pellucidum — Oliver, 1926:111. Calliostoma (Calliotropis) pellucidum spiratum Oliver, 1926: 111. New synonym Calliostoma (Calliotropis) undulatum — Oliver, 1926:111. Venustas (Venustas) pellucida — Finlay, 1926:360, 371. Venustas (Venustas) undulala — Finlay, 1926:360, 371. Maurea (Maurea) pellucida pellucida — Powell, 193764. pi 13, fig. 2. Maurea (Maurea) pellucida spirata — Powell, 1937:64. Calliostoma (Calotropis) selectum — Wenz, 1938, fig. 603 (not Dillwyn, 1817). Venustas pellucida pellucida — Dell, 19.50:49, figs. 7, 8. 12. Venustas pellucida spirata — Dell, 1950.50, fig 3 Maurea pellucida pellucida — Powell, 195788, pi. 13, fig. 2; Powell, 1979:61, pi. 19, fig 4 Maurea pellucida spirata — Powell, 1957:88; Powell, 1979:61, pi. 19, fig. 6. Maurea (Calotropis) pellucida — Fleming, 1966:38. Maurea (Calotropis) undulata — P'leining, 1966:38. Calliostoma (sensu lato) pellucidum — Ben & Maxwell, 1990: 404. Calliostoma (sensu lato) undulalum — Beu & Ma.xwell, 1990: 404. NOT Maurea pellucida— AhhnU & Dance, 1982:40 (C. selec- tum Dillwyn, 1817). Type Data: Trochus pellucidufi: Lectotype (here select- ed) and 6 paralectotypes MNHN; Trochus torquatus: Repository of type material unknown, "locality un- Figure 85. Map of New Zealand region showing distribution of Calliostoma (Maurea) selectum. 200 and 1000 meter con- tours indicated known '. Calliostoma undulalum: Holotype NZGS TM 5001, Castlecliff, Wanganui, formation unknown (prob- ably mid-late Castlecliffian or earK Haweran. Mid-Late Pleistocene); Calliostoma pellucidum spiratum : Holo- type MNZ Ml 1865, Spirits Bay, beach drift. Other Material E.xamined: 6 Pleistocene specimens (from unspecified horizons in the Wanganui coastal sequence) in 5 lots MNZ, NZGS, and 235 Recent specimens in 94 lots MNZ. Figures 73-77. Calliostoma (Maurea) selectum (Dillwyn, 1817), 73. Off Ahipara. 90 m, MNZ M. 72032 (40.0 x 47.8 mm). 74. Off Palliser Bay. 274 m. MNZ M 12969 (42.7 x 51.0 mm). 75. Off Timaru. 110-128 m, MNZ M.7162 (47 0 x 50.0 mm). 76. Early Pleistocene (Castlecliffian). Butler's Shell Conglomerate. Wanganui, NZGS (38.5 x 46 0 mm). 77. Off Chatham Islands, 248-236 m, MNZ M. 90061 (44 7 x .50 2 mm) Figures 78-84. Calliostoma (Maurea) pellucidum (Valenciennes, 1846). 78. Takapuna. Auckland, low tide, MNZ M. 83646 (38.0 x 38.0 mm). 79. Paua, Parengarenga Harbour, low tide, MNZ M. 80.536 (37.7 x 34.0 mm) 80. Between Pandora Bank and Cape Maria Van Diemen, ,33 m. MNZ M 74676 (35.3 x .34 5 mm) 81. Off Rarawa Beach, Northland, 40 m, MNZ M. 86657 (35.0 x 29.5 mm). 82. Reotahi. Whangarei Heads, low tide, MNZ M. 83643 (39.5 x 38,0 mm), 83. Akaroa Harbour entrance. 20 m, MNZ M.751.53 (43.1 x ,39.9 mm). 84. Foveaux Strait oyster beds, 31 m, MNZ M.21719 (48.0 X 47.8 mm). Page 112 THE NAUTILUS, Vol. 108, No. 4 Figure 86. Map of New Zealand region showing distribution of CaUiostonia (Maurea) pellucidum. 200 and 1000 meter con- tours indicated. Distribution (figure 86): Middle Pleistocene to Recent, North, South and Stewart islands, living at 0-187 m on hard substrata. Diet: Intestinal tracts of specimens examined contained thecate hydroids (Cnidaria) together with some indeter- minate organic matter. Remarks: Calliostoma pellucUliim has long been con- fused with the superficially similar species C. waikanae, with which it is locally sympatric throughout its range. Highly distinctive characters of C. peUncidum include reddish brown subsutural and peripheral maculations, strong teleoconch sculpture, consistent appearance of So before SI and S2, and strong P3 on early teleoconch whorls. The spire is shallowly coeloconoid and 1-1.5 times higher than the aperture, and the spire angle ranges from 56° to 86°. The most narrowly conical specimens occur off eastern Northland north of Whangaroa (sa 57-69°, mean 64°, SD 3.35, n = 22), especially in Parengarenga Harbour (figures 79, 81). In specimens from the north- western extremity of Northland (ligure 80), however, including the holotype of C pellucidum spinituni Oli- ver, 1926, the mean spire angle is rather constantly about 75°, and these broadly conical shells are closely similar to material from south of Whangaroa (sa 68-81°, mean 75°, SD 3.45, n = 43) (figures 78, 82-84). The periphery tends to be broadly rounded in southern specimens (fig- ures 83, 84) and more sharply angulate in northern spec- imens (figures 79, 81), but there is complete intergra- dation from north to south. The Pleistocene type material of C. undulatum Finlay, 1923 is indistinguishable from Recent specimens of C pellucidum. Calliostoma (Maurea) waikanae Oliver, 1926 (Figures 87-95, 99, 120, 139) Calliostoma (Calltotropis) waikanae Oliver, 1926:11, pi. 10, fig. 3, Maurea (Maurea) waikanae — Powell, 1937:64. Maurea (Calotropis) waikanae — Powell, 194666 Maurea pellucida morioria Powell, 1946:140, pi. 11, fig. 3; Powell, 1979:61, pi. 19,fig. 5. New synonym. Vcnustas pellucida waikanae~T)e\\, 1950:50, figs. 4,5,9,11. Venufitas pellucida fomteriana Dell, 1950:51, figs. 1,2,10. New synonym Venustas pellucida morioria — Dell, 1950:53. Venustas pellucida haurakiensis Dell, 1950:53, fig. 6. New syn- onym Maurea pellucida forsteriana — Powell, 1957:88; Powell, 1979: 61, pi 10, fig. 3. Maurea pellucida haurakiensis — Powell, 1957:88; Powell, 1979: 61. Maurea pellucida morioria— Powell, 1957;88; Powell, 197961, pi 19, fig. 5 Maurea waikanae— PoweW, 1957:88; Powell, 1979:62, pi. 19, fig. 7. Type Data: Calliostoma (Calltotropis) waikanae: Holo- type MNZ M.1603, Waikanae Beach; Maurea pellucida morioria Holotype AIM 71037, Owenga Beach, Chat- ham Islands; Venustas pellucida forsteriana: Holotype MNZ M.2120, Preservation Inlet, Stewart Island. Ven- ustas pellucida haurakiensis: Holotype NZGS TM 456, Hauraki Gull, 46 m. Other Material Kxamined: Fossil — AU1023, Kaikokopu Shell Grit, coastal section, Wanganui (early Castlecliffian, Middle Pleistocene) (1 AUG); GS4045, 4047, 4049, Kupe Formation, coastal section, Wanganui (late Castlecliffian, Middle Pleistocene) (3 NZGS); GS4 1 87, Shakespeare Cliff Siltstone, coastal section, Wanganui (late (Castlecliffian) (2 NZGS). Recent— 410 specimens in 148 lots MNZ. Distribution (figure 99): Middle Pleistocene (early Cas- tlecliffian) and possibly Middle Pliocene (Waipipian). Recent off North, South, Stewart, Snares and Chatham islands, and Mernoo Bank, 0-549 in, taken alive at 18- 549 m on bryozoan/shell sulistrata. Diet: Intestinal tracts of specimens examined contained fragments of thecate hydroids (Cnidaria) together with indeterminate organic matter Remarks: Calliostoma waikanae occurs throughout the geographic range of C. pellucidum, and the two species Bruce A. Marshall, 1995 Page 113 have been thoroughly confused taxonomically because of their variability and superficial similarity. Calliostoma waikanae was treated as a subspecies of C. pellncidum by Dell (1950), whereas forms of C. waikanae have been consistently treated as "subspecies of C, pellucidum, namely Maurea pcllucida rnorioria Powell, Venustas pellucida forsteriana Dell, and V. pcllucida haurakiensis Dell, all of which are here interpreted as local populations