fo. !? ?. S S Nil''. . J THE NAUTILUS Volume 112, Number 1 July 21, 1998 ISSN 0028-1344 A quarterly devoted to malacology. Mari JUL 3 1 7998 EDITOR-IN-CHIEF Dr. Jose H. Leal The Bailev-Matthews Shell Museum 3075 Sanihel-Captiva Road Sanibel, FL 33957 EDITOR EMERITUS Dr. M. G. Harasewych Department of Invertebrate Zoology National Museum of Natural History Smithsonian Institution Washington, DC 20560 CONSULTING EDITORS Dr. Riidiger Bieler Department of Invertebrates Field Museum of Natural Histor\' Chicago, IL 60605 Dr. Arthur E. Bogan North Carolina State Museum of Natural Sciences Raleigh, NC 27626 Dr. Robert T. Dillon, jr. Department ol Biology Coflege of Charleston Charleston, SC 29424 Dr. William K. Emerson Department of Living Invertebrates The American Museum of Natural Histor\' New York, NY 10024 Dr. Eileen H. Jokinen Institute of Water Resources University of Connecticut Storrs, CT 06269-4018 Mr. Richard I. Johnson Department of Mollusks Museum of Comparative Zoology Harvard Universit)' Cambridge, MA 02138 Dr. James H. McLean Department of Malacology- Los Angeles Counts' Museum of Natural History- 900 Exposition Boulevard Los Angeles, CA 90007 Dr. Arthur S. Merrill % Department of Mollusks Museum of Comparative Zoolog)' Harvard University- Cambridge, MA 02138 Dr. Paula M. Mikkelsen Department of Living Invertebrates The American Museum of Natural Histor)' New York, NY 10024 Dr. Gustav Paulay Marine Laboraton,' Universit)' of Guam Mangilao, Guam 96923 Mr. Richard E. Petit PO. Box 30 North Myrtle Beach, SC 29582 Dr. Edward j. Petucli Department of Geologv- Florida Atlantic University' Boca Raton, FL 33431 Dr. Gary Rosenberg Department of Mollusks The Academy of Natural Sciences 1900 Benjamin Franklin Parkway Philadelphia, PA 19103 Dr. Ruth D. Turner Department of Mollusks Museum of Comparative Zoology HaA'ard University Cambridge, MA 02138 Dr. Geerat J. Vermeij Department of Geology University of California at Davis Davis, C:A 95616 Dr. G. Thomas Watters Acjuatic Ecology Laboratory 1314 Kinnear Road Columbus, OH 43212-1194 Dr. John B. Wise Houston Museum of Natural Science Houston, TX 770.30-1799 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 $15.00 for air mail. All orders shotild be accompanied by pavment and sent to: THE NAUTILUS,' PO. Box 1580, Sanibel, FL 33957, USA. Chonff' of address: Please inform the publisher of your new address at least 6 weeks in advance. AU connnunications should include both old and new addresses (with zip codes) and state the effective date. THE NAUTILUS (ISSN 0028-1344) is published quarterly by The Bailev- Matthews Shell Museum, 3075 Sanibel-Captiva Road, Sanibel, FL 33975. Periochcals postage paid at Sanibel. FL, and additional mailing offices. POSTMASTER: Send address changes to: THE NAUTILUS PO. Box 1580 Sanibel, FL 33957 THE €7 NAUTILUS CONTENTS Volume 112, Number 1 July 21, 1998 ISSN 0028-1344 H. Lee Fairbanks Clarification of the taxonomic status and reproductive anatomy of Pliiloim/nis hatchi Branson, 1968 (Gastropoda: Pulmonata; Philonivcidae) 1 Da\id G. Reid Additions and corrections to the taxonomy of the genus Yiu-Ming Mak PcasicUa Nevill, 1885 (Gastropoda: Littorinidae) 6 Jose H. Leal Donald Richard Moore (February- 16, 1921-October 19, 1997): biographical sketch, zoological taxa, anil bibliography 34 Notices 41 THE NAUTILUS 112{l):l-5, 1998 Page 1 Clarification of the Taxonomic Status and Reproductive Anatomy of PJiilomycus batchi Branson, 1968 (Gastropoda: Pulmonata: Philomycidae) H. Lee Fairbanks Department ot Biology PennsvKania State University 100 Universitv' Drive Monaca, PA 15061-2799 USA hlfl@psu.edu ABSTRACT The mantle pattern and reproducti\'e anatom\' of specimens of Philotni/ais batchi are compared to specimens of P cnrolhii- anus- The results demonstrate that P batchi is distinct from P carolinianus and therefore P. batchi is reestablished as a sep- arate species. In the onginal description of P batchi the re- producti\e system described is that of a designated paratvpe from a locality different from that of the holotvpe. This study demonstrates that the reproductive anatomy of P. batchi from the type locality- is not the same as that in the original species description. The correct reproducti\e anatomv is described, it includes a penial pouch, located in the wall of the penis. Tliis pouch is not recorded in any other United States landsnail. Key Words: Slugs, Philomycidae, Philomtjcus, reproductive anatomy, taxonomy. INTRODUCTION The Philomycidae is a family of terrestrial slugs found primarily in the eastern half of the United States. The species in this family are characterized by a mantle that covers the entire body e.xcept for the tip of the tail and the anterior portion of the head. The length of an adult slug yaries from 25 to 100 mm, depending upon the species. There are three North American genera in this family, Philomijcus. PalUfcra and Mcgapallifcra. In terms of tlie reproductiye anatomy, the species of Philoim/cus are characterized by the presence of a dart sac and dart; the species of Pallifcra are small {approximately .30 mm long) and lack the dart sac and dart; the species oi Medmal than that of the specimens from the type loc;ility'. The data obtained from the immature holotype were consistent with the Philomijcus batchi specimens col- lected from the type locality. That is, the general shape of the penis was similar, and the location of the penial pouch and the presence of the ridge on the internal sur- face near the pro.\im;d end of the penis were the same. When Branson (1968) described Phihiimjcus batchi, he apparently assumed, based upon external appearance, that slug specimens he had collected in McCreary Coun- ty, Kentuck-y- were of the same species as the one a col- league had collected in Clark Ct)untv, Kentuck-y 135 km to the north. He then described a new species using hvo specimens. Based upon the data presented in this studv, it appears that Branson utihzed two different species to describe P. botchi. Therefore, a description of the cor- rect reproductive anatomy for P. batchi is provided. The specimen that Branson used to describe the reproduc- tive system of Philoimictis batchi appears to be an un- described species. Because only one specimen of this species is presentK a\;iilable. its description will appear in a later paper. An unexpected result of this stud\' was the disco\ery of a chamber in the penial wall of the specimens of Phi- lonu/ciis from Kentucky. The gross structure and posi- tion of the chamber is not recorded from any other group of land snails in the United States. Studies are underyvav to ascertain the number of different species of Philomijcus that have this chamber, and to try to de- termine hov\' widely spread these species are geograph- ically The function or functions of this chamber are un- knowii at this time. ACKNOWLEDGMENTS Financial support for the fiekl trips associated with this study was provided bv grants from the Research Devel- opment Grant Fund of Penns\lyania State University. LITERATURE CITED Bosc, L.A.G. IS02. Histoire naturelle des coquilles, contenant leur description, et leurs moeurs. Vol. 1. Paris, 34.3 pp. Branson, B. 1968. Two new slugs (Pulmonata: Philomvcidae: Philomijcus) from KentuckT and N'irginia. The Nautilus 8I(4):I27-I.33. Fairbanks. H. L. 1986. The taxonomic status of Phihvni/ciis togatus (Pulmonata: Philomvcidae): a morphological and electrophoretic comparison with Phih^mijcu.'i cnrohnianus. Malacolopa 27(2):271-2S0. Fairbanks, H. L. 1989. The reproducti\e anatomy and taxo- nomic status of Philonujam ventisttis Hubricht, 1953 and Fhilomi/ciis bisdosiis Branson, 1968 (Pulmonata: Philo- mycidae). The Nautilus 103(l):20-23. Fairbanks, H. L. 1990 Morphological comparisons of the spe- cies of Mi'gapnllifcra (Gastropoda: Philomvcidae). The Nautilus 104(2):71-75. Fairbanks, H. L. 1993. The reproductive anatomy of Philo- im/cus scUatus Hubricht, 1972 and Philomijcus virainiais Hubricht. 1953 (Gastropoda: Philomycidae). The Nautilus 107(1):9-13. Hubricht, L. 1974. A re\iew of some land snails of the eastern United States. Malacological Re\iew 7:3.3-.34. Pilsbry, H.A, 1948. Land Mollusca of North America (North of Mexico). Vol. 11, Part 2:754. THE NAUTILUS 112(l):6-33, 1998 Page 6 Additions and Corrections to the Taxonomy of the genus Peasiella Nevill, 1885 (Gastropoda: Littorinidae) Da\id G. Reid Department of ZooIogN The Natural Histon Museum Lxmdon S\V7 5BD,' UK D.reidCS'nhm.ac.nk Yiu-Ming Mak' Department of Ecology and Biotliversity The University of Hong Kong Hong Kong ABSTRACT Peasiella is a genus of the Littonmdae with nnnute, trochoidal shells, found onlv in the Indo-West Pacific region. A prexious monograph recognized slx Recent species. As a result of new anatomical information (reproductive anatomy, egg capsules, radulae) two of these are dixided into five and two species, respectively, to make a total of 11 species in the genus. Three of these are descnhed as new. Additional notes on distribution, variation and nomenclature are provided on otlier species. Characters of the radula and pallial oxiduct are relatively uni- form throughout the genus, hut penial shapes are often diag- nostic. Shell shape and sculpture show extreme mtraspecific x-ariabilitx' INTRODUCTION Pca.sicllo is one of the more poorly knoxvii genera of the Littorinidae. In common xxith all members of the ,sub- family Littorininae, its species occur t)n hard substrates in the httoral zone. Although easily accessible and often abundant, they are infre(juently collected oxving to their small size and crvptic habit in rock crexaces and among oyster and barnacle shells. All knoxvn species occur in the Indo-West Pacific province and most are entirely tropical, although several extend into the temperate zones of southern Africa and the northxx'esteni Pacific. Among Utt(jrinids, they are readily recognized by their small (1-7 mm diameter), trochoidal or depressed shells, lacking spines, with open umbihcus and multispiral oper- culmn. These characters are superficially similar to those of trochids (but lacking the nacreous interior of that group), with which they were often contused by eark authors (see review of taxonoinic histon- bv Reid, 1989a). Other littorinids with trochoidal sliells include the genera Bciiihicimu and m.scllop.sis irom Australia and New Zealand (Reid, 1988), and Tcctiiniis and C.i'u- chritis from the Indo-West Pacific and western Atlantic (Ro.sevvater, 1972; Reid, 1989b; Reid & Celler, 1997). ' Present address: 1.3/F, Canton Road Covemment Offices, Ag- riculture and Fisheries I^epartinent, Hong Kong For long these small littorinids were neglected ta.\o- nomicallv. The most recent monograph of e.xtant and some fossil Pca.sicllo species was that by Reid (1989a); before this the onlv compilations were a fist by Nevill (1885) and a monograph bv Trvon (1887). The anatomy (male and female reproductive tracts, paraspermatozoa, egg capsule, alimentary tract, head-foot pigmentation), radula, protoconch and teleoconch were described in de- tail bx' Reitl (1989a). As a result, the genus xxas defined not onlv bv shell and opercular characters, but iilso by the uni(jue combination of the elongate penis with single mamilliform peni;il gland and closed vas deferens, to- gether xvith the double-looped form of the pidfial ovi- duct, and other anatomical features. These same char- acters were included in a phvlogenetic an;ilysis of the morphology of the genera and subgenera of the Litto- rinidae (Reid, 1989b). However, since none of the coded anatomical characters appeared to be apomoqihic xxithin the famil), the (juK' formal svnapomorphies of the genus in this analysis were the trochoidal shell shape and mul- tispiral operculimi, and neither of these xx'as unitjue. Nexertheless, the close moiphological similarities among its members leaves little doubt that the genus is a mono- phvletic one; the arrangement of the loops of the p;dli;d oxiduct is one unitjue character, but oxving to difficulties of coding such structures it did not appear as such in the formal auiilvsis. As a result of Reid's (1989b) phylo- genetic analysis, it was clear that Peasiella was a member of the derived subfamilv Littorininae, and probably a relatively basal member, although its precise relation- ships xxith such genera as Maimvarin^ia, Ccnchhtis and Tcctariiis were not resolved. A recent molecular study included a species of Peasiella. vxith members of Tcctar- iiis. Ccncliriti.\ and Sodiliftorina. but did not find any close relationships xxith these other genera (Reiil & C;ell- er, 1997). In th(- previous systematic account ni Peasiella species (Reid, 1989al, anatomical details were provided for all the six species then recognized. Hovxexer, the amount of material available vxas limited; for example, for P. 'isseli' onlx one preseived male was dissected, and tor /' 'iiifraciKtalii' oulx seven. As a result, it was loiuluded D. G. Reid and Y.