of C waikanae. Although they are similar in shape, size, colour, and colour pattern, C. waikanae is easily distin- guishable from C. pcllucidum by the constant appear- ance of SI and S2 well before S3 instead of vice versa (figures 119, 120). C. waikanae differs further in having finer pustules on the side of the foot and tends to have a more lightU pigmented shell with finer, more finely beaded spiral cords, and the summit of P4 is usually partly or entirely exposed on the spire instead of being almost entirely covered by succeeding whorls. In other words the suture is bordered by P4 in C. waikanae and by So in C. pellucidum. S3 is seldom nodular in C. waikanae, but constantly becomes strongl)' nodular in C. pellucidum from an early stage of growth. Most specimens from off the Three Kings Islands and North Cape (figure 88), the north-western North Island southward to off Kawhia (figure 89), and the north-east- ern North Island as far south as East Cape (figure 87) have narrower and correspondingly higher spires than specimens from further south (sa 68-76°, mean 72°, n = 27, as against 73-92°, mean 82°, n = 76)(figures 90-95). They differ further in that P3 frequently remains con- siderably stronger than PI and P2 onto later whorls, often until the second to last adult whorl, and typically sur- mounts a prominent, narrow suprasutural bulge together with S3 and P4. Such specimens intergrade completeK' with specimens in which the primaries become similar through progressive enlargement of PI and P2 and weak- ening of P3 at an earlier stage of growth and in which the suprasutural bulge is correspondingly weak or absent. Weakness or absence of a suprasutural bulge is a char- acteristic of type and topotypic material of V. p. hau- rakiensis and most material from southern localities, though some specimens from Fiordland (figure 94) have a well-developed and persistent peripheral bulge. Spec- imens from north of Cape Reinga attain larger size (di- ameter up to 52.5 mm) and are more darkly and exten- sively pigmented than specimens from elsewhere off the northern North Island but are otherwise identical. Calliostoma waikanae is exceptionally variable in the area off the west coast of the southern North Island and in Cook Strait. In many specimens from this area, in- cluding the holotype of C. waikanae, the spiral cords and nodules weaken with increasing shell size, and the nodules may become almost obsolete (figure 91). There is, however, complete gradation between weakly and strongly sculptured forms both w ithin and between pop- ulations. Moreover, these specimens show exceptionally wide variation in the development of S3, which may fuse with P4, become as large as P3, remain weak throughout, or entirely fail to appear. Again there is complete mosaic intergradation between the extremes. Pronounced vari- ation in shell morphology in this area is probably at least partly the result of repeated divergence and remixing of populations isolated to the east and west of Cook Strait land bridges that formed during Pleistocene glaciations (Fleming, 1962). This explanation probably also accounts for extreme local variation in direct developers such as buccinids of the genera Penion Fischer and Cominella Gray. The only significant differences that I am able to detect between specimens from the Cook Strait area and the southern South Island (Jorsteriana) is the more general tendency for the southern form to have a rounded pe- riphery at maturity and a slightly narrower spire angle (73-86°, mean 79°, n = 23 as against 75-92°, mean 82°, n = 53). Again, however, there is complete intergrada- tion both within samples and clinally between the north- ern and southern forms. Specimens from the Chatham Islands {rnorioria} (fig- ure 95) are indistinguishable from mainland specimens. Existing collections suggest that C. waikanae has a rather patchy distribution (figure 99) and the differences between specimens from off north-eastern North Island/ Kawhia and the Wanganui/Cook Strait area are certainly greater than between specimens from Cook Strait/Kai- koura area and East Otago, suggesting that the northern population may be more strongly isolated than the other mainland populations are from each other. In view of the tendency for this and other New Zealand calliosto- matids to form morphologically discrete local popula- tions and to show pronounced clinal geographic and bathymetric variation, it seems clear that all of the nom- inate forms should be interpreted as local populations of a single polymorphic species. Middle Pleistocene specimens from the Kaikokopu Shell Grit (AU1023, AUG) and the Kupe Formation, Wang- anui (GS4045, 4047, 4049, NZGS), differ from Recent specimens in being thicker at maturity and in the early appearance and more rapid enlargement of the spiral cord between the suture and PI, which develops to re- semble PI. The two subadult specimens from the Shake- speare Cliff Siltstone (GS4187, NZGS) are very lightly built and have sculpture that becomes obsolete on the sixth w horl and gradually reappears on the eighth w horl. All of these fossils are indistinguishable from Recent spec- imens in early teleoconch morphology, and they are in- terpreted as C. waikanae that is both undergoing genetic drift through time and varying in response to changing sea temperature and bathymetry. A single fragmentary specimen from below the Te Aute Limestone, 2.5km south-east of Takapau, Takapau Surve) District, Hawke Bay (GS2320, NZGS) — age either Nukumaruan or (more likely) Waipipian (Middle Pliocene) (A.G. Beu, pers. comm.) — is even more similar to Recent C. waikanae than the Pleistocene material, but better material is re- quired to ascertain its status. Calliostoma (Maurea) turnerarum (Powell, 1964) (Figures 96, 100, 121) Page 114 THE NAUTILUS, Vol. 108, No. 4 Bruce A. Marshall, 1995 Page 115 Maurea turnerarum Powell, 196411. pi 3, fig. 1-3; Powell, 1979:62, pi. 19, fig. 9; Horikoshi, 1989:pl. 4, figs. 8,9. Type Data: Holotype AIM 71239, off Mayor Island, Bay of Plenty, 366 m. Other IVIaterial Examined: 37 specimens in 19 lots MNZ. Distribution (Bgure 100): Three Kings Islands, off Ninety Mile Beach, and north-eastern North Island as far south as Cape Runaway, 186-805 m, taken alive at 312-529 m on muddy substrata with shells. Diet: The intestinal tracts of specimens examined con- tained numerous fragments of thecate hydroids (Cni- daria) together with some indeterminate organic matter. Remarks: Calliostoma turnerarum resembles C. wai- kanae in gross fades, and the two species are undoubt- edly closely related. Calliostoma turnerarum differs in having a more lightly built shell with finer sculpture on adult whorls and in having pale, wavy axial bands, within which the strongest spiral cords are streaked deep reddish or vellowish brown. Compared with specimens of C. waikanae from within its geographic range, C. turner- arum is more broadly conical, the spire angle ranging from 83° to 95° (mean 87°, n = 5) instead of 68° to 75° (mean 72°, n = 27). Although the early teleoconch sculp- ture is similar in both species, P3 remains similar to P2 in C. turnerarum, whereas P3 becomes markedly stron- ger and more strongly nodular in C. waikanae. Calliostoma (Maurea) penniketi Marshall, new species (Figures 97, 98, 100, 123, 140) Description: Shell up to 57 mm high, about as high as broad at maturity, spire 1.19-1.78 x as high as aperture, spire outline evenly conical or shallowly coeloconoid, last whorl slightly contracting at maturity, spire angle 65- 78°, rather thin but strong, anomphalous throughout. Pro- toconch and 1st teleoconch whorl white. Ground colour on subsequent whorls pale yellowish brown or buff white, rich yellowish brown on last adult whorl of a few spec- imens. Spiral cords spotted and spire whorls typically maculated with yellowish or reddish brown, maculations absent from a few specimens. Spiral cords becoming spotted after 2nd-4th w horl, typically on each alternate spiral on early whorls and on each spiral on later whorls, nodules predominantly white on last 2 adult whorls, most Figure 99. Map of New Zealand region showing distribution of Calliostoma (Maurea) waikanae. 200 and 1000 meter con- tours indicated. deeply pigmented on maculations. Maculations irregu- lar, subquadrate, occupying rather discrete subsutural and suprasutural zones, becoming obsolete on last whorl. Subsutural maculations typically more darkly pigmented than suprasutural ones, almost entirely traversing early whorls, narrowing to occupy adapical half of later whorls, numbering 7-9 on adult penultimate whorl. Suprasutural maculations small, occupying narrow band throughout, numbering 13 or 14 on adult penultimate whorl. Inner lip white, aperture nacreous. Protoconch 370 nm wide. Figures 87-95. Calliostoma (Maurea) waikanae Oliver, 1926. 87. Paratype of Venustas pellucida haurakiensis Dell, 1950, Hauraki Gulf, 46 m, MNZ M.15910 (36.9 x 38.3 mm). 88. Off North Cape, 178-167 m, MNZ M.74670 (51.0 x 51.3 mm). 89. Off Kawhia, 83 m, MNZ M.74656 (36.5 x 40.0 mm). 90. Erie Bay, Tory Channel, Wellington, c. 20 m, MNZ M. 17931 (41.0 x 42.0 mm). 91. Between Mana Island and Cape Jackson, Cook Strait, 256-186 m, MNZ M. 49913 (32.0 x 36.0 mm). 92. Off Oamaru, c. 73 m, MNZ M. 102589 (50.5 x 510 mm). 93. Foveaux Strait oyster beds. 37-46 m, MNZ M. 80567 (48.0 x ,52.5 mm). 94. Bligh Sound, Fiordland, c, 20 m, D. Gibbs collection (42.5 x 41.2 mm). 95. Off Kaingaroa, Chatham Islands, craypots, MNZ M.92395 (36.2 x 39,0 mm). Figure 96. Calliostoma (Maurea) turnerarum (Powell, 1964), North of Three Kings Islands, 348-312 m, MNZ M. 71984 (47.5 X 56.7 mm). Figures 97, 98. Calliostoma (Maurea) penniketi Marshall, new species, 97. Holotype, off Three Kings Islands, c. 55 m, MNZ M.74817 (57.0 x 53.0 mm). 98. Ranfurly Bank, East Cape, 76-71 m, MNZ M. 64782 (27.2 x 27.0 mm). Page 116 THE NAUTILUS, Vol. 108, No. 4 Figure 100. Map of New Zealand region showing distribution of Calliostoma (Maurea) penniheti (circle) and C. (M.) tur- nerarum (star). 200 and 1000 meter contours indicated. sculptured with a network of fine threads that enclose roughly hexagonal spaces, terminal varix strong. Teleo- conch of up to 10 whorls, 1st 2.5-3 whorls convex, sub- sequent whorls more or less flat, last 2 whorls weakly convex. Periphery tightly rounded, becoming rather broadly rounded on last whorl; base almost flat, becoming weakly rounded at maturity First 3 whorls sculptured with spiral cords, axial riblets, and a few crisp spiral threads, axials and spiral threads weakening and vanish- ing over next whorl. P1-P4 commencing immediately, PI thread-like on 1st whorl, gradually enlarging to re- semble P2 and P3, which are similar throughout; P4 peripheral, covered by succeeding whorls, becoming ex- posed by descent of last whorl. SI- S3 commencing on 3rd whorl, enlarging to resemble P1-P3; tertiary spirals commencing on 5th whorl or late on 4th whorl, 1 in each interspace and between IM and suture, enlarging to re- semble secondaries and primaries. Additional finer spi- rals in each interspace on last few whorls. Spirals about as high as broad with considerably wider interspaces and prominent rounded nodules on ist 5 whorls, then flat- tening, becoming smooth, and broadening until sepa- rated by shallow grooves; finally becoming strong, crisply nodular and widely spaced on penultimate or last adult whorl. P3 more strongly nodular than PI and P2 on 3rd and 4th w horls. Adult base with about 18-20 major, spot- ted, nodular spiral cords, additional finer threads in most interspaces. Aperture subquadrate to subcircular. Outer lip thin at rim, thickened within, strongly so at base. Inner lip a spreading glaze. Parietal glaze thin, restricted to a narrow outer spiral band. Type Data: Holotype M.74817 (height 57.0 mm, width 53.0 mm, c. 10 teleoconch whorls) and 2 paratypes MNZ, off Three Kings Is., c.55 m, alive in craypots, coll. A.D. Howell, pres. M. Sanson. Paratypes (9) ; C763, 33°58'S, 172°17.6'E, off Three Kings Is., dead, 73-99 m, 18 Feb- ruary 1962, m.v. Viti (1 NZOI); B93, 34°00'S, 172°30'E, alive, 55-91 m, 22 September 1958, m.v. Viti (1 NZOI); BS392, 34°08.5'S, 172°11'E, dead, 102 m, 18 February 1974, r.v. Acheron (1 MNZ); BS902 (0648), 34°10.5'S, I72°11.4'E, dead, 153 m, 1 February 1981, r.v. Tangaroa (1 MNZ); BS901 (0647), 34°14.1'S, 'l72°09'E, dead, 192- 202 m, 1 February 1981, r.v. Tangaroa (1 MNZ); BS910 (0656), 34°18.8'S, 'l72°18.5'E, dead, 93-88 m, 2 February 1981, r.v. Tangaroa (2 MNZ); BS911 (0657), 34°20.2'S, 172°21.8'E, dead, 121 m, 2 Februarv 1981, r.v. Tangaroa (1 MNZ); BS769 (R127), 37°33.2'S, 'l78°50.3'E, Ranfurly Bank, East Cape, alive, 76-71 m, 25 January 1979, r.v. Tangaroa; BS678 (R36), 37°36.3'S, 178°53.1'E, dead, 74 m, 17 January 1979, r.v. Tangaroa (1 MNZ). Other Material Examined (mostly juveniles): Off Three Kings Islands — 50 specimens in 1 1 lots MNZ; Ranfurly Bank, East Cape — 8 specimens in 4 lots MNZ. Distribution (figure 100): Three King Islands and Ran- furly Bank, East Cape, 55-622 m; taken alive at 55-76 m on rugged, bryozoan/shell substratum with sponges, corals, gorgonians, etc. Diet: The alimentary tract of a paratype from off the Three Kings Islands at 55 m contained fragments of thecate hydroids and indeterminate organic material. Remarks: Calliostoma penniketi is closely related to C. waikanae and C. pelhicidum, which it resembles in size, colour, and colour pattern. It is readily separable from them by the finer, more numerous nodules and weaker P3 on the early spire whorls, and in passing through an intermediate growth stage in which the spiral cords be- come low, broad, and smooth. Specimens from off the Three Kings Islands and East Cape are indistinguishable. Calliostoma penniketi may eventually be discovered at intermediate localities, which are as yet inadequately sampled at appropriate depths on hard substrata. Etymology: Named in honour of the late J. R. (Bob) Pen- niket, a connoisseur of New Zealand Calliostomatidae, and in recognition tor his fostering ot New Zealand nial- act)logy, both amateur and professional. Subgenus Olitkaia Ikcbe, 1942 Otukaia Ikebc, 1942:277. Type species (by original designa- Bruce A. Marshall, 1995 Page 117 Figures 101, 102. Calliostoma (Otukaia) alertae Marshall, new name. 101. Off C^ape Karikari, Northland, 743 m, NZOI F913 (27,0 X 25 7 mm). 