-M. Mak, 1998 Page 7 that the genus was anatoinicallv rather uniform, without the striking differentiation in reproductive characters that has proved so useful for the discrimination of spe- cies complexes elsewhere in the faniilv (e.g. Reid, 1986a, 1988, 1996). Instead, it was necessar\' to base species definitions largely on the shape, sculpture and color of the shell (although these characters were evidently sari- able), and on geographical distributions. Anatomical characters such as head-toot pigmentation, the number of cusps on the outer margiuiil radular tooth, and pres- ence or absence ot a copulatory bursa provided only mi- nor input. Our attention was once more drawn to the svstematics of PcasicUa as a result of a stud\- iiv one of us (Y-MM) of the littorinids ot Hong Kong. The fauna of Hong Kong was thought to include three TcasicUa species (Reid, 1992), the highest number then known to occur s\in- patricallv. However, it was tound that two distinct shell forms of P 'infracostata' scnsti Reid (1989a), both rare in Hong Kong and not yet found svnitopically (i.e. on the same shores), produced egg capsules of different shape. This prompted a reexamination ot this stipposed species throughout its wide range in the IncUan and Pacific Oceans. Meanwhile, further collections were made by Y- MM throughout the Far East. During the past decade much new material has also accumulated in the major museums. Using this new material, we found that penial torm was, after all, a character showing useful interspe- cific variation. As a result, we have revised the taxonomy of two species complexes; PcasicUa 'infracostata' scnsii Reid (1989a) is here showai to consist of five species, and P. 'isscli' scnsti Reid (1989a) of two. These seven species are fully described. We also record additional information on the variability, distribution and nomen- clature of other PcasicUa species. The number of species recognized in the genus is therefore increased from six to eleven. MATERIALS AND METHODS The descriptions and records are based on material in the following museums: Natural History Museum, Lon- don (BMNH); Museum National d'Histoire Naturelle, Paris (MNHN); Instituut voor Svstematiek en Populatie- biologie, Amsterdam (ZMA); Nationaal Natuurhistorisch Museum, Leiden (NNML); Institut Royal des Sciences Naturelles de Belgique (IRSNB); National Museum of Natural History, Smithsonian Institution, Washington, DC (USNM); Academ%- of Natural Sciences of Philadel- phia (ANSP); Museum of Comparatixe ZoologN, Harvard Universit}- (MCZ); Los Angeles Countx' Museum of Nat- ural History (LACM); Natal Museum (NM); National Science Museum, Tok-yo (NSMT): Department of Earth and Planetary' Science. Kyushu Universit)- (KU); Austra- lian Museum, Sydney (AMS) and the private collections of H. Dekker, The Netherlands (HD) and J. Le Renard, Paris (LR). In addition, t\pe specimens have been ex- amined from Oxford Uni\ersit\ Museum (OUM); Mu- seo Civico di Storia Naturale Giaconio Doria', Genoa (MGD) and Zoological Survey of India. Calcutta (ZSI). Unless otherwise indicated, all figured specimens are in BMNH. LectoKpes ha\e been designated from svmtypic series onK' for \-alid species names. Shell height (H) was measured parallel to the axis of coiling, and maximum shell diameter (D) perpendicular to this iLxis. The height/diameter ratio was used as a sim- ple index of shell shape. The number of whorls of the protoconch was counted as described by Reid (1996: 9). Where counts of the number of ribs above the periphery are given, this includes the strong rib at the periphery; this rib is not included in counts of the basal ribs. Li\ing material was preserved in 80% ethanol, and dissected under a binocular microscope. PiJUal oxiducts were ex- amined by dissection; in \ery small animals the copula- tory bursa can be impossible to find unless filled with recentK deposited sperm; to confirm absence of the bur- sa woukl re(juire histological preparation, but this was not done in the present study. The loops of the albumen and capsule gland follow a similar path in all species examined, and are therefore not re-described (see Reid, 1989a, tor description and figures). Radulae were cleaned ot tissue in a cold, ililute solution of hypochlorite bleach, thoroughly rinsed in distilled water, and mount- ed on a thin laver of polyvinyl acetate glue on glass cox'er slips. Ratlulae were examined with a scanning electron microscope. Unworn radular sections were photo- graphed from three orientations: vertically above flat radula (to show shape of teeth), at an angle of 45° from front end of radula (to show shape of tooth cusps), and at an angle of 45° from the side of the radula (to show relief). For three species, egg capsules were obtained from living animals. Females were placed inchvidually in plastic vials (20 X 47 mm) half filled with filtered sea- water Vials were covered and kept at room temperature (24-26° C) with no aeration. The water was renewed daily, and eggs were collected from the bottom of the containers. Eggs were preserved in 5% formalin in sea- water, for subsequent examination and measurement us- ing light microscop\-. The fists ot material examined are those records used tor the cUstribution maps. Numbers of specimens dis- sected are incUcated by M (male), F (female); R indicates radula preparation, and P protoconch examination. For the seven .species described in detiiil, the material listed mcludes that examined during the previous study by Reid (1989a), and the descriptions supercede the earlier account. This earfier work should still be consulted for detailed descriptions of the histology of the reproductive system and of the sperm For those species for which only additional notes antl records are given, full descrip- tions and other records can be found in Reid (1989a). SYSTEMATIC DESCRIPTIONS OF REDEFINED AND NEW SPECIES OF PEASIELLA PcasicUa infracostata (Issel, 1869) (Figures 1-5, 35-41, 76-79, 108-109, 131) Trochus sp, Audouin, 1826:42 Risella infracostata Issel, 1869:19.5-196, .348 [refers to Savigiiy, Page 8 THE NAUTILUS, Vol. 112, No. 1 1817: pi. 5, fig. 40.1, 40.2; 16^01)^6 (Reid, 1989a; fig.41) + .3 paraleetot)pes MNHN, seen; Suez (Issel)]. Pallary, 1926:86, pi. 5, fig. 40.1, 40.2. Bouchet & Danrigal, 1982: 13, fig. 64. Risella (Peasiella) infrocoslnta. — Ne\ill, 188.5:160. Tryon, 1887: 264, pi. 50. ficrs. 41, 42. Peasiclhi infracostata— -Reid. 1989a:61-64, figs. 106-109,112 [in part; includes P fasciata. P fuscopiperata. P. habei, P patula]. Bosch et al. 1995:46, fig. 118. ? Risella (Peasiella) tantillus var. subinfracostata Nevill, 1885: 160 [lectonpe (Reid, 1989a; fig. 35) + 2 syntypes ZSI; lectotype seen; Nicobar Islands]. Nomenclature: In his description of this species, Issel (1869) referred to Savigny's (1817) unlocalized figures of Egyptian material, and ;ilso mentioned three specimens of his own from Suez; the latter were not found by Bouchet & Danrigal (1982) in Issels collection in MGD. Reid (1989a) therefore designated one of four shells in the Savignv Collection in MNHN as the lectotype. The identit\- oi Risella (Peasiella) tantillus var. subin- fracostata Nevill, 1885, is uncertain. The lectot\pe (Fig- ure 35) measures 2. .3 mm in diameter by 2.2 mm in height (H/D 0.96), is almost smooth above the periph- ery, but for fine spiral microsculpture and traces of about four spiral grooves near the periphei;)'; the base bears four subequal ribs around a small umbihcus; the color is cream with a single brown spiral line on the first whorl of the teleoconch, and no other pattern. Of the known species of the genus, this shell can be compared with P. infracostata and P. fasciata. In outline it resembles P. infracostata, for the whorls and aperture are less round- ed than in P fasciata; however, it lacks the enlarged mid- basal rib that is often present in the former. The sparse color pattern is unlike any k-nown specimens of either of these .species. The locafity of Nevills shells, from the Nicobar Islands, is far from the clo.sest known occur- rence of P. infracostata in Bombay, and from that of P. fasciata in Borneo. The only other Peasiella species re- corded from the Andaman Sea is P. roepstorffiana (Reid, 1989a), but that .species usually has stronger spiral sculp- ture, flatter whorls and a strong peripheral keel. Until more information is a\iiilable, the name is tentatively included in the synonvmy of P. infracostata. Reid (1989a) confu.sed five species under the name P infracostata (here distinguished as P infracostata s.s.. P. fasciata, P. fuscopiperata. P. Iiahei and P. patula), misled by their similar and confusingly variable shells. Shell (Figures 1-5, 35-41): Adult size range 1.1-2.6 mm diameter. Shape: thickness varies from dehcate and translucent, to thick-walled and opaque; equilaterally conical to depressed-globular (H/D 0.7-1.0); outline domed; whorls usuallv rounded or with rounded shoul- der, becoming almost flat-sided on last whorl of largest, thick-walled shells; suture always distinct; periphery oc- casionally uniformly rounded, but usually marked by sharp angle or rib, rarelv de\eloped as a narrow flange; base usually shghtly rounded, becoming flat or even shghtlv concave in largest, thick-walled shells; umbilicus small to large, occasionally closed in thick-widled shells; columella narrow, uniformly rounded. Sculpture: proto- conch not seen, apex usuallv eroded; teleoconch whorls always lacking ribs above periphery, microscopically smooth or with fine spiral microstriae (rarely, about 6 striae mav be more prominent, but not developed into ribs. Figure 39); 1—4 ribs on base (rarely absent, or 5- 6), of which that at mid-point of radius is usually most prominent (producing shght angulation of base). Color: translucent yellow-browni, opaque white where shell is thicker at suture and periphery (entirely white in thick- walled shells); pattern of p;ile or dark brown or black, oblique or zigzag, narrow lines from suture to periphery, numbering .5-19, darkest at periphery and sometimes idso at suture; pattern occasionally represented only by peripheral stripes; spire mav be brown, or frequently marked only by single spiral brown line; base usu;dly pale with brown line in outermost groove, line some- times absent, base rarely brownish with paler ribs; col- umella and parietal area usually not colored, rarely browni. Animal: Head-foot: front of head blackish, with a sep- arate black band across base of snout; tentacles unpig- mented; foot may have black pigment around opercu- lum. Penis (Figures 76-79): filament long (50-60% total length), tapering; base slightly thicker, with single small mamilliform penial gland. Pallial oviduct: bursa not seen. Radula (Figt/rc.v 108, 109): cusps pointed, central cusp of rachichan tooth slightly apiculate; outer marginal tooth with 6-7 long, pointed cusps, outermost more rounded, becoming smaller toward micUine. Distribution: Habitat: Among barnacles in upper eu- littoral on surf beach (Oman; J. D. Taylor); rock pool on hmestt)ne, in upper eulittonil of sheltered shore (Oman; E. Glover); under rocks and stones in intertickil (Oman; R. G. Moolenbeek & H. Dekker); under coral blocks in eulittoral (Eg\pt; D. G. Reid). Abundant at some local- ities in Oman and at Karachi, but elsewhere appears to be scarce. Figures 1-5. Peasiella infracostata. 1. Has ;il-Hapt (BMNH 1882.8.7.316). .3()-.34. Pca.siclla m/iiiritiana .'50. Ll■c•to^pe o( Ri.sclla isseli var maiiritiana Viader, 1951, Grand Bav, Mamitius (BMNH 19S9()()4i 31. Arabian C;ult (BMNH 1996.396). .32. Inhaca I., Movamhiiiue (NM L3435). 