102. Off Bounl> Islands, 47.5 ni. NZOI 1704 (31.5 x 25.5 mm). Figure 103. Calliostoma (sensu lato) limatulum Marshall, new species. Holotype, King Bank, north-east of Three Kings Islands, 128 m, MNZ M. 90129 ( 8.05 x 5 50 mm) Figure 104. Calliostoma (sensu lato) onustiim Odhner, 1924 King Bank, north-east of Three Kings Islands, 128 m, MNZ M. 50551 (9.85 X 5 50 mm) Figure 105. Calliostoma (sensu lato) kopua Marshall, new species Holotype, off Cape Campbell, 454-424 m, MNZ M. 59698 (5.35 x 4.95 mm). tion): Calliostoma kiheizicbisu Otuka. 1939; Recent, Ja- pan. Alertalex Dell, 1956:46. Type species (by original designation): Alertalex blacki Dell, 1956 {Calliostoma alertae Marshall, new name); Recent, New Zealand Remarks: Most members of this group are characterised by a moderate to large-sized shell (height up to 36 mm); strong, smooth or weakly beaded P2 and P3; and the extreme weakness of PI. All have a highly nacreous shell due to transiucency of the colourless outer shell layers, and the group (as currently interpreted) occurs world- wide at bathyal depths The radula, jaw, external anat- omy, and development of sculpture on the early teleo- conch are essentially similar to those in Calliostoma (sen- su strict a), and I am unable to justify segregation of the group at genus level. Since I am not able to justify place- ment of Otukaia as a synonym of Calliostoma either, I follow McLean and Andrade (1982) in interpreting it as a subgenus. Calliostoma (Otukaia) alertae Marshall, new name (Figures lUl, 102, 108, 122, 141) Alertalex blacki Dell, 1956:46, figs 61, 120, 260; Dell 196275 (secondary homonym of Venustas blacki Powell, 1950) Otukaia hlack'i—DeW. 1963208; Powell, 197963, pi. 19, fig. 13. Page 118 THE NAUTILUS, Vol. 108, No. 4 Figures 106, 107. Radula of holotype of Calliostoma (sensu lata) kopua Marsfiall, new species. 106. Central and lateral teetfi. 107. Marginal teeth, innermost tooth indicated (x). Scale bars = 10 fjm. Type Data: Holotype MNZ M.9767 and 1 paratype MNZ: 44°04'S, 178°04'W, Chatham Rise, alive, 476 m, 10 Feb- ruary 1954, m.v. Alert. Paratypes (4 MNZ): 43°42'S, 179°55'E, Chatham Rise, alive, 512 m, 24 January 1954, m.v. Alert. Other Material Examined: 17 specimens in 12 lots MNZ, 30 specimens in 20 lots NZOI. 165 I : ^ 175 I Distribution (figure 108): Off Cape Brett southward to off The Snares, Challenger Plateau, Chatham Rise, Boun- ty Plateau, and Pukaki Rise (34°43.5'S-49''02'S), living at 280-861 m on muddy substrata with shells. Diet: Gut contents of the few specimens e.xamined com- prise much indeterminate organic matter together with fragments of thecate hydroids (Cnidaria). Remarks: Since Otukaia is here interpreted as a subgenus of Calliostoma, C. {Otukaia) hlacki (Dell, 1956) becomes a homonym of the prior C. (Matirea) hlacki (Powell, 1950). In the absence of junior synonyms it is here ren- amed C. (O.)alertae. Should Otukaia prove to be worthy of genus-group status, the specific name will revert to the original (ICZN Art 59d). This species is strongly characterised by its iridescent, weakly nodular shell, very strong P2 and P3, the late appearance and extreme weakness of PI, and the strong development of S2, which enlarges to resemble P2. Calliostoma alertae shows slight southward narrowing of the spire angle, which ranges from 61° to 70° (mean 65.7°, u = 7) in material from north of East Cape (figure 101) and 55° to 60° (mean 57.8°, n = 7) in specimens from the Bounty Platform (figure 102). Specimens from these extreme northern and southern populations also have respectively the most lightly and heaviK built shells. Material from geographically intermediate localities are intermediate in spire angle and shell thickness. Calliostoma (sensu lata) onusluni Odhner, 1924 (Figures 104, 108, 124, 142) Figure 108. Map of New Zealand region showing distributions ol (.'(illiiistonia (Otukaia) alcrlac (dot), both C (slat.) lima- litlutu and ('. (.s. lat .) (inttsliuii (star), and C. (.s. lat.) kopiia (open circle). 200 and KHK) meter contours indicated. Bruce A. Marshall, 1995 Page 119 Figures 109-1 17. Earl) whorls of Calliostoma species. Figure 109. Calliostoma (Maurea) gibbsorum Marshall, new species. Off Three Kings Islands, craypot, MNZ M. 74663. Figure 1 10. Calliostoma (Maurea) tigris (Gmelin, 1791 ). Whangaroa Harbour entrance, 20 ni, MNZ M. 41060. Figure 111. Calliustonia (Maurea) regale Marshall, new species. Holotype, off Three Kings Islands, 153 m, MNZ M. 86730, Figure 112. Calliostoma (Maurea) aupouriarmm Marshall, new species. Holotype, off Three Kings Islands, 2.52 m, MNZ M. 86731. Figure 1 13. Calliostoma (Maurea) spectabile (A. Adams, 18.55). Off Auckland islands, 42-44 m, NZOI D71. Figure 114. Calliostoma (Maurea) joveauxarnuii (Dell, 1950), Off Otago Peninsula, 220 m, MNZ M.8939. Figure 115. Calliostoma (Maurea) blacki (Powell, 19.50). Off Otago Peninsula, 476-549 m, MNZ M.8891. Figure 116. Calliostoma (Maurea) simulans Marshall, 1994. Paratype, wall of Pegasus Canyon, off Banks Peninsula, .329-183 m, MNZ M. 64654 Figure 117. Calliostoma (Maurea) antipodense Marshall, new species. Paratype, off Antipodes Islands, 103 m, MNZ M. 80469. Scale bars = 1mm Calliostoma onustum Odhner, 192416, pi 1, fig 4. Calliostoma (Maurea) onustum — Oliver, 1926:108. Fautor onustus—Fin\si\. 1926:360; Powell, 1979:63, pi. 19, fig. 12. Description: Shell up to 6.6.'3 mm high, of moderate thickness, considerably higher than broad; spire narrowly and evenly conical, up to .3.4 x higher than aperture; juveniles with minute umbilical chink, larger specimens anomphalous. Iridescent nacreous through translucent outer shell layer, which is colourless when fresh, dead specimens white, base and P4 pale buff in some speci- mens, protoconch and 1st teleoconch whorl pink in some specimens. Protoconch .320-.'3.30 /um wide, sculptured with fine threads that enclose hexagonal spaces, terminal vari.x strong, rounded. Teleoconch of up to 6.50 whorls, 1st whorl rounded, next 2 angulated at P.3; subsequent whorls rounded over abapical half, weakly concave adapically; periphery narrowly rounded, base more or less flat. First Page 120 THE NAUTILUS, Vol. 108, No. 4 Figures 118-126. Karly whorls of Calliusluniu species. Figure 118. Calliosluma (Maurea) aclccttim (Dillw\n, 1817). Tasman Bank, Golden Bay, 26-24 m, MNZ M. 50.508 Figure 1 19. Calliostoma (S4aureaj pellucidum (Valenciennes, 1846). Off Stephens Island, (^ook Strait, 183-187 m, MNZ M 50269. Figure 120. Calliostoma (Mattrea) waikanae Oliver, 1926 Pelorus Sound mouth, 29 rn, MNZ M. 51450. Figure 121. Calliostoma (Maurca) turncrarum (Powell, 1964) Off Great Island, Three Kings Islands, 440 m, MNZ Ml 18.380 Figure 122. Calliostoma (Ctiikaia) alertae Marshall, new name Wall of Pegasus Canyon, off Banks Peninsula, 1(K)6-512 m, MN'Z M 52781 Figure 123. Calliostoma (Mattrea) pennikvti Marshall, new species. Off Three Kings Islands, 187 m, MNZ M. 34246 Figure 124. Calliostoma (scnsu lato) onustnm Odhner. 1924 Off Three Kings Islands, King Bank, north-east of Three Kings Islands, 128 rn, MNZ M 50.551. Figure I2.'>. Calliostoma (scnsu lato) kopua Marshall, new species. Holot\pe, off Cape Campbell, 454-424 m, MNZ M. 59698 Figure 126. Calliostoma (sensii lato) limalitlum Marshall, new species Holotype, King Bank, north-east of Three Kings Islands, 128 m, MNZ M. 90129. Scale bars = Inim. c. 0.3 whorl demarcated by growth scar, not .sculpturally differentiated from .succeeding half whorl. First 1.5 whorls with fine, crisply defined spiral threads. Spiral cords rounded, sharply shelved adapically, nodular where tra- versing axial costae, multiplying by intercalation from 3 {P2-P4) to 7 (P1-P4, S1-S3); interspaces concave, about as wide as each spiral. Spiral cords enlarging Irom tlie spiral threads. PI commencing later than P2-P4, which are similar throughout, PI as large as them after 2nd v\ liorl. Summit of P4 exposed on spire throughout. Sec- ontlary spirals ra[)itlly enlarging to resemble primaries; SI commencing on late 2rid to late 3rd whorl, S2 on mid Bruce A. Marshall, 1995 Page 121 Figures 127-135. Protoconchs of Calliostoina species Figure 127. Calliostoma (Maurea)tigris (,(.inrliii, 1791). Whangaroa Harbour entrance, 20 m, MNZ M. 41060 Figure 128. Calliostoma (Maurea) punctulalum (Martyn, 1784) Cook Strait, 256-254 m, MNZ M. 54911 Figure 129. Calliostoma (Maurea) granti (Powell, 1931). Off Stephens Island,' Cook Strait, 183-187 m, MNZ M.50266. Figure 130. Calliostoma (Maurea) gibbsonim Marshall, new species. Off Three Kings Islands, 102 m, MNZ M. 34249. Figure 131. Calliostoma (Maurea) osbornei Powell, 1924 Off Wanganui, 33-35 m, MNZ M 118381 Figure 132. Calliostoma (Maurea) regale Marshall, new species. Off Three Kings Islands, 102 ni, MNZ M. 80699 Figure 133. Calliostoma (Maurea) aupourianum Marshall, new species. Middlesex Bank, north-west of Three Kings Islands, 246-291 m, MNZ M 80675 Figure 134. Calliostoma (Maurea) spectabile (A Adams, 1855) Off Auckland Islands, 42 m, NZOI D71 Figure 135. Calliostoma (Maurea) foveauxanum (Dell, 19.50) Off Otago Peninsula, 220 m, MNZ M.8939. Scale bars = 100 ^m. 2nd to early 3rd whorl, S3 on late 1st to mid 2nd whorl. Axial costae commencing late on 1st whorl, gently pro- socline, non-collabral, entirely traversing all subsequent whorls, very strong between P2 and P4, numbering 16- 20 on .5th whorl. Base with 7-9 similar, rounded spiral cords, their outer edges beveled, at first smooth, becom- ing weakly nodular with increasing shell size; interspaces wider than each spiral in most specimens but narrower in a few specimens Collabral growth lines prosocline on spire, opisthocyrt on base. Aperture subquadrate, inner and outer lips thin and simple, parietal glaze very thin. Type Data: Lectotype (Odhner s described and illus- trated specimen here selected) and 1 paralectotype. Zoo- logical Museum, Copenhagen : 10 miles NW of Cape Maria van Diemen, dead, 91 m, 4 January 1915. Other Material Examined: 48 specimens in 12 lots MNZ. Distribution (figure 108): Off Three Kings Islands and off Cape Reinga (33°57.0'S-34°25.0'S), 55-310 m, taken alive at 102-202 m on rugged, bryozoan/shell substratum with sponges, gorgonians, and hydroids. Diet: L'nknown. Remarks: Among previously described ta.xa, Calliosto- ma onustum most closely resembles the southern Aus- tralian species C. retiarium Hedley & May, 1908, from which it differs in having a flattened or concave shoulder on the teleoconch whorls and broader spiral cords. The smaller specimen (paralectotype) recorded by Odhner (1924) is an immature specimen of Thysanodonta wai- rua Marshall, 1988 (Calliostomatidae : Thysanodontin- ae). Marshall (1995) will refer this species to a new genus based on the highly distinctive shell morphology and Page 122 THE NAUTILUS, Vol. 108, No. 4 Figures 136-144. Protoconchs of Ca//tos(oma species. Figure 136. Calliostoma (Maurea) blacki {PoweW, 1950). Off Otago Peninsula, 476-549 m, MNZ M 8891. Figure 137. Calliostoma (Maurea) simtilans Marshall, 1994. Paratype, wall of Pegasus Canyon, off Banks Peninsula, 329-183 m, MNZ M. 64654. Figure 138. Calliostoma (Maurea) antipodense Marshall, new species Paratype, off Antipodes Islands, 103 ni, MNZ M. 80469 Figure 139. Calliostoma (Maurea) waikanae Oliver, 1926. Pelorus Sound mouth, 29 m, MNZ M. 51450. Figure 140. Calliostoma (Maurea) penniheti Marshall, new species. Off Three Kings Islands, 187 m, MNZ M. 34246. Figure 141. Calliostoma (Otukaia) alertae Marshall, new name. Wall of Pegasus Canyon, off Banks Peninsula, 1006-512 m, MNZ M. 52781. Figure 142. Calliostoma (sensu lato) onustum Odhner, 1924. Off Three Kings Islands, 102 m, MNZ M. 34251, Figure 143. Calliostoma (sensu lato) kopua Marshall, new species. Holotype, off Cape Campbell, 454-424 m, MNZ M. 59698 Figure 144. Calliostoma (sensu lato) limatulum Marshall, new species. Off Three Kings Islands, 102 m, parat>pe, MNZ Ml 18382. Scale bars = 100 Mm. degenerate radula (figures 106, 107). C. retiarium and the two new taxa described below are congeneric. Calliostoma (sensu lato) limatulum Mar.sliall, new spe- cies (Figures 103, 108, 126, 144) Description: Shell up to 6.60 niin high, of niotleratc thickness; liigher than broad, considerably so at maturity; some juveniles with minute umbilical chink, adults an- omphalous; spire narrowly and evenly conical, up to 2.9 X higher than aperture. Iridescent nacreous through translucent outer shell layer, which is colourless when fresh, dried specimens becoming white; occasionally with a few scattered irregular patclies of pale buff; protoconcli and 1st 1 or 2 teieoconch whorls sometimes pink. Pro- tocorich 330 ^m wide, sculptured with fine threads that enclose hexagonal spaces, terminal varix strong, rounded. Teieoconch of up to 6.10 whorls, 1st whorl rounded, subsequent whorls flattened adapically, angulate at P3 on 2nd and 3rd whorls, rounded thereafter, periphery narrowly rounded, base more or less flat First c 0 3 w horl demarcated by growth scar, not sculpturally ilifterenti- ated from succeeding half whorl. First 1.5 whorls with fine, crisply defined spiral threads. Spiral cords enlarging from fine threads, rounded, sharply shelved adapically, multiplying b\ intercalation troin 3 (P2-P4) to .5 or oc- casionally 6 (P1-P4, S2±S1), interspaces narrower than each spiral, P3 and P4 becoming most closely spaced. S[)iral cords enlarging from fine threads. PI commencing later than P2 and P3, alter mid 2nd-4th whorl as large as P2; P2 and P3 of similar size on 1st whorl, P3 stronger thereafter; summit of P4 exposed on spire throughout. When present, SI either developing in parallel with PI and rescnibliiig it throughout or commencing as late as mid 4th whorl and enlarging to resemble PI; S2 com- Bruce A. Marshall, 1995 Page 123 mencing on late 1st or on 2nd whorl, as large as PI and P2 after late 3rd whorl. Spirals nodular on early whorls, strongly so on P3 and P4, on 4th or 5th whorl nodules weakening and vanishing from all spirals, v\hith are smooth thereafter. Early spire whorls either entirely tra- versed by strong axial costae, or