33. Sadh, Oman (BMNH 1996397) 34. •/an/il)ar, Tanzania (BMNH 1904.10.20.97). Scale bar = 2 unn. D. G. Reid and Y,-M. Mak, 1998 Page 11 Page 12 THE NAUTILUS, Vol. 112, No. 1 ery uniformly rounded or marked by slightly thickened rib onlw in thicker shells peripher\' angled and marked bv sharp keel or narrow flange; base rounded; umbilicus of moderate size; columella rounded, shghtK thickened and angled at base. Sculpture: protoconch 240 |xm di- ameter, 2.5 whorls, protoconch II sculptured by scat- tered granules and narrow raised ridges (as in Figures 106, 107 of P. fasciata): teleoconch whorls usually mi- croscopically smooth above periphery, or with fine spiral microstriae, rarely with 2-6 slight or indistinct ribs; 2-6 (rarely 0, 1 or 8) ribs on base, outermost strongest, be- coming weaker toward umbilicus; basal ribs fine, indis- tinct or absent in thin-walled shells; rarely minute per- iostracal bristles present on basal and dorsal ribs. Color: thin-walled shells translucent pale yellow-brown with faint pattern of 6-14 oblique or waw bro\\ni lines (some- times present onlv as faint marbling), distinct only at suture and periphery, base unmarked; thicker shells opaque white or beige, with highly variable pattern: red- brown to black coarse, irregular pattern of bands, chev- rons and zigzags, usually leaxdng an unpigmented sutural band; sometimes a continuous broad spiral band on shoulder; sometimes a paler brown pattern of diffuse marbhng, with or without 9-12 darker peripheral spots; spire orange-brown, often with dark red-brown spiral band; base sometimes red-brown with 6-10 dark spots on white peripheral keel; lower columella and parietal callus sometimes red-brown. Animal: Head-foot: front of head blackish, with a sep- arate black band across base of snout; tentacles unpig- mented; foot with some black pigment on sides or un- pigmented. Penis (Figures 80-83): filament small (10- 20% total length), tapering; base thicker, sometimes wrinkled, with single very large mamilliform penial gland occupying most of its length. Pallial oviduct: bursa not seen. Radula (Figures 110. Ill): cusps pointed, major cusp of lateral and inner marginal tooth sometimes sUghtly rounded, central cusp of rachidian tooth sUghtly apiculate; outer marginal tooth with 5-8 long, pointed cusps, outermost more rounded, becoming smaller to- ward midline; base of rachidian tooth sometimes devel- oping a pair of small posterior denticles. Di.stribution: Habitat: Eufittoral; low, undercut lime- stone cliffs with fringing reef platform beyond (Kenya; D.S. Brown): alive in coralline algal debris washed up on strand line (South Africa; D. Herbert). Range (Figure 132): East coast of South Africa to Ke- nya. Records and material examined: South Africa: Port Al- fred (NM, OUM); East London (NM); Mbot\i. Pondo- land (NM); Port Edward (NM); Umdhloti, Natal (NM); Mapelane, Zululand (NM, 2M); Leven Point, Zululand (NM, IM). Mozambique: Benguera Island, Bazaruto Ar- cliipelago (NM); Conducia Bay (NM). Kenya: Tiwi Beach, 30 km S Mombasa (BMNH, 7M, 4F, 4R, IP). Remarks: Reid (1989a) identified smooth-shelled forms of this species as P. infracostata, and those rare examples with spiral sculpture above the periphery as P. isseli (with which P. mauritiana was then included). The form of the penis pro\dded the first evidence that this species is distinct; the short filament is found elsewhere in the genus only in P. patula, but there the mamilhform gland is of a different shape. The present species is svm- patric with tvvo others, P. mauritiana and P. infracostata, in East Africa, and penial shape is diagnostic of each. The shell is most similar to that of P. infracostata, and separation of the two has been discussed in the Remarks on that species. In both, the shell shows parallel variation in thickness, opacity, spire height, angulation of the mar- gin and intensitv^ of color pattern. Occasional shells with spiral sculpture abo\e the periphery' could be confused with those of P. mauritiana; P. fuscopiperata lacks the shoulder angulation of that species, is usually a more dehcate shell and rarely shows the same pattern of strong oblique radial lines. PeasicUa patula new species (Figures 11-13, 52-56, 84-87, 112, 113, 123, 127, 128, 133) Peasiella infracostata. — Reid, 1989: 61-64, fig. 110 [in part; in- cludes P. infracostata, P. fuscopiperata. P hahei. P. fa.scia- ta: not P infracostata (Issel, 1869)]. Etymology: Latin patulus, open, in reference to the relatively large aperture. Types: Holotype BMNH 1996276 (Figure 11); 5 para- types BMNH 1996277, 49 parat\pes in alcohol BMNH 1996278 (Figures 87, 112, 113)'. Type locality: Changi Point, Singapore. Shell (Figures 11-13, 52-56): Adult size range 1.2- 2.0 mm diameter Shape: depressed to coniciil (H/D 0.75-0.85); somewhat patulous, last whorl enlarged, ap- erture relatively large; protoconch (if present) projects as papillose tip to spire; whorls well rounded or inflated, often with rounded shoulder, suture impressed; angled peripheiT marked by shaq3 but bareh' projecting keel; base slightly rounded; umbihcus wide; columella narrow, cuned at base; viewed from above or below, apertural Figures 35—41. Peasiella itifracostala. .'$5. Lcctoh pi* u{ Risellu [Peasiella) lunlillus vm. subinfracostata Ne\ill, 1885, Nicobar Is (ZSI). 36, 37, 39, 40. Karachi. Pakistan (liMNH l'996383). 38. 4 km S Ras Qudufah, Masirali I., Oman (BMNH 1996.385). 41. Lectohpe of Risella infracostata Ls.sel, 1869, Red Sea (MNHN). 42^8. Peasiella fuscopiperata. 42, 43, 46, 48. Tiwi Beach, 30 km S Mombasa, Kenya (BMNH 1996.386). 44. Leven Point, Zuhiland Sontli Africa (NM E2745). 45. Mzaniba, Transkei, South Africa (NM .3028). 47. Mapi'lanc, Znhiland South Africa (NM D2415). Scale bar = 1 mm. D. G. Reid and Y.-M. Mak, 1998 Page 13 Page 14 THE NAUTILUS, Vol. 112, No. 1 edge is curved (i.e. growing edge is prosocyrt). Sculp- ture: protoconch 260 ^I,m diameter, 2.7 whorls, sculpture not preserved; teleoconch whorls smooth or with 6-7 indistinct to strong spiral ridges; 4— .5 fine rihs on base; surface smooth or with fine spiral microstriae; basal ribs occasionally bear minute periostracal bristles. Color cream to vellow ochre, with 4—7 fine bnmTi spiral lines above periphery' (including dark line at suture; some- times increasing to 9 at end of last whorl); 7-12 browm obhque short stripes at periphery and suture, often faint or absent, occasionally continuing indistinctly across dor- sal surface; base with 1-2 spiral brown lines at peripherv', margin of umbificus sometimes puiplish brown; colu- mella and parietal area purplish brown. Animal: Head-foot: usually entirely unpigmented; sometimes slight blackish pigmentation behind head. Pe- nis (Figures 84-87): filament slender, tapering, 30-40% total length; base thick, single large mamilliform penial gland (occup)ing 60-80% total length) with narrowly elongate reservoir; subepithelial glandular tissue at base of filament and around base of reservoir. Pallial oviduct: bursa not seen. Egg capsule (Figures 123. 127, 128): di- ameter 221.7 (j.m (SD 13.1 |jLm, n=10; ovum diameter 66.0±1.6 Jim; Singapore; this study), of depressed cu- pola type with 3 concentric rings on upper side and pleated margin (like a thick cogwheel), containing single ovTim. Radula (Figures 112. 113): all cusps shaiplv point- ed, central cusp of rachidian tooth slightly apiculate; base of rachidian tooth with two sharp posterior denti- cles; outer marginal tooth with .3—4 pointed cusps, be- coming smaller toward midline. Distribution: Habitat: Among barnacles and oysters in upper euUttoral, on rocky shores and sea walls (Sin- gapore). Range (Figure 133): Singapore, Gulf of Thailand, northern Borneo. Records and material examined: Singapore: Changi Point (BMNH; 1M,4F, 3R); Changi South (BMNH; 4M, 2F); St. Johns Island (BMNH; 2R, lP).Thailand: Pattava (Le Renard Colin). Sabah: Berhala Channel, Sandakan (USNM). Remarks: The expansion rate of the shell is larger in this species than in other members of the genus, so that it achieves a sUghtly, but tlistinctly, patulous shape which is characteristic. The color pattern resembles that of F. fasciata, ;ilthough the fines are usually less pronounced and the sutural and peripheral spots more developed. These two species are distinguished b\ their penial form. the filament being slender and short in P. patula, and the mamilfiform gland relatively larger. Their egg cap- sules differ in shape in samples of P. patula from Sin- gapore and P fasciata from Hong Kong The\ have not yet been found s\ntopically, but their known ranges ap- pear to overlap in northern Borneo; additional collecting in the region may well rev'eal wider SMiipatn. The radula of P. patula is peculiar, showing a p;ur of shaip posterior denticles on the base of the rachicfian tooth (Figures 112, 113). Among other littorinids, similar denticles have been described only in Melarhaphe neritoides (Bandel, 1974: figs. 58, 59), but the stnictures are clo.ser to the posterior edge of the tooth in P. patula, and are probably not homologous. Sfight development of basal denticles has been found in one example of P. fuscopipcrata and one of P. isseli. One specimen from the tvpe locality showed both a fulK- formed pallial oxiduct and a well developed penis (see remarks on pseudo-hermaphrodit- ism under P. habei). This is the rarest of all the Pcasiella species in museum collections, perhaps because of its limited geographical range; in Singapore it occurs abun- dantly. Pcasiella fasciata new species (Figures '14-17, 49-51, 8.8-91, 106, 107, 117, 122, 125, 126, 134) Peasielln infracostnta—Keid. 1989a:61-64, fig. 12, fig. .30 (rad- vila), fig. 48 (penis), fig, 112 [in part, includes P infracos- tata, P. fuscopipcrata, P patula, P. habei; not P. infracos- tata (Issel. 1869)]. Reid, 1992:194, fig. Ic (penis), fig. 2c (oviduct), pi. 1, figs, i, 1, 111 [in part, includes P. habei]. Etymology: Latin fasciatus, banded, in reference to color pattern. Types: Holotvpe BMNH 1996272 (Figure 15); 5 para- types BMNH 1996273; 1 parat>pe AMS C203254. Type locality-: Turtle Bay, Cape Ferguson, Queensland, Aus- tralia. SheU (Figures 14-17, 49-51, 106, 107): Adult size range 1.3-2.5 nun diameter. Shape: turbinate to de- pressed turbinate (H/D 0.78-1.12); outhne domed; whorls well roundeil, suture distinct; peripher\' uniform- 1\ rounded, or angled, or marked by a more or less prominent rib; base rounded; umbificus snuill or almost closed; columella narrow, uniformly rounded. Sctdpture: protoconch 240 [xm cfiameter, 2.6 whorls, protoconch I smooth, protoconch II with scattered small granules and narrow wavy or straight ridges, of which 6 are visible above suture (Figures 106, 107); teleoconch whorls Figures 49-51. Pcasiella fasciata, \}\o\ Mission. Cape Danipier, New Bntam, I'apua New Guinea (AMS C163128). 52-56. Pcasiella patula. 52. Changi Point, Singapore (BMNH 1996278). 53-55. St. Johns I.. Singapore (BMNH 1996387), 56. Pata\a, Thiulaiid (LR 990f2), 57-«l. Pca.'iiella habei. 57. 4-5 km SW Tsutsu, Tsushima I., Nagasaki Pref , Japan (BMNH 1996389). 58, 59. Nejiko, Hirado I., Nagasaki Pref., Japan 60. Ma Liu Shi, Tolo Harbour, Hong Kong (BMNH i996.393). 61. Zanpa Point, Okinawa, Japan (AMS C146702). Scale bar = 1 mm D. G. Reid and Y.-M. Mak, 1998 Pa^e 15 Page 16 THE NAUTILUS, Vol. 112, No. 1 smooth above periphery, with fine spiral microstriae, rarelv with .3—4 faint groo\-es: 3—4 ribs on base (rarely 1- 6), becoming stronger toward periphen: Color translu- cent cream to opatjiie gra\ish white; pattern of brown, dark red-brown or black lines or bands, numbering 2-8 above periphery on last whorl (only 2—4 on spire whorls), bands often altematelv thick and thin on last whorl; in addition sometimes 6-13 f;unt brownish spots at suture and periphery ; 2—4 dark lines in grooves at periphery of base; base of columella and umbilical area sometimes dark purphsh brown. Animal: Head-foot: front of head blackish, with a sep- arate black band across base of snout; tentacles unpig- mented or with single longitudinal black line; toot un- pigmented or with blackish sides. Penis (Figures 88-91): filament 40—50% total length, cylindrical and rounded at tip; base thicker, with single maniillitorm penial gland of moderately large size (30—40% total length). Pallial ovi- duct: bursa present. Egg capsule (Figures 122. 125, 126): diameter 215.4 ^.m (SD 5.8 jim, n = 10; ovum diameter 77.8±2.9 |jim; Hong Kong; this study), of depressed cu- pola Kpe with 4 concentric rings on upper side and pleated margin with peripheral keel, contiiining single ovum. Radula (Figure 117): cusps pointed, central cusp of rachidian tooth shghtly apiculate; outer marginal tooth with 5-6 long, pointed cusps, outermost more rounded, becoming smaller toward midline. Distribution: Habitat: among algae in rock pools (type locahtv; A. Mitchell); among short calcareous green algae and Caidctya, on wave-exposed rocky plat- form (Cape Dampier; P. H. Colman); among euhttoral cobbles, cre\dces in granite and sedinientarv rocks, among Saccostrca, on fairly sheltered and wave-exposed coasts (Hong Kong; J. D. Taylor, Y.-M. Mak). Appears to be generally scarce. Range (Figure 134): Queensland, New Guinea, Bor- neo, Vietnam, Hong Kong, Taiwan. Records and material examined: Queensland, Austra- Ua: Facing Island, Port Curtis (AMS); Turtle Bay, Cape Fergu.son^AMS; IM, IR); ILdfmoon Bay, Cairns (AMS); Green Island (AMS); Port Douglas (AMS). New Britain, Papua New Guinea: Uvol Mission, near Cape Dampier (AMS; 3M, 2F, 2P); Duke of York Island, Ral)aul (AMS). Sabali: Bak-Bak, 9.6 km N Kudat (USNM). Vietnam: La Table Island. Tonkin (MNHN). Hong Kong: Wu Kwai Sha (BMNH; 2M, 2R); Peng Chan (BMNH; IM); Hoi Ha (BMNH; 2F); C:ape d'Aguilar (BMNH). Tmwan: Shi Cheng, Taipei (BMNH). Remarks: The first evidenc-e that this species was chs- tinct from P. hahei (both confused under P. infracostata by Reid, 1989a, 1992) was provided bv their differently shaped egg capsules in Hong Kong (Figures 122, 124). Differences in shell shape and penial lorni were then recognized, as describi'il in the Remarks on P. hahei. The shell of P. fasciata is easily recognized by its com- bination of dark spiral lines, tinliinate shape with well- rounded whorls, and small umbihcus. Spiral color bands are found in some other Peasiella species, especially on the spire whorls, but are onlv well-developed on the last whorl in one other species, P. patula. which is distin- guished by its patulous shape. Anatomically, P. fasciata is characterized by its penis with long, cylindrical fila- ment and moderatelv large mamilhform gland. As in P. infracostata and P. fuscopiperata. the shell can be opaque and solid, or translucent, although it is never as dehcate as in some examples ot those species. This spe- cies is rare in collections. Peasiella hahei new species (Figures 18-24, 57-61, 92-98, 114-116, 124, 129, 130, 135) Litlorinn-capsuln hahei Tokioka, 1950:151-1.52, fig. 6.1 (egg capsule) [Ago and Taiiabe Bavs, Japan; name una\ailable, see below]."Habe, 19.56:117-121. Peasiella roepstorffiana. — Habe. 1956;lliS-121, fig. A (egg cap- sule). Oyama & Takemura, 1961: Peasiella and Littorinop- sis pi., figs. 1-3. Yaniamoto & Habe, 1962:16, pi. .3, figs. 3, 4, figs. 34, 35 (egg capsule). Amio, 1963:.303, figs. 22a, b (egg capsule). Habe, 1964:28, pi. 9, fig. 23. Higo, 1973: 46. Habe, 1984:11. fig. 1. Ohtsuka & Yoshioka, 1985:232, fig. 4B. Okutaiii, 1986:71, unnumbered pi Fukuda, Ma- shino & Sugimura, 1992:57, pi. 43, fig. 126. Higo & Goto, 1993:74. [All not P. roepstotfiana (Ne\ill, 1885)[. Peasiella infracostata— Reid, 1989a:61-64, fig. 10, fig. 29 (rad- ula), fig. 47 (penis), fig. 64 (egg capsule), fig. Ill [in part; includes P infracostata, P. fuscopiperata. P patula. P. fas- ciata]. Choe, 1992:290, .591, fig. 56. Reid, 1992:194, pi. If [in part: includes P fasciata]. Choe & Park, 1993: 19-20, text fig. 2. [All not P infracostata (Issel. 