costae strong between S2 and P4 and weaker adapicalK . A,\ials obsolete after 3rd-5th whorl. Base with 7 or 8 broad, smooth, rounded spiral cords w ith interspaces considerably narrower than each spiral, occasionally with up to 12 finer cords. Col- labral growth lines prosocline on spire, opisthocyrt on base. Aperture subquadrate, inner lip thin. Type Data: Holotype MNZ M. 90129 (height 8.05 mm, diameter 5 50 mm, 7.00 teleoconch whorls), BS 904 (0650), 33°57.0'S, 172°19.0'E, King Bank, NE of Three Kings Is., dead, 128 m, 1 February 1981, r.v. Tangaroa. Paratypes (31 MNZ): BS 898 (0644), 34°01.2'S, 17r44.4'E, dead, 206-211 m, 31 January 1981, r.v. Tangaroa (2); BS 637 (P485), 34°05.0'S, 172°24.6'E, dead, 200 m. 24 June 1978, r.v. Tangaroa (1); BS 392, 34°0S.5'S, 172°11.0'E, alive, 102 m, 18 February 1974, r.v.. Ac/if ron (8); BS 901 (0647), 34°14.1'S, 172°09.0'E, ahve, 192-202 m, 1 February 1980, r.v. Tangaroa (11); BS 911 (0657), 34°20.2'S, 172°21 8'E, alive, 121 m, 2 February 1981, r.v. Tangaroa (6); BS 912 (0658), 34°22.8'S, 172°24.6'E, dead, 121 m, 2 Februarv 1981, r.v. Tangaroa (2); BS 631 (P441), 34°24.0'S, 172°16.8'E, dead, 120 m, 20 June 1978, r.v. Tangaroa (1). Other IMaterial Examined: 72 specimens in 10 lots MNZ. Distribution (figure 108): Off Three Kings Islands and off Cape Reinga, 91-805 m, taken alive at 102-310 m on rugged, bryozoan/shell substratum with sponges, gor- gonians, and hydroids. Diet: I'nknown. Remarks: Calliostorna limatuliau is closely similar to the sympatric species C. onustuni. differing principally in being more broadly conical, in having axial costae confined to the early spire whorls, in having an extra spiral cord on the spire (S3), and in that the spiral cords on the spire become broader and smooth with increasing shell size. The two species have identical protoconch and first teleoconch whorl morphologies, and they are similar in shell structure and shell size relative to the number of whorls. Their geographic and bathymetric distributions are similar, and they commonly occur together in dredge samples. I have not overlooked the possibility that they may be forms of a single sexually dimorphic species, but without knowledge of the reproductive anatomy it is impossible to tell. Calliostorna (sensu lato) kopua Marshall, new species (Figures 105, 106, 107, 125, 143) Description: Shell up to 5,35 mm high, slightly higher than broad, with minute umbilical chink; spire evenly conical, up to 1.78 x higher than aperture. Iridescent nacreous through colourless, translucent outer shell layer. Protoconch 320 jitm wide, sculptured with fine threads that enclose hexagonal spaces, terminal varix strong, rounded. Teleoconch of up to 4.70 whorls, 1st whorl convex, subseciuent whorls flat sided, periphery angulate; base almost flat at first, becoming weakly convex. A growth scar almost immediately after protoconch. First 1.5 whorls with fine, crisply defined spiral threads. Spiral cords on spire angulate in section, sharply beveled adapically, with rounded conical nodules where traversing axials, mul- tiplying by intercalation from 3 (P2-P4) to 6 (P1-P4, SI, S2), summit of P4 exposed on spire; interspaces concave, broader than each spiral, P3 and P4 becoming closer than others with increasing shell size. PI very weak on 1st whorl, by end of 2nd whorl as large as P2 and P3, which are similar throughout, P3 peripheral, summit of P4 exposed on spire throughout. Secondary spirals de- veloping to resemble primaries, SI commencing on early to late 3rd whorl, S2 commencing on 1st half of 2nd whorl; axial costae commencing at end of 1st whorl, strong, prosocline, entirely traversing all spire whorls to P4. Base with 7 or 8 similar, strong, rounded spiral cords, at first smooth then with small rounded nodules, inter- spaces about as wide as each spiral or wider. Surface minutely granulate throughout. Collabral growth lines prosocline on spire, opisthocyrt on base. Aperture subquadrate. Radula (figures 106-107) with formula c.6 + 3 + 1 + 3 + c.6, highly degenerate, all teeth other than innermost 2 pairs of marginals flexible, almost ge- latinous. Central and lateral teeth slender, narrowly ta- pered, extremely thin in section, tips finely serrate In- nermost marginal shortest, stoutest, cutting area angu- late. strong terminal cusp and 3 or 4 secondary cusps on each side. Outer marginals slender, innermost pair lon- gest, cutting area with long series of stout, narrowly ta- pered, curved cusps; outer marginals weakly developed. Type Data: Holotype MNZ M. 59698 (height 5.35 mm, width 4.95 mm, 4.70 teleoconch whorls): BS 668 (R26), 41°55.9'S, I74°43 2'E, SE of Cape Campbell, alive, 454- 424 m, 14 January 1979, r.v. Tangaroa. Paratypes (4 MNZ): BS 969 (R27), 41°55.8'S, 174°40.7'E, SE of Cape Campbell, alive, 434-446 m, 14 January 1979, r.v. Tan- garoa. Other Material Examined: 1 incomplete shell MNZ, 45°45'S, 171°02'E, off East Otago, 600-520 m, r.v. Miin- ida. Distribution (figure 108): Off Cape Campbell and off East Otago, 424-600 m, taken alive at 424-454 m from rugged substrata with shells and corals. Diet: Unknown. Remarks: Compared with the Australian species Cal- liostorna retiarium, which it most closely resembles, C. kopua differs principally in being more strongly nodular, in being larger relative to the number of whorls, and in having a less excert protoconch. The single incomplete shell from off East Otago is 6.55 mm in diameter (esti- mated height 7 mm), suggesting that the type specimens are immature. As in other members of this group the Page 124 THE NAUTILUS, Vol. 108, No. 4 radiila is degenerate through developmental retardation, C. kopua having the most degenerate radula of any known cailiostomatid. Etymology: Deep (Maori). ACKNOWLEDGMENTS I am grateful to the following collectors who provided much of the material: M. Eraser (Wellington), D. Gibbs (Auckland), G.S. Hardy, CD. Paulin, CD. Roberts, and A.L. Stewart (Museum of New Zealand, Wellington), P.R. Jamieson (Wellington), S. O'Shea (Wellington), JR. Penniket (late of Warkworth), J. Raven (Wellington), and D. Watson (Otaki). Thanks to A.G. Beu (Institute of Geological and Nuclear Sciences, Lower Hutt) and J. A. Grant-Mackie and J. Sutherland (University of Auck- land) for the loan of fossil material, to A.G. Beu for advice on stratigraphic problems, and to P. Anderson and W. Main (National Institute of Water and Atmospheric Re- search, Wellington), P. Bouchet (Museum National d'Histoire Naturelle, Paris), W.O. Cernohorsky and B. Haywood (Auckland Institute and Museum), R.G. Mool- enbeek (Zoological Museum, Amsterdam), T. Schiotte (Zoological Museum, Copenhagen), K.M. Way (The Nat- ural History Museum, London), and Sorting Center staff of the National Museum of Natural History (Washington D.C) for the loan of Recent material. Eor comments on the manuscript I thank A.G. Beu and J.H. McLean (Los Angeles County Museum of Natural History). For pho- tography and photographic printing, I am grateful to N. Heke, M. Hall (Museum of New Zealand, Wellington). Thanks also to W. St. George (Institute of Geological and Nuclear Sciences, Lower Hutt) for access to the scanning electron microscope. LITERATURE CITED Abbott, R. T. and S. P. Dance. 1986. Compendium of seashells (Third revised printing). Dutton, New York. 41 Ip. Abbott, S. T. and R. M. Carter. 1994. The sequence archi- tecture of mid-Pleistocene (c. 1. 1-0.4 Ma) cyclothems from New Zealand; facies development during a period of or- bital control on sea-level cyclicity. Pp. .367-394 In: P L. DeBoerand D. G Smith (eds). Orbital forcing and cyclical sequences. 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Dillwyn, L W 1817, A descriptive catalogue of Recent shells arranged according to the Linnaean method; with partic- ular attention to the synonymy. 2. M'Creery, London 1092 pp. Finlay, H. J. 1923 Some remarks on New Zealand Callios- Bruce A. Marshall, 1995 Page 125 tomatidae, with descriptions of new Tertiary species. Transactions of the New Zealand Institute 5-4:99-105 Finlay, H. J. 1926. A further commentary on New Zealand molluscan systematics. Transactions of the New Zealand Institute 57:320-485. [Issued separately 23 December 1926] Finlay, H J. 1927. Additions to the Recent molluscan fauna of New Zealand. -No.2. Transactions of the New Zealand Institute 57:485-487. Fischer, P. 1873-1879. Species general et iconographie des coquilles vivantes . . . (continuation of L. C Kiener, 1834- 1870) 10, 11. Bailliere, Paris Fleming, C. A 1944 Molluscan evidence of Pliocene climatic change in New Zealand. Transactions of the Royal Society of New Zealand 74:207-220. Fleming, C. A 1951. Some post-Miocene changes in New Zealand environments. New Zealand Science Review 9:166- 171, Fleming, C. A. 1953 The geology of Wanganui Subdivision New Zealand Geological Survey Bulletin 52 362 pp Fleming, C A. 1962. New Zealand biogeography: a paleon- tologist s approach. Tuatara 1053-108. Fleming, C. A 1963. A moa-bone from the sea-floor in Cook Strait. Records of the Dominion Museum 4231-233 Fleming, C. A. 1966 Marwick s illustrations of New Zealand shells, with a checklist of New Zealand Cenozoic Mollusca. New Zealand Department of Scientific and Industrial Re- search Bulletin 173. 456 pp. Gmelin, J. F. 1791. Caroli a Linne . Systema naturae per regna tria naturae, secunda classes . . . Editiodecima tertia, aucta, reformata. 1(6). Vermes testacea. Beer, Lipsiae, p.3021-4120 Gray, J. E. 1834 Alphabetical list of the figures of Mollusca. Pp, 595-601 In: E Griffith and E Pidgeon The Animal Kingdom, arranged in conformity with its organisation, by the Baron Cuvier 12. The Mollusca Whittaker, Lon- don. 601 pp. Gray, J. E. 1843. Catalogue of the species of Mollusca Pp.228-265 In: E. Dieffenbach, Travels in New Zealand ... 2. Murray, London. Gray, M, E. 1842. Figures of molluscous animals selected from various authors .1. Longman and Bailliere, Lon- don, p. 1-40. Herrmannsen, A N. 1846-52. Indicisgenerum malacozoorum primordia . , . 1(1);232 pp; supplement et corregenda: 140 pp. Fischer, Cassellis. Horikoshi, M 1989 Sea shells of the world The shapes and patterns designed by nature from the Ninomiya collection. Natural History Museum and Institute, Chiba 86 pp Hutton, F W. 1873. Catalogue of the marine Mollusca of New Zealand Government Printer, Wellington, 116 pp. Hutton, F. W. 1875. Description of three new Tertiary shells, in the Otago Museum. Transactions and Proceedings of the New Zealand Institute 7:458. Hutton, F. W. 1880. Manual of the New Zealand Mollusca Government Printer, Wellington, 224 pp Hutton, F. W 1882. Notes on some branchiate Mollusca. Transactions of the New Zealand Institute 14162-167 Hutton, F W 1884. Revision of the Recent rhipidoglossate and docoglossate Mollusca of New Zealand. Proceedings of the Linnean Society of New South Wales 9:354-378. Hutton, F. W, 1885, Descriptions of new Tertiary shells. Parts 1, 2 Transactions and Proceedings of the New Zealand Institute 17:313-332. Hutton, F. W 1893. The Pliocene Mollusca of New Zealand, In J. Fletcher (ed). The Macleay Memorial Volume, Lin- nean Society of New South Wales:35-92 ICZN Opinion 479, 1957. Validation under the Plenary Powers of specific names for nine species of the Class Gastropoda occurring in the New Zealand area as published by Martyn (T. ) in 1784 in the work entitled "The Universal Con- chologist Opinions and Declarations rendered by the International C^ommission on Zoological Nomenclature 16: 365-416 Ikebe, N. 1942. Trochid Mollusca Calliostoma of Japan, fossil and Recent. Japanese Journal of Geology and Geography 18:249-282. Kiener, L, C. 1834-1879. Species general et iconographie des coquilles vivantes . . . (continue par P Fischer), 1-11. Rousseau and Bailliere, Paris. Lamarck, J. B P A de M. de, 1822. Kistoire Naturelle des animaux sans vertebres .7 Paris 711 pp. Marshall, B. A. 1978. The genus Nei/o in New Zealand (Mol- lusca:Bivalvia), New Zealand Journal of Zoology .5:425- 436. Marshall. B. A. 1979. The Trochidae and Turbinidae of the Kermadec Ridge (Mollusca: Gastropoda). New Zealand Journal of Zoology 6: 521-552. Marshall, B A. 1981. New records of Conidae (Mollusca: Gastropoda) from the New Zealand region. New Zealand Journal of Zoology 8493-501 Marshall, B. A 1988. Thysanodontinae: A new subfamily of the Trochidae (Gastropoda). Journal of Molluscan Studies 54:215-229, Marshall, B. A 1994. A new species of Calliostoma from New Zealand (Mollusca: Gastropoda: Trochoidea). Molluscan Research 15:67-69, Marshall, B A 1995 Calliostoniatidae (Mollusca: Gastropoda: Trochoidea) from New Caledonia, the Loyalty Islands, and the northern Lord Howe Rise Memoires de la Museum National d'Histoire Naturelle, Paris (A), (in press), McLean, J H and V. H. Andrade. 1982. Large archibenthal gastropods of central Chile: collections from an expedition of the R.V. Anton Bruun and the Chilean shrimp fishery. Los Angeles County Museum Contributions in Science 342: 1-20. Martyn, T. 1784. The universal conchologist, exhibiting the figure of every known shell . . . with a new systematic arrangement .1 London Matsukuma, A , T. Okutani and T Habe. 1991 World sea- shells of rarity and beauty. National Science Museum, Tokyo. 206 pp. Mayr, E. 1963. Animal species and evolution. Belknap Press, Cambridge. Odhner, N H 1924 New Zealand Mollusca Papers from Mortensen's Pacific Expedition 1914-16, 19 Videnskabe- lige Meddelelser f ra Dansk Naturhistorisk Forening i Kjob- enhavn 77:1-90. Oliver, W. R B. 1926. New Zealand species of Calliostoma. Proceedings of the Malacological Society of London 17: 107-115. [30th December 1926] Pantin, H M. 1957. Fossiliferous concretions from the shelf south-east of Cape Campbell, New Zealand. New Zealand Journal of Science and Technology B38:781-791. Pantin, H. M. 1963. The significance of a living Chlamys delicatula (MolluscaBivalvia) from Cook Strait New Zea- land Journal of Science 6507-512. Philippi, R A 1846-1855 Trochoideen (Turbo. Trochiis. So- larium, Botella, Delphinula, Phasianella) In: H. C. Kiis- ter and W, Kobelt (eds. ), Systematisches Conchylien-Cab- Page 126 THE NAUTILUS, Vol. 108, No. 4 inet von Martini und Chemnitz ... 2. Bauer and Raspe, Leipzig. 