1869)]. Etymology: To honor Prof T Habe, who has contrib- uted much to the studv of Japanese httorinids. Types: Holotxpe BMNH 1996274 (Figure 20); 9 para- types ni alcohol BMNH 1996275. Type locidity: Esu Cape, Shirahama, Wakayama Prefecture, Japan. Nomenclature: Uniquely for a Uttorinid, the first name for this species was applied to its egg capsules. Tokioka (1950) described two types of httorinid egg cap- sules in plankton hauls from Ago and Tanabe Bays in southeastern Honshu, to which he gave Latin names. Since the httorinids that produced the capsules were not kniowni, he coined a new genus Littoiina-capsula. Sub- secjnentlv, Habe (1956) showed that one of these cap- sides, Littorina-capsula hahei, was the egg of the com- mon Japanese species of Peasiella (which he misidenti- fied as P. roepstoiffiana). However, it is debatable wlu'ther the specific name gi\en by Tokioka is nonien- ilaturalK a\ailable. Habe (1956) apparenth' considered it so, printing it in italics. Others have not done this, and thus appi^ar to have judged it unaviiilable (Amio, 1963; Reid, 1989a). In its favour it can be argued that the name is uniuubiguous, since the egg capsule is distinctly different from those of other known Japanese httorinids. A name based on ouK one stage in a fife cycle, while undesirable, is not iherebv made unaNaikdile (ICZN Art. D. G. Reid and Y.-M. Mak. 1998 Page 17 17). The generic name Littorina-capsula is una\;dlable (since it is not accompanied bv fixation of a t\pe species, ICZN Art. 13b), but this does not affect the standing of a specific name combined with it (ICZN Art. llh). The inclusion of a hvphen in the generic name is an incorrect spelling (ICZN Art. 32c), and does not indicate that the name is non-binominal. On the other hand, it appears that Tokioka did not intend to introduce new names for httorinid species, for he stated: For con\enience of re- cording these capsules met with during our plankton studies, I propose to give them the following provisional names'. This, we believe, debars the names from no- menclatural consideration, since the ICZN Code specif- ically excludes names proposed "as means of temporan. reference and not for formal taxonomic use as scientific names' (ICZN Art. lb). This case is a complex and un- usual one, and is open to altemati\e interpretations. If Toldoka's name were to be accepted, the egg capsules studied by him would have the status of type material; inquiries from the Seto Marine Laboratorv have shown that no such material exists there (S. '^'amato, pers. comm.). In this case, our choice is to re-describe the species as new, based on type material of adult aniniiils collected near the locality of Tokioka's capsules. How- ever, in recognition of Tokioka's discoverv of the egg cap- sules, his intention to honor Prof T Habe, and to pro- vide continuitv, we use the same specific name. In the Japanese literature this species has generallv been misidentified as P. rocpstotjfiana, while Reid (1989a, 1992) incorrectlv included it under P infracos- tata. Shell (Figures 18-24, 57-61): Adult size range 1.6- 3.8 mm diameter Shape: equilaterally conical or sfightK- more depressed (H/D 0.60-1.03); outhne domed; whorls almost flat-sided or rounded or shghtlv shouldered, su- ture inconspicuous or impressed; peripheral keel prom- inent, often a projecting flange, rarelv slightlv undulat- ing; base flat to slightlv rounded; umbilicus usualK' nar- row; columella narrmv, cun'ed at base. Sculpture: pro- toconch 230 |xm thameter, 2.5 whorls, sculpture not preserved, apex usually eroded; teleoconch whorls usu- ally smooth, with spiral microstriae, sometimes with 6- 11 equidistant spiral grooves above peripherv in largest specimens; .3-.5 (rarelv 6) shaip ribs on base, equidistant or becoming closer around umbilicus; basal ribs (and oc- casionaUy dorsal ribs) may bear periostracal bristles. Col- or: variable; fawn, cream or ochre, paler at peripherv, with obhque (sometimes axial or zigzag) anastomosing browm stripes (sometimes faint, especiaUv in Hong Kong), always darkening to form single row of 7-12 more or less conspicuous large brown or black spots near periphery (occasionallv fusing to give a sofid spiral band) which become narrower as they extend over pale pe- ripheral keel; pattern on spire whorls darker, often blackish where eroded (especiallv in Japan); occasionallv 2^ spiral Isrown lines on spire whorls (common in Oki- nawa), usuallv disappearing on last whorl, but rarelv per- sisting as 5-9 spiral lines; base ochre to brown, darker near umbilicus and in peripheral groove adjacent to keel; sometimes shell more darklv patterned, spire and base black, last whorl with obhque anastomosing black stripes on whitish ground (some from Japan and northern Chi- na). Animal: Head-foot: front of head blackish, with sepa- rate black band across base of snout, occasionallv almost unpigmented; tentacles unpigmented; sides of foot sometimes slightly pigmented. Penis (Figures 92-98): fil- ament long (50-60% total length), vemiiforni; base shghtly thicker, 1 (rarely 0 or 2) small mamilhform penial gland. Pallia] oviduct: bursa present. Egg capsule (Fig- ures 124. 129, 130): diameter 261.3 |xm (SD 0.5 |xm, n=10; ovum diameter 68.3±2.3 |xm; Hong Kong; this study), 150-250 jjim (Japan; Tokioka, 1950; Amio, 1963; Ohtsuka & Yoshioka, 1985), of cupola Upe with 4 con- centric rings on upperside, containing single ovum. Rad- ula (Figures 114-116): cusps pointed, central cusp of rachidian tooth shghtly apiculate, major cusp of lateral and inner marginal teeth sometimes shghtlv rounded; outer marginal tooth with 4-6 (rarelv 3) long, pointed cusps, outermost more rounded, becoming smaller to- ward midline. Distribution: Habitat: In Japan this species is abun- dant in crevices and among barnacles in the middle and upper eulittor;il zone, on sheltered and moderatelv ex- posed rock"\ shores; on exposed shores it shows a pref- erence for surfaces protected from wave action (Mori et al. 1985a, b; Tanaka et ai, 1985). In Hong Kong and Taiwan it appears to be rare, and is found among bar- nacles and oysters in the uppermost euhttoral zone on sheltered shores. Range (Figure 135): China, Korea, Taiwan, Rvukxii Is- lands, Japan. Records and material examined: China: Ma Liu Shi, Tolo Harbour Hong Kong (RMNH; IM, IF, 2R); Lok Wo Sha Bay, Tolo Channel Hong Kong (BMNH); Pao- tai, Xiamen (BMNH; IM, IF); Gu Leng Yu, Xiamen (BMNH; IM, 3F,1R, IP); Tai Ping Bay, Qingdao (BMNH; 2M, 2F, IR); Hui Quan Bav, Qingdao (BMNH). Korea: Taesori, Sangch'uja I. (Choe, 1992; Choe & Park, 1993). Taiwan: Yehhu, Taipei (BMNH). Japan: 1.5 km WNW Onna, Okinawa (AMS); Bolo Point, Okinawa (AMS, IF, IR; USNM, NSMT); Baten, Oki- nawa (BMNH; IM, 2F, 3R); Tomioka. Kumamoto Pref (NSMT, 2R); Nagasaki (USNM); Omura Bav, Nagasaki Pref (NSMT); Goto, Nagasaki Pref (AMS); Nejiko, Hir- ado I., Nagasaki Pref (BMNH; IM, IF); Tsut.su, Tsush- ima I., Nagasaki Pref (BMNH); Fukoivoshi, Fuk-iK)ka Pref (KU); Kure, Hiroshima Pref (KU); Hanazura, Ko- chi Pref (MNHN); Tatsukoislri, Kochi Pref (USNM, NSMT); Shirahama, Tanabe Bay, Wakavama Pref. (BMNH; 5M, 2F, IR; USNM, NSMT); Shionomisaki, Wakavama Pref. (NSMT); Kuchino, Shiznoka Pref. (BMNH, 4M, 2R); Arasaki, Kanagavva Pref (USNM, NSMT); Oga Peninsula, Akita Pref (KU); Asamushi, Ao- Page 18 THE NAUTILUS, Vol. 112, No. 1 mori Pref. (USNM, NSMT); 2.5 km N Tomari, Aomori Pref. (AMS); Kominato, Aomori Pref. (LACM). Remarks: This .species is variable in conspicuous fea- tures of the shell including color, spire profile and sculp- ture, but consistent characteristics are the row ol dark spots above the periphery', which e.\tend onto the pale peripheral keel, the darker and often black spire whorls, and the prominent keel at the periphery. There are some recognizable geographical trends in shell characters. In Japan and northern China the shape is taller, with flatter whorls and a sharper peripheral keel; periostracal bris- tles are present at least on the base of most well-pre- served specimens; die spire, usually eroded, is black or darklv patterned (Figures 18-21, 57-59). Shells from the southern parts of the range are distinctly different. Those from Okinawa have a pale pattern; the dark pe- ripheral spots are most conspicuous on the spire whorls, vv'hich are also often marked bv 2-4 spiral brown lines; the whorls are sometimes rounded at the shoulder, and usually bear strong spiral grooves above the periphery, and the peripher;il keel is less pronounced; periostracal bristles have not been seen (Figures 23, 61). Shells from Hong Kong and southern China are similar in shape to those from Okinawa, but are usually smooth above the periphery, sometimes bear periostracal bristles on the base, and the color is ochre with strong peripheral spots and darkly patterned spire (Figures 24, 60). However, these southern populations are also variable, and inter- mediates apparentlv connect them with the typical northern forms (Figure 22). Additional material, partic- ularlv from the R\-uk-\ii Islands, would be desirable to investigate this \ariation in more detml, and to confirm our interpretation. The penial shape is similar through- out the range. The egg capsules from Hong Kong (fig- ures 124, 129, 130) are almost identical in shape to those figured from Japan by Toldoka (1950; reproduced b\ Ya- mamoto & Habe, 1962; Reid, 1989a), Habe (1956) and Ohtsuka & Yoshioka (1985). However, at a diameter of 261 fjim they are slightlv larger in size than Japanese capsules, measured as 150-2()0(xm (Tokioka. 1950), 150 |xm (Amio, 1963) and 220-250 (xm (Ohtsuka & Yoshi- oka, 1985). The rounded shells with lined pattern on the spire that can be found in Okinawa bear some similaritii- to those of P. fcisciata from Hong Kong. This led Reid (1989a) to cemsider them conspecific (as P. infracostata) . However, new material from Hong Kong has showni that both species occur there, although so tar the\ have not been collected syntopicallv (i.e. on the same shores). In Hong Kong the shells of P. luihci only occasionally bear 1-2 indistinct spiral brown lines above the periphery on the last whorl, and their conspicuous peripheral spots and darkl\' patterned spire differ from the 4—8 strong spiral lines of local examples of P fa.sciata. Anatomically, the mamilliform penial gland of P. Iiahei is smaller and the penial filament relatively narrower and shghtly more elongate than in P. fasciata. The egg capsules of these two species (sampled in Hong Kong) also differ (Figures 122, 124). Two other species are sometimes svnipatric with P. luibci in the southern part of its range and might be confused with it, P. hitulcnta in Hong Kong and P. roep- stoijftana in the southern Rviik-yu Islands, Taiwan and Hong Kong. PcasicUa hitulcnta has 5-9 strong spiral ribs above the peripherv, usuallv a gap between inner and outer series of ribs on the base (ribs all eijuidistant in P. liabei), the columella has a thickened angulation at the base, and the color pattern is of 7-13 broad and irreg- ular, dark axial stripes (see Reid, 1989a, 1992). Anatom- icallv the two are doselv similar; the penial filament of P. Iiahci is shghtly narrower and more elongate, and there are 4-6 cusps on the outer marginal tooth, but only 3 in P lutulcnta. Peasiella rocpstotjfiana (Figures 142-145) has 4—13 strong spind grooves above the pe- ripherv, frefjuently shows radial plications near the su- ture and periphery, which give rise to strong crenulations of the peripheral keel; the innermost one third of the basal rathus lacks ribs; the color is often bright chrome vellow or orange, with peripheral brown spots or band. Anatomically, P. rocpstoijfiana from Hong Kong lacks the mamilliform peniid gland, and the copulators- bursa is absent. Some abnormalities of the reproductive tract of P. lui- bci have been found. A single specimen from Shiznoka Prefecture had no maminiform penial gland (Figure 98), and one (parasitized h\ trematodes) from Tanabe Bay had two glands of normal size (Figure 97); in each case normal specimens occurred in the same lots. One other specimen from the latter sample had both a small, fully formed penis and a small oxaduct. Genuine hermaph- roditism has been described in onlv one littorinid genus, Mainwariufiiii (Reid, 1986b), but pseudo-hermaphrodit- ism (involving development of a penis and other male characters in females) is known in several littorinid gen- era (Reid, 19S6b, 1996), and is sometimes a response to pollution bv organotin compounds (Bauer ct at.. 1995). Peasiella species are not known to be tnily hermaphro- ilite, antl tiiis example is probably a case of pseudo-her- maphrocbtism. Peasiella i.s.seli (Semper in Issel, 1869) (Figures 2.5-29, 62-67, 99-101, 118, 119, 136) Trochiis sp. Audonin, 1826:42 Riselhi isseli Semper in Issel, 1869:194, .347 [refers to Savigiiy, Figures 62-67. Pctisicllti isseli. 