372 pp. Pliilippi, R. A. 1848. Centuria altera testaceorum novoriim Zeitschrift fiir Malakozoologie 5;123-128. Pilsbry, H. A. 1888-1889. Manual of Conchology, structural and systematic. With illustrations of the species. 10, 11. Academy of Natural Sciences, Philadelphia Powell, A. W. B. 1926 Descriptions of six new species and a new genus of gasteropod Mollusca from northern New Zealand. Transactions of the New Zealand institute 56: 591-596. Powell, A. W. B. 1931. Waitotaran faunules of the Wanganui System : and descriptions of new species of Mollusca from the New Zealand Pliocene. Records of the Auckland In- stitute and Museum 1:85-112. Powell, A. W. B. 1937. The shellfish of New Zealand: an illustrated handbook. Unity Press, Auckland 100 pp Powell, A. W B 1939 The Mollusca of Stewart Island Re- cords of the Auckland Institute and Museum 2211-238. Powell, A W B. 1946 New species of New Zealand Mollusca from the South Island, Stewart Island, and Chatham Is- lands Records of the Auckland Institute and Museum 3: 137- 144. Powell, A. W. B. 1950. Mollusca from the continental shelf. Eastern Otago. Records of the Auckland Institute and Mu- seum 4:73-81. Powell, A, W B. 1952. New Zealand molluscan systematics, with descriptions of new species, Part 1 Records of the Auckland Institute and Museum 4169-185. Powell, A. W. B. 1955. Mollusca of the southern islands of New Zealand. Scientific results of the New Zealand sub- antarctic expedition, 1941-45. Cape Expedition Series 15. Department of Scientific and Industrial Research, Wel- lington. 151 pp. Powell, A. W B. 1957 Shells of New Zealand An illustrated handbook. Whitcombe and Tombs, Christchurch. 202 pp. Powell, A. W. B. 1964. New Zealand molluscan systematics with descriptions of new species: Part 4. Record of the Auckland Institute and Museum 6:11-20. Powell, A. W. B. 1976 Shells of New Zealand. An illustrated handbook. WhitcouUs, Christchurch. 154 pp Powell, A. W. B. 1979. New Zealand Mollusca Marine, land and freshwater shells. Collins, Auckland. 500 pp. Preston, H. B. 1907 Descriptions of Cypraca bcrnardinae and Calliostoma carnicolorn.sp. The Nautilus 20:139-140. Quoy, J. R. C. and J. P. Gaimard. 1826, 1827, 1828, 1829. Voyage de decouvertes de I'Astrolabe, execute par ordre du Roi, pendant les annees 1826-29, sous le commande- ment de M.J. Dumont d'Urville. Zoologie, Mollusca 3:1- 366. Tastu, Paris. Reeve, L. 1861-62. Monograph of the genus Truehus. Con- chologia Iconica : or illustrations of the shells of molluscous animals. 13. Reeve, London : 16 captions and plates. Reeve, L. 1863. Monograph of the genus Zizijphmus Con- chologia Iconica: or illustrations of the shells of molluscous animals 14. Reeve, London : 8 captions and plates Riiding, P. F 1798. Museum Boltenianum . . sive catalogus cimeliorum e tribus regnis naturae olim collegerat Joa. Fried Bolten .2. John Christi Trappii, Hamburgi. 199 pp. Shikama, T and M Horikoshi 1963 Selected shells of the world illustrated in colour Hokuryu-kan, Tokyo 154 pp. Shikama, T 1964 Selected shells of the world illustrated in colour. 2 Hokuryu-kan, Tokyo. 212 pp Spengler, L. 1776. 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The Mollusca of Macquarie Island: Gastropods and Bivalves. British Australian and New Zea- land Antarctic Research Expedition Reports (ser. B) 5223- 232. Turner, G. M & P J J Kanip 1990. Palaeomagnetic location of the Jaramillo Subchron and the Matuyama- Brunhes transition in the Castlecliffian stratotype section, Wanga- nui Basin, New Zealand Earth and Planetary Science Let- ters 100:42-50. Valenciennes, A 1846. In: A. du Petil-Thours, Voyage autour du monde sur la Venus, pendant 1836-1839. Atlas de Zoologie (no text). Gide and Cie, Paris. Vella, P. 1954. Tertiary Mollusca from south-east Wairarapa. Transactions of the Royal Society of New Zealand 81:539- 555. Wenz, W. 1938. Gastropoda. Teil LAllgemeinerteil und Pros- obranchia 1(1). In: O. H. Schindewolf (ed), Handbuch der Paliiozoologie 6 240pp. Wood, W. 1825. Index testaceologicus; or a catalogue of shells, British and foreign, arranged according to the Linnean system . . . Wood, London. 188pp. Bruce A. Marshall, 1995 Page 127 APPENDIX CHECKLIST OF RECENT CALLIOSTOMATIDAE FROM THE NEW ZEALAND REGION WITH NEW SYNONYMS CALLIOSTOMATINAE Calliostoma (Matirea) antipodense Marshall, new species. C.(M.) aiipourianum Marshall, new species. C.(M.)bcnlhicola (Dell. 1950). C.(M.} blacki (Powell, 1950) {= couperi Vella, 1954 = profunda Dell, 1956). C.(M.) cmiiiens Marshall, new species C.(M.) fovcauxanum (Dell, 1950). C.{M.) gibbsonim Marshall, new species. C.{M.) granti (Powell, 1931) (= ampla Powell, 1939 = multigemmata Powell, 1952). C.(M.) jamiesoni Marshall, new species. C.fM.j maui Marshall, new species. C.(M.} mcgaloprepcs (Tomliii, 1948). C.{M.) oshornei Powell, 1926. C.{M.) pclbicidum (Valenciennes, 1846) (= nndulatum Finlay, 1923 = spiratum Oliver, 1926). C.(M.) pcnnikcti Marshall, new species. C.(M.) punctulatum (Martyn, 1784). C.(M.) regale Marshall, new species. C.(M.) selectum (Dillwyn, 1817) (= hodgci Hutton, 1875 = ponderoms Hutton, 1885 = caniicolor Preston, 1907). C.(M.) simulans Marshall, 1994 C.(M.) spectabile (A. Adams, 1855). C.(A4.) tigris (Gmelin, 1791) (chathamensis Dell, 1950). C.(M.) turjierarum (Powell, 1964). C.(M.) waikanac Oliver, 1926 (= morioria Powell, 1946 = forsteriana Dell, 1950 = haurakiensis Dell, 1950). C.(Olukaia) alertac Marshall, new name {blacki Dell, 1956). C. (sensti tato) kopua Marshall, new species. C {sensu lata) limatulum Marshall, new species. C. (sensu lata) onustum Odhner, 1924. New genus A, new species (Marshall, 1995). New genus B, new species (Marshall, 1995). THYSANODONTINAE Thijsanudonta aucklandica Marshall, 1988. T. wairua Marshall, 1988. Carinastelc coronata Marshall, 1988. C. jugosa Marshall, 1988. C. krisiellae Marshall, 1988. Five additional calliostomatines are known from the Kermadec Islands, to the north-east of New Zealand (Marshall, 1979): Calliostoma new species A = C. (Trislwhotrocluis) sp, cf. simplex of Marshall, 1979, 6gs. C, D, not Schepman, 1908 (Marshall 1995). C. new species B = C. (T.) sp cf simplex of Marshall, 1979, figs. E, F, not Schepman, 1908 (Marshall, 1995). C. (Trislichotrochus) gendalli Marshall, 1979, C. (T.) species cf tosaense (Kuroda & Habe, 1961). C. (Fautor) consobrinum (Powell, 1958). INSTRUCTIONS TO AUTHORS Tllli NAUTILUS piil)lislu'.s papcM's on all aspects of the biolog\' and svstematics of molhisks. Manuscripts do- scrii)int; oriijinai, nnpublisiied lescarc-li as well as r('\iew articles will he considered. Brief articles, not exceeding 1000 words, will he pnhlisiied as notes and do not re- quire an abstract. Notices of meetings and other items of interest to niaiacologists will appear in a news and notices section. Manuscripts: Each original manuscript and accompa- nving illustrations should he submitted in triplicate. Text must he tv'ped on one side of S!^ X 11 inch white paper, double spaced throughout (including literature cited, ta- bles and figure captions), with at least 1 iiu'h of luargiu on all sides. 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