62. Lectotype of Risella isseli Semper in Issel, 1869. Red Sea (MNHN). 63, 65, 66. Sue/., Egvpt (BMNII ISSS.IO 14) 64, 67. llmijliada, Eirvpt (BMNH 1996.39,5). 68-75. Peasiella mauritiana 68. Tiwi Beach, 30 km S Mombasa, Kenya. 69, 70, 73. Sedli. A! Kliassali, Oman. 71. Kuwait. 72, 74. Bandar Kliasran, Oman. 75. Bale Temay, Mahe, Seychelles. Scale bar = 1 mm. D. G. Reid and Y.-M. Mak, 1998 Page 19 Page 20 THE NAUTILUS, Vol. 112. No. 1 1817: pi 5. figs. .35.1, .35.2; lectohpe (Reid, 1989a: fig. 62) + 2 paralectohpes MNHN, seen; Suez (Reid, 1989a); 2 paralectohpes in MGD ex-Seniper Colin, seen, from Zan- zibar are P mauritiana]. Pallarv, 1926:84, pi. 5, fig. 35.1. 35.2. Laniv, 19.38:71. Moiizzo, 1939:183. Bouchet & Dan- rigal, 1982:13. fig. 65. Risella {PeasieUa) isseli. — Nevill, 1885:160-161 [in part: in- cludes P. mauritiana]. Trson, 1887:263, pi. 50. figs. .39, 40. PeasieUa ).s.se/i.— Franc, 1956:25. Mastaller. 1979:40. Reid, 1989a:60-61, fig. 13, fig. 50 (penis), fig. 56 (sperm), figs. 97, 98, 101, 103 [in part, includes P. mauritiana]. Trochus sismondae Issel, 1869:225-226, pi. 2, fig. 13 [holotype MGD, seen; Suez]. Risella isseli var. carinata Pallarv', 1926:84-85, pi. 5. fig. 36 [re- produced from Savngnv, 1817: s\mt\pe MNHN, seen; Suez]. Risella i.sseli van undata Pallarv', 1926:84, pi. 5, fig. 35.3 [re- produced from SaMgnv, 1817; t\pes not in MNHN; Suez]. Nomenclature: Throughout its taxonomic history this species, endemic to the Red Sea, has been confused with P. mauritiana, and the two taxa have not previously been distinguished at the specific level. In the original de- scription, Issel referred to the figures of Sa\dgnv (1817) based on Eg\ptian specimens. He also mentioned his own material from Suez and Semper's from Zanzibar, of which the latter (in MGD) is P. mauritiana. Reid (1989a) designated one of the shells from the Sa\ignv Collection as the lectot)'pe, and restricted the type locality to Suez, thereby fixing the identity of this species. Shell (Figures 25-29, 62-67): Adult size range 1.7- 4.2 mm diameter Shape: conical to depressed conical (H/D 0. 5.5-1. 23); outhne domed; whorls usually with rounded or angled shoulder, occasionalh' flat or only gently rounded; suture usually distinct; periphery sharply angled, with prominent rib or flange; dorsal sculpture sometimes slightlv nigose, but only rarelv does periph- eral flange show slight crenulation; base flat or slightly rounded; umbilicus small to moderate; columella round- ed, sometimes with a slightly thickened angle at base. Sculpture: protoconch 240 |xm diameter, 2.3 whorls, protoconch II v\ath scattered small granules and narrow wavy or straight ridges, of which 5 are visible above su- ture (as in Figures 106, 107, of P. fasciata): teleoconch whorls with 6—15 narrow ribs, uniform or unequal in size, above periphery; rib at shoulder sometimes slightlv enlarged; surface above periphery' covered with fine spi- ral microstriae; base with 4-6 (rarely 2-7) ribs, larger toward peripherv: Color: extremely variable; uniform cream, ochre, orange-browai, dark chestnut brown or en- tirely black; often with darker pattern of fine dots or lines in dorsal grooves; dorsal surface sometimes with fine white and brown marbled pattern; rarely a dark spi- ral band on dorsal surface; sometimes 7-15 brown spots on white peripheral keel; rarely pale sutural spots in ad- dition to darker peripheral spots, but only very- seldom does an indistinct pattern of broad radial bands appear; spire sometimes pink or crimson; apex often lilac or blackish; base lacking color pattern or with small spots in grooves. Animal: Head-foot: head unpigmented or blackish in front, separate black band across base of snout; tentacles usually with 2 longitudinal black fines; sides of foot with some black pigmentation or unpigmented. Penis (Fig- ures 99-101): filament long (509f total length), tapering at tip; base thicker, with single mamilfiform penial gland of moderate size. Pallial oviduct: large anterior bursa present. Radula (Figures 118, 119): cusps pointed, major cusp of lateral and inner marginal teeth slightK rounded, central cusp of rachidian tooth sometimes slightK- apic- ulate; base of rachicfian tooth occasionally with slight de- velopment of pair of posterior denticles; outer marginal tooth with 4 pointed cusps, outermost more rounded, becoming smaller toward midline. Distribution: Habitat: Abundant under coral blocks in upper eulittoral on a moderatelv sheltered shore (Egypt; D. G. Reid). Range (Figure 136): Red Sea and Aden. Records and material examined: Israel: Eilat (BMNH, IM, 2F; USNM, ANSP). Egypt: Faraun Lsland, Gulf of Aqaba (HD); Ras el Kura, Daliab, Sinai (HD); Suez (BMNH, IP, 2R; MNHN); Suez Canal (Tillier & Bavay, 1906; Moazzo, 19.39); Hurghada (BMNH, 3M, 2F 2R; HD). Sudan: Dungunab Bay (BMNH); Port Sudan (BMNH; ANSP). Saudi Arabia: Jeddah (USNM, MNHN). Eritrea: Mas.sawa (BMNH, NM); Dahlak Ar- chipelago (BMNH). Yemen; Perim Island (MNHN); Aden (MNHN). Remarks: The shells of P. i.s.seli and P. mauritiana are very similar, and the two have not previously been dis- Figures 76-79. Penes of PeasieUa infracostata 7fi, 77. Tivvi Beach, 30 km S Mombasa, Ketna. 78. Karachi. Pakistan. 79. Ras al-Hadd, Oman. 80-83. Penes of PeasieUa Juscopiperata. 80, 81. Tiwi Beach. .30 km S Mombasa, Kenva. 82, 83. Mapelane, Zululand, South Africa (NM D5822). 84-87. Penes of PeasieUa patula 84-86. Cliangi South, Smgapore' 87. Paratvpe, Changi Point, Singapore (BMNH 1996278). 88-91. Penes of PeasieUa faseiata 88, 89. U\ol Mission. Cape bampier. New Britain, Papua New Guinea (AMS C163128), 90. Wu Kwai Sha, Hong Kong. 91. Peng Cliau, Hong K.ong. 92-98. Penes of Pea.siella habei. 92. Tai Ping Bay, Qiugdao, China. 93. Sliirahama, VV'akayama Pref , Japan. 94. Batcn, Okinawa, Ja]ian 95-97. Fujisliima, Tanabe Ba\. VVakayama Pref., Japan. 98. Kuchino, S\miga Bay, Sliiznoka Pref., Japan. 99-101. Penes of PeasieUa isseli. Hurghada, Egvpt. 102- 105. Penes of PeasieUa mauritiana 102. Khor Hajar, Ras al-Hadd. Oman. 103. Anse La Mouche, Mahe, Seychelles. 104. Mera- wwah I., Abu Dhabi. 105. Tiwi Beach, .30 km S Mombasa, Kenva. Note that stnictnre of mamilliform penial glands is visible by transparency; mucous reservoir is shown by solid line, and extent of subepithelial glandular tissue by dotted line. The penial base is .sometimes wrinkled. D. G. Reid and Y.-M. Mak, 1998 Page 21 76 77 78 79 80 81 82 83 92 93 94 95 96 97 98 99 100 101 102 103 104 105 Page 22 THE NAUTILUS, Vol. 112, No. 1 tinguished as separate species. From the new anatomical material av;dlable for each species, it is clear that they can be diagnosed by the form of tiie penis. This is of the common type for the genus in P. isseli, with long filament and small mamilliform gland, whereas in P mauiitiana from throughout its large range it has a short filament with mucronate and hooked tip, and a large gland. Having separated the two using this character, correlated differences in shell shape and coloration can be recognized. In both, the development of the shoulder angulation is variable; in P. isscli it is often absent or represented only by an angulation of the profile, more rarely bv a prominent rib; in P iiiaiiritiana the shoulder is usuidly marked by a shaip angulation, often with an enlarged or even carinate rib. The coloration is also dif- ferent; in P. mauiitiana there is usually a pattern of obliquely ratlial lines or stripes, often darker at the su- ture and periphery; in P. isseli the color is highly variable, but radial stripes rarely appear, and the apex is often pink or blackish. Using these characters, together with knowledge of the regional variation in the shell of P. mauritiana, it has been possible to identify shells with confidence. So far, it appears that P. isseli is restricted to the Red Sea and Aden, whereas P. mauiitiana is wide- spread in the Indian Ocean and Persian Gulf The two apparently occur sympatrically just at the mouth of the Red Sea. Two dead shells of P. isscli have been recorded from Aden (MNHN), from which locality five other sam- ples contained only P. mauritiana. Both have been found together in a single lot of dead shells from Perim Island, South Yemen (Jousseaume Colin, MNHN), which con- tained 50 P. isseli and 8 P. mauritiana, readily separated by their .shell characters. The only other species to occur sympatrically with P. isseli is P. infracostata. These two are readily distin- guished by the presence of spiral ribs above the periph- ery in P isscli (absent in P. infracostata}, anil by the color pattern of obh(jue or zigzag lines in P. infracostata. The shape of the penis is, however, similar in both. Most Peasiella species show variation in color pattern, sometimes on a regional or inteqiopuhitional scale. This species is unique in the genus m its extreme intrapop- ulational variation; shells from beneath the same rock mav range from cream to p;itterned to black. Further- more, occasional specimens can be found in which the color changes from cream on the spire to entirely black on the last whorl (Figure 26). Peasiella mauritiana (Viader, 1951) (Figures .3()-.34, 68-75, 102-105, 120, 121, 1.37) Ri.sella mr/i — Lssel, 1869:194 [in part; includes P. mr/i]. Riwlta {Peasivlla) me/i— Ne\-ill, 188.5:160-161 [in part; in- cludes P issi'ti]. MeK-ill & Standen, 1901:.364 [not Semper in lssel. 1869[. Peasiella me/i.— Reid, 1989a:60-61, figs. 14, 1.5, fig. 28 (rad- ula), fig. 49 (fiead), fig. 97, 99, 100, 102, 104, 105 [in part; includes P isseli]. Bosch et ai, 1995:46, fig. 119 [not P. isseli (Semper in lssel, 1869)]. Ri.selln infracostata. --Ddutzenherg, 1929:496 [not lssel, 1869]. Risella is.seli van mauritiana Viader, 1951:149, pi. 3, figs 10, 11 [lectotype, here designated. Figure .30, -I- 2 paralecto- types, 'seen, BMNH 1989004; Grand Bay, Mauritius; 2 probable paralectotvpes Mauntius Institute, not seen]. Nomenclature: Hitherto, this species has always been confused with P isseli. It was first named, as a variety of that species, bv Viader (1951). Twt) specimens of Ri- sclla isseli var. mauritiana were exhibited in the natural history museum of the Mauritius Institute (R. Gajeelee, pers. comm. 1985); these have not been examined, but are likely to have been syntypes. In addition, there are 3 specimens in the Winck-worth Collection in BMNH, labeled 'Peasiella isseli v. mauritiana Viader MS' in Winckworth's hand; these are from Grand Bay, one of the two lociilities mentioned in Viader's (1951) descrip- tion. It is knowai that Winck-worth received specimens from Viader (see also acknowledgement of Winck-worth's assistance in determinations, Viader, 1951:1.36), and it is likely that this material was sent by Viader before Winck- worth's death in 1950. Since Viader (1951) chd not des- ignate tvpe specimens, a lectotype is here designated from the BMNH materiid (ICZN art. 74; recommen- dation 72B). Shell (Figures 30-34, 68-75): Adult size range 1.6- 5.6 nun diameter Shape: highly variable; depressed con- ical to Hattened (H/D 0.42-1.03); outline ilomed, straight or spire sfightly concave; shoulder usually sharp- Iv angled, or with projecting rounded carina (some Per- sian Gulf and northern Oman specimens), or sometimes becoming rounded or even flattened on last whorl; su- ture usuallv distinct; periphery sharply keeled or flanged, or with flaring rounded carina (some Persian Gulf and northern Oman specimens); shoulder occasionally reg- idarlv nigose and peripheral flange with 7-8 slight cren- ulations (some East African specimens); base flat to rounded; umbilicus small to large; columella rounded, sometimes indistinctlv thickened and angled at base. Figures !()(>, 107. Frotoconch of Peasiella fasciatn, Uvol Mission. Cape Danipier, New Britain, Papua New Guinea (AMS C163128). Figures 108, 109. Radulae oi Peasiella infracostata. 108. Karachi, Pakistan (flat view, shell D = 1.5 nun). 109. Ras al- Hadd, Oman (flat view, shell D=1.9 mm). 110, 111. Two \iews (flat and at 45°) of radula of Peasiella fiiscopiperata, Tiwi Beach, 30 km S Mombasa, Kenya (slieii D=1.7 mm) 112, 113. Two views (flat and at 45° from side) of radula of paratype o( Peasiella patula. Changi Point, .Singapore (BMNH 199627S, shell D=1.7 mm); note posterior denticles on base of rachidian tooth (arrow). Scale Inirs: 106, 107 = 100 |jlui; 108-113 = 20 jxm. D. G. Reid and Y.-M. Mak, 1998 Page 23 Page 24 THE NAUTILUS. Vol. 112. No. 1 Sculpture: protoconch 220 [im diameter, 2.2 whorls, protoconch II with .scattered small granules and narrow wa\y or straight ridges, of which 6 are visible above su- ture (as in Figures 106. 107. of P. fasciata); teleoconch whorls usually with 6-12 fine spiral ribs above periphery; sometimes onlv single rib at shoulder, or absent; surface above peripher\' covered with fine spiral microstriae; base with 4-7 (rarely 1-3) ribs, becoming weaker and more closely spaced around umbiHcus; periostracum thicker than in other species of the genus, and may flake off from dn- shells; ribs on base and rarely on dorsal surface mav bear periostracal bristles (some Persian Gulf and northern Oman specimens). Color: cream to ochre; usually with dark brown oblique (opisthocline) hnes or narrow stripes, numbering 6-11. often darkest or broad- est at suture and periphery; in northern Oman and Per- sian Gulf oblique lines may number up to 23 on last whorl, anastomosing to produce 8-15 spots at suture and periphen-; diagon;il stripes occasionally present only at suture and periphery (some East African specimens); di- agonal pattern only rarely entirely absent or developed only on spire (most specimens from Seychelles); often a single brown spiral line on early spire whorls; base un- pattemed or with small brown spots on ribs, occasionally forming radial fines; parietal callus, columella base and umbifical margin sometimes browaiish. Animal: Head-foot: front of head blackish or with nar- row black band only, separate black band across base of snout; tentacles with 2 longitudinal black lines; sides of foot with some black pigmentation or unpigmented. Pe- nis (Figures 102-105): filament short (20-30% total length), with elongate, mucronate tip, distinctly hooked or minutely hammer-shaped at very tip; base thicker, finelv wrinkled, with single large mamilliform penial gland (rarely absent) with elongate reservoir. Pallial ovi- duct: bursa present, apparently at posterior end of straight section. Radula {Figures 121, 122): cusps point- ed, major cusp of lateral and inner marginal teeth slight- ly rounded, central cusp ol rachidian tooth slightly apic- ulate; outer marginal tooth with 4 pointed cusps, out- ermost more rounded, becoming smaller toward mid- fine. DLstribution: H(i]>itat: intertidal rocks, pools and Tlwl- as.sodr lid roil Hats in a sheltered bay (Inhaca Island; R. N. Kilburn); barnacle zone on beach rock (Aldabra; J. D. Taylor); among barnacles in high eufittor;il, on surf beach (Masirah Island; J. D. Taylor); crevices in fime- stone, in upper eulittoral, on sheltered coast and in tidal lagoon (Ras al-IIadd; E. Glover); among baniacles on mangrove pneumatophores (Abu Dhabi; D. George); among Crassostrea (Kuwait; D. Jones). Ran^e (Fiffire 137): Zululand to Persian Gulf, Mauri- tius, Madagascar, Seychelles; excluding Red Sea. Records and material examined: South Africa: Mission Rocks. Zululand (NM). Mofambique: Inhaca Island (NM, IM); Benguera Island (NM); Mozambique Island (NM). Tanzania: Zanzibar (BMNH). Kenya: Tiwi Beach, 30km S Mombasa (BMNH, 2M. 2F). Madagascar: Tu- lear (MNHN); Maliajanga (IRSNB); Nossi-Be (USNM. ANSP MNHN. ZMA); Diego-Suarez (IRSNB). lie Ma- yotte (MNHN, IM). Mauritius: Grand Bav (BMNH, NM); Pointe Radeau, S of Roches Noires (NM); Pointe d'Azur (NM). Seychelles: Aldabra (BMNH); Baie Ter- nay, Miilie (BMNH, 2R); Anse la Mouche, Mahe (BMNH, IM, IF; USNM); Northwest Bay, Mahe (ANSP). Djibouti (MNHN). Yemen: Perim Island (MNHN); Little Aden (BMNH, 2F); Aden (BMNH, MNHN). Oman: Hamran. Zufar (BMNH); Sudh (BMNH); Sadh (BMNH); Kuria Muria Islands (ZMA); Masirah Island (BMNH. IF; ZMA. 2F); Sedli. Al Khav- sah (BMNH); Khor Hajar, Ras al-Hadd (BMNH. IM, 2F, IR); Bandar Kliayran (BMNH); Al Bustan, 3.2 km E Muscat (ZMA); Muscat (NM); N of Sharm (BMNH). Abu Dhabi: Merawwah Island (BMNH. 3M, IF). Qatar: Khor. Doha (BMNH). Kuwait (BMNH). Remarks: Although shape and coloration are highly variable in this species as a whole, some geographical patterns can be tfistinguished. In southern Arabia the spire is a little taller, the shoulder more rounded, the peripheral keel less pronounced, and spiral sculpture on the dorsal surface is weaker or even absent (Figures 33. 69. 73). In the Persian Gulf and northern Oman there is a tendency to flattening of the spire, and dexelopment of heaw carinae at the shoulder and peripher\-. while the dark obli(jue fines become more numerous (Figures 31, 71, 74). Specimens from Kenya and Zanzibar are more nigose. sometimes with regular racfial folds and slightlv crenulate peripheral flange (Figures 34. 68). In the flattened and weakly shouldered specimens from the Sevchefles (Figure 75) the fined color pattern is absent or occasionallv present on the spire only. In typical spec- imens from the Mascarene Islands. Madagascar and Mo- 9ambi(jue, the shoulder is scarcely marked and the color pattern is pronounced (Figures 30, 32). The shell of this species is most likely to be confused with tliat of/' (.s,st7( (see Remarks on that species), which Figures 114-116. Radulae of Peasiella halm. 114. Baten, Okinawa, Japan (flat view, shell D=3.2 mm). 115. Ma Liu Shi. Hong Kong (view at 45°. shell D = 2.2 mm). 116. Gu Leng Yu, Xiamen. China (view at 4.5° from side, shell D=1.8 mm). 117. Radula of paratvpe of Pcasidla fasciat a. Turtle Bay. Cape Ferguson, Queensland. Australia (BMNH 1996273; view at 45°, shell D = 2.1 mm). 118,'ll9. Radulae of PrasirUa mc/i. 118. Hunj;liada, Kgvpt (flat view, slid! D = 2.4 mm). 119. Hvirghada. Egvpt (view at 45°, shell D= :?,() luuii. 120, 121. Two views (flat and at 4.5°) of radula of AY;.vic//« inmiritiana. Khor Hajar. Ras al-Hadd. Oman (shell D=3.6 mm). Scale hars = 20 (Jini. D. G. Reid and Y.-M. Mak, 1998 Page 25 Page 26 THE NAUTILUS, Vol. 112, No. 1 122 123 124 -T^^X 100 ;jm Figures 122-124. Egg capsules of Pcnsiella species. 122. Pcasiclla fnsciata. Cape d'Aguilar, Hong Kong. 123. PeasieUa patida. Changi Point, Singapore. 124. PensU-Un Itahri. Ma Liu Shi, Tolo Harbour, Hong Kong. is endemic to tlie Red Sea; the two are only known to be sympatric close to the mouth of the Red Sea. Two other species, P. infracostata and P fuscopipcrota. can be found s\nitopically with P nmuiitiana, and all three have diagnostic penial shapes. The shell of P. infracos- tata is alwavs smooth abo\'e the periphery, and is thus easily distinguished from all ribbed and shouldered forms off matiritiana. However, examples of the latter from southern Oman and Yemen may lack ribs above the peripherv, and if the shoulder angulation is poorly developed mav be confused with P. infracostata. The ba- sal ril)s may then be useful; in P. mmiritiana the outer- most rib is the largest, but in P. infracostata the mid- basal rib is most prominent. Rare examples of P. fiisco- piperata which ha\'e spiral sculpture above the periphery- could lie confused with this species, lint lack the shoul- der angulation. ADDITIONAL NOTES ON OTHER SPECIES OF PEASIELLA PeasieUa roepstorffiana (Nevill, 1SS5) (Figures 142-145) Di.stribution: Additional records: Indonesia: Ambon (NN.ML). T;iiwan: Kcnding, Kaohsiang (BMNH); Yehhu, Taipei (BMNH). Japan: Kauuon Saki, Ishigaki, Rvuk-yu Islands (BMNH). Remarks: The range of this species is now extended from Hong Kong (Reid, 1989a, 1992) to T;iiwan and the .southernmost RvTikyu Islands (Figures 142, 144, 145), and also includes Ambon in Indonesia. A curious feature of this species is the apparent dimoiphism ol penial form; a mamilliform gland is present in material from Queensland, but not in specimens from Asia (Reid, 1989a). The absence of a penial gland has been con- firmed in m;it('rial from Ishigaki and .Xmbon. .Vdditional anatomical material has also been seen from Lautoka, Viti Levu, Fiji (BMNH); penial glands were found to be absent in two specimens, but in a third a nidimentary gland was present, with a cluster of subepithehal tissue but no reservoir or papilla. This supports the earlier sug- gestion that the peni;d dimorphism is indeed intraspe- cific. A similar case of geographic;d \ariation in presence or absence of a single penial gland has since been re- corded in Littoiina kasatka (Reid, 1996). The new Fijian material also supports the identification of the distincti\'e darklv tessellated shell from the eastern part of the range as merelv a color variant of this species. Sb; additional radulae examined (Hong Kong, Singapore, Tiiiwan, Ja- pan) each have 3 cusps on the outer marginal tooth, as recorded by Reid (1989a). The egg capsule of this spe- cies was described bv Mak (1995) from Hong Kong; it is nuich larger than those capsules described here, 360- 390 |xm in diameter, and similar in shape to that of P. hahei, although without the h\'o inner concentric rings on the upper surface. Pcasiclla conoidalis (Pease, 1868) (Figures 138-141) Remarks: AdtlitionJ anatomical materi;il of this species has been seen from Taiwan and Ishigaki, Ryuk)ii Islands (BMNH). The penial shape is the same as that illustrated b\- Reid (1989a), with a small and slender penial gland. Shells from these loc;Jities are often darker in cok)r than those found elsewhere, with brown dots (or occasionally fine lines) in the spiral grooves abcne the peripher)- and strong brown marks in the folds behveen the peripheral creiHilations (Figin-e 138). In some shells the pattern con- .sists of 1(>-17 dark browni axial stripes (Figiue 140). Peas- ieUa rocpstoiffjana ;ilso occurs at these localities, and shells of the t\\t> are ea.silv confused since both are veUovv, D. G. Reid and Y.-M. Mak, 1998 Page 27 ' i. 125 126 im ] I' 127 ''^■P .^' ^' i %' : ' s" ■ 128 ^ 129 130 Figures 125-130. Egg capsules oi' Ptnsirlla species. 125, 126. Peasiella fasciata. Cape d'Aguilar, Hong Kong. 127, 128. Peasiella pattiln. Chaiigi Point, Singapore. 129, 130. Pensiclln hahci. Ma Liu Shi, Tolo Harhour. Hong Kong Scale bar = 200 \xm. have radially plicate sculpture and peripheral crenulation.s (Figures 138, 141, 142, 145). They are separated by the following characters: in P. conoidalis the spiral grooves above the periphery usually contain sni;ill lirovMi dots rather than continuous lines and the periphenil lirovvni marks are weaker; die middle of the base of P. conoklalis shows 1-2 rows of small brown spots, whereas that of P. roepstoijfiana is unmarked but for dark blotches on the underside of die peripheral keel and puiple-tinged colu- mella; the wide unsculptured area surrounding the mn- bilicus is characteristic of P. roepstoijfiana: anatomically, the single mamilliform penial gland of P. conoklalis is lacking in P. roepstorffiana (note that these characters do not necessarily apply to the two species in other parts of their ranges, see Reid, 1989a). These two species are only occasionally found SMiipatriciillv elsewhere (e.g. Lizard Is- land, Australia; soutlieasteni Papua New Guinea; Ambon, Indonesia; some islands in Fiji and the Philippines), since P. conoidali.s has an oceanic distribution while that of P. roepstoijfiana is more continental in character. An un- identified Peasiella species from Chichijima, in the Oga- sawara (Bonin) Islands, illustrated bv Fukoida (1993), is a Page 28 THE NAUTILUS, Vol. 112, No. 1 small, worn example of this species (specimen now in BMNH). PeasieUa corwidalis has also now been recorded on die African mainland for die first time, from Tiwi Beach, Kenva (BMNH), where it occnrred v\ith P. infra- costata, P. fuscopipcrata and P. mauiitiona. Another new record is from Ambon, Indonesia (NNML). Pca.siclla lufiilcuta Reid, 19S9 Remarks: This species is now recorded from Singa- pore (BMNH), where it appears to be rare. One of the five specimens found was a female with a mature paUial oviduct as well as a small penis (see Remarks on P. habci and P. pattda). Two additional radulae examined each have 4 cusps on the outer marginal teeth, whereas Reid (1989a) found 3 cusps. PeasieUa petiti Le Renard, 1994 Remarks: This species from the Eocene of France was described and figured as P. minitta (Deshayes, 1824) bv Reid (1989a); this name is preoccupied, and the spe- cies was renamed by Le Renard (1994). DISCUSSION Snf:i,l, V.\RIATION Our reassessment of the taxonomy of PcasicUa has not altered the earUer conclusion (Reid, 1989a) that the shell characters of these littorinids are unusualK- variable. Elsewhere in the family, species with likewise plankto- trophic development do not normally show such con- spicuous variation (e.g. Reid, 1986a, 1996). The varia- tions in whorl outline and sculpture are most striking, since these characters are traditionallv used to diagnose httorinid species. For example, in P. injracostata and P. fuscopiperata the periphery may be strongly keeled or smoothlv rounded. In P. isscli and P. maitritiana the whorl profile may be almost flat or may develop a prom- inent carina at the shoulder In P. fuscopiperata, P. pa- tula, P. habei and P. mauritiami the surface above the periphery may be smooth or bear spiral ril)S. Shell thick- ness is also \ariable, ranging from solid and opaque to thin and translucent in both P. injracostata and P fus- copiperata. One sculptural character that is more useful for identification is the number and size of the ribs on the base. Manv littorinids arc conspicuously variable in shell color and pattern, and this is also the case in some PeasieUa species, notably P isseli. Nevertheless, we have found the number and arrangement of the obUque or spiral color bands t(j be one of the more useiul charac- ters for their identification. The cause and adapti\e sig- nificance of shell variation in non-planktotrophic littorin- ids has been the subject of much researcli, but that oi the plank-totrophic species is less well understood (re- views by McQuaid, 1996; Reid, 1996). Anatomy Throughout the family Littorinidae the shape of the pe- nis is known to be one of the most useful taxonomic characters, and probably plays a role in species recog- nition in nature (e.g. taxonomic accounts and reviews by Reid, 1986a, 1989b, 1996). It was therefore surprising that the prexaous account oi PeasieUa (Reid, 1989a) con- cluded that penial shape was similar throughout the ge- nus. All the six species for which anatomical materiid was then available showed a penis with a long filament and a single mamiHiform penial gland (although in P rocp- storffana the gland was found to be absent over much of the geographical range). The new material available to us in the present study has re\'ealed that s\nipatric species do in fact frequenth' show diagnostic differences in penial form, particularly in the relative sizes oi fila- ment and penial gland. The gross moipholog\' of the female reproductive sys- tem does not, however, show useful diagnostic charac- ters. The arrangement of the loops of the egg groove through the glands of the palfial oviduct is the same in all 11 species oi PeasieUa (as illustrated bv Reid, 1989a: figs. 57, 58, 65, 66). The copulaton bursa is apparently absent in six species, but this character is not useful for identification since it can be difficult to trace without histological preparation. Furthermore, in at least one ht- torinid, Littorina saxatilis. there is intraspecific variation in this feature, for the bursa is sometimes vestigial or absent in small, but mature, adults (Reid, 1996: 312). The egg capsules of four PeasieUa species have now been described (P habei, P. patula, P. fasciata, herein; P rocpstotffana in Mak, 1995). At least in P habei the form of the capsule is constant over a wide geographical range, although there is some variation in size (see de- scription of this species). These four species each show a unique capsule shape, and there is an almost twofold range of diameters among them. Similar interspecific variation is present in the genera Wxhlittoiina and Lit- toraria (reviews bv Bandel 6c Kadolsky, 1982; Reid, 1986a, 1989b; Mak', 1995), although in most cases intra- specific variation is poorly kiiovvni. The radial plication of the capsules of P patula and P. fasciata has hitherto been described onlv in Sodilittorina species In general, the radula of fittorinids is taxononiicallv useful for the diagnosis of generic groups (Reid, 1989b), lint not at the species level (e.g. Reid, 1986a, 1996). As in tlu' earlii'r studv, we have found the radula of PeasieUa species to be relatively uniform in the genus, showing interspecific variation onlv in the mmiber of cusps on Figures 131-135. Distribution maps tA l','/isirlln species, compiled from records listed in text. 131. PeasieUa iufnuosltitn. 132. PeasieUa fiisiopipmild 1.33. Pi-asiillii patula 1.34. Priisii-lhi fiisciata 1.3.5. Pitisiclla hahei. D. G. Reid and Y.-M. Mak, 1998 Page 29 Page 30 THE NAUTILUS, Vol. 112, No. 1 Figures 136, 137. Distrilnition nmp.s of Piasiella species, compiled t'roni records listed in te.\t. 136. Peasiella isseli. 137. Pensiella mnuritiana. Figures 138-141. I'casirUa ronoidalis. Kcndint;, Kaolisiaiii;, Taiwan (BMNH 199(i-59.S). 142-145. I'laswUa nvpstorffiana. 142. Kannon Saki. Isliiijaki. Japan (HMNd 1996:5991. 143. Ma Lin Slii, Honj; Kong (BMNH 1996400). 144. Yelilin, Taipei, Taiwan (BMNH 1996401). 14.5. Kciidini;, Kaolisianj;, Taiwan (BMNH 1996402). Scale bar = 2 mm. D. G. Reid and Y.-M. Mak, 1998 Page 31 tile outer marginal tooth, ami m the development oi ha- s;il denticles on the raehiilian tooth. Phvi,(ic;env and Bi()(:i'.()(;hai'ii\ The phvlogenetic relationships of PcasicUa within the Littorinidae remain uncertain. In the nioqihological phvlogenetic analvsis of Reid ( iy89h), the gcTius ap- peared as the sister-taxoii ot Maimcahngia in the con- sensus of all equallv parsimonious trees; however, no unique s\iiapomoq)hv tor these two taxa was found. An- atomicalK, thev are similar in their penial form (single mamilliform gland and closed sperm duct), pallial om- duct loops (although these are shghtlv more complex in Mainwaringia. Reid, 1986b) and egg capsule shape (compare Reid, 199():fig. 3j with Mak, 1995: fig. 2d), hut some of these resemhlances mav he plesiomoqihic. Thev are so different in their shell and opercular shape, rad- ular teeth and in the unique hermaphroditism oi Main- waringia, that a close relationship might seem unlikely. Nevertheless, prehminar\' molecular data offers some support for their sister-group relationship, and is cur- rentlv under investigation (R. H. Thomas, pers. comm.). Morphological features do not provide sufficient in- formation for a cladistic analysis at the species level, and in the absence of an une(juivocal sister-group the polar- ities of the few aviulable characters are doubtful. The five species confused by Reid (1989a) under the name P. infracostata (P. infracostata. P. fiiscopipcrata. P. pa- tula. P. hahci, P fasviata) are a possible clade, showing some similariU in shell form (tendencv to reduction of peripheral keel and spiral ribs, presence of spiral cokjr bands), and in radular cusps (4^8 cusps on outer mar- ginal tooth, except 3—4 in P. pattila: cf 3-4 in all re- maining species). Four probable pairs of sister-species can be suggested. PeasicUa infracostata and P. fuscopipcrata are closeK similar in shell characters; both lack a copulators' bursa, and radular cusps are similar. Their geographical distri- butions are knowm to overlap only between Mo^ambi(jue and Kenva, which might suggest an allopatric mode of speciation followed b\ limited secondarv contact. These two have strikingly dissimilar penial shapes, as would be predicted if this were either an isolating or species-rec- ognition mechanism (Reid, 1996). A second likely sister- species pair is P. isscli and P. mauritiana. Again these are similar in shell characters, but differ in penial shape. These are aliuost completel)' allopatnc; P isscli is re- stricted to the Red Sea, while P. mauritiana occurs wide- ly in the Indian Ocean and Persian Gulf, and thev over- lap only in the vicinih' of Aden and at the mouth of the Red Sea. While endemism in the Red Sea fauna is high in some groups, among mollusks it is generally low (Sheppard et al, 1992). During the substantial falls in sea level produced bv Pleistocene glaciation, the Red Sea became hvpersaline, and recolonization from the Arabian Sea has occurred only since the Holocene trans- gression. Speciation in the Red Sea mav therefore be a recent phenomenon. At present, the currents at the mouth of the Red Sea do not appear to prevent passage of plankionic lanae, but the cold upweUing on the southern Arabian coast during the summer months may cause a partial isolation of the Red Sea from the Arabian Sea and Indian Ocean (Sheppard rf al., 1992). The third likelv pair is P hahci and P. fasciata. vs'hich both belong m the infracostata group'. These are known to overlap only between Hong Kong and Taiwan, and show a shght but consistent difference in penial shape. Finally, P. co- noidalis mvX P tantilla mav be sister-species. Their rad- ula, o\iduct and penis are identical, shells are similar in sculpture, and both occur only on oceanic islands (Reid, 1989a). The former is widespread throughout the Indo- West Pacific, whereas the latter is restricted to the Ha- w;dian Islands, suggesting a case of speciation by pe- ripheral isolation. The species status of P. tantilla is based only on its distinct shell. The two are not biown to occur svmpatricallv and, assuming that penial shape is an isolating mechanism, there has presumablv been no selection for its divergence. ACKNOWLEDGMENTS This work was carried out while Y-MM was in receipt of a post-doctoral fellowship from the Croucher Foun- dation, Hong Kong. We thank E. Glover and J. D. Tavlor (BMNH), S-Y. Chan (Singapore), H.L. Strack (Dor- drecht, The Netherlands), Y. Shikancj, (Tokx'o) and E. Platts (Winchester, U.K.) for providing specimens. For assisting Y-MM with field collecting we thank K. S. Tan (National University of Singapore), Fu-xue Li (Xiamen University), C. K. Tseng (Institute of Oceanography, Qingdao), Huang Shong (National Taiwan Normal Uni- versity), Michael Ilin-kiu Mok (National Sun Yat-sen University, Taiwan), S. Yamato and S. Ohgald (Kyoto University, Shinihama), Y. Takada (Ishigaki Tropical Sta- tion) and all their colleagues. For providing loans of ma- terial we are grateful to P. Bouchet (MNHN), H. Dekker (The Netherlands), H. Fukiida (Tok-vo Metropofitan University), R. N. Kilburn (NM), I. Loch (AMS), R. Moolenbeek (ZMA) and ). Pickering (OUM). For their expert photography of shells we thank H. Tavlor and N. Hayes (BMNH). P. York (BMNH) kindly assisted with the photomicroscopy of egg capsules. We appreciate the helpful comments of two anon)mious referees. LITERATURE CITED Amio, M, 196.3. A comparative embrvologx' ot niariiie gastro- pods, witli ecological consideration. Joiinia! of tlie Shi- monoseki Uni\ersih' of Fisheries 12:229-.:i5S. Audouin, V. l.S2fi. Explication sommaire des pjanclies de mol- lus(jiies (fe I'Egvpfe et de la Syria, publiee par Jules Cesar Savigiiy. Description de rEg>pte. Histoire Natnreile 1(4): 7-56. ■ Bandel, K. 1974. Studies on Littorinidae from {W Atlantic The Veliger 17(2):92-114. Page 32 THE NAUTILUS, Vol. 112, No. 1 Bandel, K. and D. Kadolskv. 1982. Western Atlantic species of yodilittorina (Gastropoda: Prosobranchia): comparati\e morphologs' and its functional, ecological, ph\logenetic and taxonomic implications. The \'eliger 25(1):1— 42. Bauer, B., P. Fioroni, I. Ide, S. Liebe, J. Oehlniann, E. Stroben and B. \\'atermann. 1995. TBT effects on the female gen- ital system oi Littorina littorea: a possible indicator of tri- but\ltin pollution. Hvdrobiologia 309:15-27. Bosch, D. T, S. P Dance', R. G. Moolenbeek and P G. Oliver. 1995. Seashells of eastern Arabia. Motivate Publishing, Dubai. 296 pp. Bouchet, P. and F. Danrigal. 1982. Napoleon's Egyptian cam- paign (179.8-1801) and the Savignv collection of shells. The Nautilus 96(1) :9-24. Choe, B. L. 1992. Illustrated encyclopaedia of fauna and flora of Korea, Vol. .33: MoUusca (II). Ministry of Education, Republic of Korea. Choe, B. L. and J. K. Park. 1993. Se\en imrecorded mesogas- tropodous species (Gastropoda: Mollusca) from Korean waters — superfamilies Littorinacea, Tomacea, Rissoina- cea, and Cerithiacea. Korean Journal of MalacologN' 9:17- 26. Dautzenberg, P. 1929 .Molkisques testaces marins de Mada- gascar. Faune des Colonies Fran^aises 3:322-636. Franc, A. 19.56. Resultats scientifiques des campagnes de la Cah/pso. II.- Campagne 1951-1952 en Mer Rouge. 9. Mollusques marins. Annales de I'lnstitnt Oceanogra- phique 32:19-60. Fukuda, H. 1993. Marine Gastropoda (Mollusca) of the Oga- sawara (Bonin) Islands. Part 1: Archaeogastropoda and Neotaenioglossa. Ogasawara Research 19:1-86. Fuk-uda, H., Mashino, K. & Sugimura, T 1992. A review of the molluscan fauna of Yamaguchi Prefecture, Western Ja- pan. Yamaguchi Museum, Hahe, T. 1956. The floating egg capsules of the Japanese per- iwinkles (Littorinidae). Venus 19(2):1I7-121. Habe, T 1964. Shells of the western Pacific in color, \"ol. 2. Hoikiisha, Osaka, 2.33 pp. Habe, T 1984. Peasiella roepstorffiana (Nevill) and P oaidei (Montrouzier). Chiribotan 15:11-12. Higo, S. 1973. A catalogue of molluscan fauna of the Japanese Islands and the adjacent area. 61 p. Higo, S. and Goto, Y. 1993. A ,s\stematic list of molluskan shells from the Japanese Islands and the adjacent area. Em Mal- acological Publishers.Yao, Osaka, 693-1-148 pp. Issel, A. 1869. Malacologia del Mar Rosso. Pisa. Lamy, E. 1938. Mission Robert Ph. Dollfus en Egypte. VII. Mollusca Testacea. Memoires de I'Institute d"Eg\pte .37:1-89. Le Renard, J. 1994. Revision des mollusques paleogenes du bassin de Paris. I. Rectifications de nomenclature d'especes. Cossmanniana 3:.35-40. Mak, Y.-M. 1995 Egg capsule morpholog\- of five Hong Kong rocky shore littorinids. Hvdrobiologia .3()9:.53-.59. Mastaller, M, 1979. Beitriige zur Faunistik und (ikologie der Mollusken und Echinodermen in den Koralleiiritten bei Aqaba, Rotes Meer PhD thesis, Ruhr-Uni\ersitat Bo- th i mi, .344 pp. McQuaid, C. D. 1996. Biology of the gastropod family Litto- rinidae. I. Evolutionar)' aspects. Oceanography and Ma- rine Biology Annual Review 34:2.33-262 Mebill, J. C. and R. Standen 19()L Tlie Mollusca of the Per- sian Gulf Gulf of Oman, and Arabian Sea, as evidenced mainlv through the ((illections of Mr F, W, Tovvnsend, 189;3-19G0; with descriptions of new species. Proceedings of the Zoological Society of London 1901(2):327-460. Moazzo, P. G. 19.39. Mollusques testaces marins du canal de Suez, Memoires de I'Institut d'Egvpte 38:1-283. Mori, K., S. Nishihama and M. Tanaka, M. 1985a. Conimunih' structure of a rockT shore in Tsuji-shima Island, Amakiisa. III. The analysis of relationships between distribution ot organisms and micro-topographical conditions using small quadrat. Publications of the Aniakusa Marine Biological Laboratory 8:4.3-63. Mori, K., M. Tanaka and S. Nishihama. 1985. Community' structure of a rocky shore in Tsuji-shima Island, Amak-usa. II. Vertical distribution of dominant species and its zo- nation pattern. Publications of the Amak-usa Marine Bio- logical Laboratory 8:27—41. Nevill, G. 1885. Hand fist of Mollusca in the Indian Museum, Calcutta. Part 2. Indian Museum, Calcutta, 306 pp. Ohtsuka, S. and E. Yoshioka. 1985. A preliminarv' note on the pelagic eggs of marine invertebrates. Special Publications of the Mukaishima Marine Biological Station 1985:225- 2.35. Okaitani, T 1986. Colored illustrations of living things: mol- luscs. Sekaibuukasha Co., Tokyo. Oyania, K. and Y. Takemura. 1961. The molluscan shells. Vol 5. Resources Exploitation Institute, Tokyo. Pallary, P. 1926. Explication des planches de J. C. Savignv. Me- moires de I'Institiit d'Egvpte 11:1-1.38. Reid, D. G. 1986a. The littonnid moUusks of mangrove forests in the Indo-Pacific region: the genus Littorarin. British Museum (Natural Histon), London, xy-l-228 pp. Reid, D. G. 1986b. Mainwaringia Nevill, 1885, a littorinid ge- nus from Asiatic mangrove forests, and a case of protan- drous hermaphroditism. Journal of Molluscan Studies 52: 22.5-242. Reid, D. G. 1988. The genera Bembicium and Risclhypsis (Gas- tropoda: Littorinidae) in Austraha and New Zealand. Rec- ords of the Australian Museum 40:91-1.50. Reid, D. G. 1989a. Systematic revision of the Recent species of Peasiella Nevill, 1885 (Gastropoda: Littorinidae), with notes on the fossil species. The Nautilus 103(2):43-69. Reid, D. G. 1989b. The comparative morphologv', phylogenv' and evolution of the gastropod family Littonnidae. Philo- ■sophical Transactions of the Royal Society of London. Se- ries B ,324:1-110. Reid, D. G. 1990b. A cladistic phylogeny of the genus Litiorina (Gastropoda): implications for evohition of reproductive strategies and for classification. Hydrobiologia 193:1-19. Reid, D. G. 1992. The gastropod family Littorinidae in Hong Kong. In: Morton, B. (ed). Proceedings of the Fourth International Marine Biological \\"orkshop: The marine flora and fauna of Hong Kong and .southern China III. Hong Kong University Press, Hong Kong, pp. 187-210. Reid, D. G. 1996. Systematics and Evolution oi Litiorina. Ray Society, London, x -1-463 pp. Reid, D. G. and J. B. Geller 1997 A new ovoviviparous species of Tectarins (Gastropoda: Littonnidae) from the tropical Pacific, with a molecular phvlogeny of the genus. Journal of Molluscan Studies 63(2):207-2.33, Rosewater, J. 1972. The family Littorinidae in the Indo-Pacific. Part II. The subfamihes Tectariinae and Echinininae. Indo-Pacific Mollusca 2:507-5,33. Savignv, J. C. 1817. Description de I'Egvpte, Histoire Natu- relle. Planches, \ol. 2. Imprinierie Royale, Paris. Shep|iard, C , A, Price and C. Roberts 1992. Marine ecologv D. G. Reid and Y.-M. Mak, 1998 Page 33 of the Arabian region. Patterns and processes in extreme tropical enxironments. Academic Press, London, 359 pp. Tanaka, M., K. Mori. S. Nojima, T. Kikuchi, T. Shibata, T. Ni- shino and K. Omori. 1985. Communitv' structure of a rockv' shore in Tsuji-shima Island, Aniakusa. I. Horizontal and vertical distribution pattern of common animals. Pub- lications of the Amakusa Manne Biological Laborator\- 8: 1-26. Tillier, L. and A. Ba\av. 1906. Les moUusques testaces du canal de Suez. Bulletin de la Societe Zoologique de France 30: 170-181. Tokioka, T. 1950. Droplets from the plankton net V. New names for egg capsules of littorinid gastropods. Publica- tions of the Seto Marine Biological Laborator\ 1:151-152. Tryon, G. W. 1887, Manual of conchology. Vol. 9, Philadelphia, 488 pp. Turton, W. H. 1932. The marine shells of Port Alfred, South Africa. Oxford Uni\ersit)' Press, O.xford, .331 pp. Viader, R. 1951. New or unrecorded shells from Mauritius and its dependencies. Mauritius Institute Bulletin 3:127- 153. Yamamoto, G. and T. Habe. 1962. Fauna of shell bearing mol- lusks in Mutsu Bay. Bulletin of the Biological Station of Asamushi, Tohoku University 11:1-20. THE NAUTILUS 112(l):34-4(), 1998 Page 34 Donald Richard Moore (February 16, 1921-October 19, 1997): Biographical Sketch, Zoological Taxa, and Bibliography Jose H. Leal The Bailev-Matthews Sliell Museiini 3075 Sanibel-Capti\u Road Sanibel, FL 33957 USA leal@water.net Donald R. Moore was born in West Palm Beach, Flor- ida, son of Walter R. and Estelle K. Moore. He attended elenientar\- school in Venice, C^alifornia and Hobe Sound, Florida, and high school in Stuart, Florida. Growing up in Florida provided Don with, in his words, "plentv of sea, swamp, lagoon, and woods to roam in". He joined the U.S. Marine Coq^s in September 1941. Don's love for shells and marine life consoUdated after his stay in the Central and Southwest Pacific, something that happened with so many young naturahsts who fought the Second World War in that part of the world. Don took his dive mask witli him to the Pacific, and dived as much as events would permit, at Midway, New Hebrides, and the Solomon Islands. After receiving an honorabk- discharge in October 1945, Don entered the undergraduate program at the Universitv' of Miami in June 194S, and graduated in Feb- marv 1954 with a Bachelor of Science degree in Zoology. He joined the Graduate School of the Universit\' of Mi- ami shortly thereafter, aiming for a M.S. degree in Ma- rine Science. In the summer of 1954 Don traveled to Houston, Texas, taking on a temporaiy assignment with the Exploration and Product Research Dixdsion of Shell Oil Company as field biologist. A job offer as Research Scientist at the Institute of Marine Science, University of Texas, caused Don to put his academic efforts tem- porarily on hold. From Texas he moved to Ocean Springs, Mississippi, to join the staff at the Gulf Coast Research Laboratory, working on oyster culture with Gordon Gimter as an Assistant Marine Biologist It was there that he published his first scientific article (Moore, 1956). Taking advantage of his proximity to the Univer- sitv of Southern Mississippi camp\is in Hattiesburg, he enrolled in the graduate program at that institution with Gordon Counter as his advisor, earning the Master of Sci- ence degree in June 19fi(). In September of 1960, he again entered the Graduate School at the Institute of Marine Science (now the Ro- .seustiel School of Marine and Atmospheric Science, RSMAS), University of Miami, graduating under the guidance of Clilbi'rt L. Voss on June 7, 1964, with a Ph.D. degree in Marine Science. His Ph.D. dissertation revised the systematics of western Atlantic \itrincllid gastropods. Don once told me of his first meeting with the great Henry Pilsbry. In the early stages of his dis- .sertation work, Don visited Pilsbrs at the Acadeins of Natural Sciences in Philadelphia. In Pilsbry's office, he brieflv introduced himself and his research goals. After hearing Don's desire to revise the western Atlantic Vi- trinellidae, Pilsbrv' offered his hand, saving, "Here, I sa- lute a hero! ". His unpubli.shed dissertation remains the best source of information about this difficult faniilv of gastropods. From the historic perspective of the entl of the twen- tieth centur)', the timing of Don's Ph.D. defense could not have been better: the "golden age" of tropical west- ern Atlantic marine science, centered at the lab in Mi- ami was under full sail in the mid-196()s. Don was hired in 1964 as an Assistant Professor of Marine Science at that institution. There he shared an extremely produc- tive academic environment with marine biologists such as (Gilbert Voss, Ravmond Manning, Frederick Bayer, C. Richard Robins, and Hillary B. Moore, as weD as with geologists of the caliber of Robert Ginsburg and Cesare Emiliani. During the sixties, Don cniised extensively on the Institute's research vessels in the Caribbean Basin, northern South America, Brazil, Straits of Florida and the B;iliamas, and the eastern seaboard of the U.S. In 1971 Don was promoted to Associate Professor of Ma- rine Geology, remiiining in that position until he retired to become Professor Emeritus in 1984. Don continued working and participating in students' committees until the dav of his death. Don showed great appreciation of other cultures anil was proud of his travels and abilitv to read and ,speak several foreign languages. Among his many travels and professional visiting appointments, he v\'as a visiting sci- entist at the Institnto de Investigaciones de Punta Betin in Santa Marta and taught for Sea Grant in Cartagena, both in Colombia. In 1980, Don worked at the Natural Histon Museum in Basel, Switzerland, by invitation of the Swiss National Science Foundation. An active par- ticipant in the American Miilacological Union, he sen'ed as Conniillor-at-Large of that organization in 1968-69, Vice-President in 1973-74, and Presulent in 1974-75. Following an invitation by then-editor R. Tucker Abbott, l^on joined the first group of consulting editors of The Naiifilit.s in julv 1972, a position he retained for the rest of his life (l')(in's first three peer-reviewed articles were piiblislicd m rlir Saiitilus). He was also a member and strong supporter ol the Miami Geological Society, serv- in<' as tluir Vice-President in 1967, President in 1968- J. H. Leal, 1998 Page 35 Young Donald R. Moore in 1954, suneving Gull ot Mexico niolliisks on the coast ol Texas for Shell Oil Co., as featined on the cover of Shell News Magiizine {Anonviiious, 1955). 69, and Secretai-v in 1977-78. He wa,s a .strong .snpporter of Unita.s Malacologica a.s a global organization, and played a key role, as a Scientific Committee member, in the formation of the Committee for Latin American Congresse.s in 1990. Don Moore sened on more than one hundred Uni- versit)' of Miami graduate .student committees, niiiinly at RSMAS, but also at the University of Miami's Depart- ment of Biolog) and Department of Geological Sciences, on the Cora! Gables Campus. He was committee chair or co-chair for the students: Richard S. Houbrick (M.S.), Jose H. Leal (Ph.D.), John Meeder (Ph.D.), James Quinn, Jr. (Ph.D.), Nidia Homer (M.S.). Don iilso was a committee member for the following malacologists: George Darcv (M.S.), Thomas Brackoniecld (M.S. and Ph.D.), Charles DAsaro (Ph.D.), Steven He.ss (Ph.D.), Edward J. Petuch (M.S. and Ph.D.), James Quinn, Jr. (M.S.), Ronald Toll (Ph.D.). Both as mv personal friend and academic advisor, Don was always present and supportive, in good and bad times. A large portion of my personal librarv' comprises used books and reprints that were gifts from Don (and I know the same to be tnie for many of his students) An a\id reader and staunch book"\vorm, Don used to spend a great deal ol his time in bookstores and book k ■ ■'• ' -J Dr Donald R. Moore m 1989, Coral Cables, Flonda Photo courtesN RSMAS Archi\'es. lairs. He also used to gauge the merit ot small towns by the presence or absence of good second-hand book- stores. Although most of his scientific producti\itv took place in the 1960s and 1970s, Don was allowed to retain for hfe his office in the North Gro.sxenor Building at RSMAS. His office doors were always open, a s\nnbolic depiction of his view of the world and of the special affection and warmth he had for students, no matter how inexperienced, or how distant their homelands. Don's immediate family survive him: his wife C\iithia is Researcli Associate with the Division of Marine and Atmospheric Chemistry- at RSMAS, and son Walter is Systems Administrator with Eckerd College at St. Pe- tersburg, Florida ZOOLO(;iCAL TAXA Abbreviations im repositories of Wpe material: AMNH American Museum of Natural Historv, New York ANSP The .Academv of Natural Sciences, Philadel- phia BMNH The Natural History Museum, former British Musemn (Natural Histoi-y), London, En- gland CNMS C^olombian National Museum of Science, Bo- gota, Cokjmbia DMNH Delaware Museum of Natural Histon, Wil- mington FMNH Field Museum of Natural Historv, Chicago FSBC Florida Marine Research Laboratorv, St. Pe- tersburg Page 36 THE NAUTILUS, Vol. 112, No. 1 MNHN Museum national d'Histoire naturelle, Paris, France MNRJ Museu Nacioniil, Rio de Janeiro, Brazil (also MNHSL) MCZ Museum of Comparative Zoolog\\ Harvard University', Cambridge MORG Museu Oceanografico, Funda^ao Universida- de do Rio Grande, Rio Grande, Brazil NNM Nationaal Natuurhistorisch Museum, Leiden, The Netherlands (formerly Rijksmuseum van Natuurlijke) UF Florida Museum of Natural Historv, Univer- sity of Florida, G;iines\ille UMML Marine Invertebrate Museum, Rosenstiel School of Marine and Atmospheric Science, Uni\ersit\ ol Miami, Miami USNM National Museum of Natural History, Smith- sonian Institution, Washington, DC Family-Level Taxon CYCLOSTREMELLIDAE Moore, 1966, Bulletin of Marine Science, 16(.3):4S0— 4S4 (diagnosis on page 4S1). For Cij- clostremella Bush, 1897. Robertson (1973) indicated that this genus belongs to the PyTamidellidae. Ponder & Waren (198S) included the Cvclostremellinae as as a subfamily o) P\TaniidelIidae. Genus-Level Taxon Dimijella Moore, 1969, Journal de Conchvlioloepe 107(4):137- 141 (diagnosis on page 137). T\pe-species: Dinu/rlhi star- cki Moore, 1969, by nionotvpy. Dinividae. Species-level Taxa All type-specimens collected by D. R. Moore except where otherwise indicated. bulli.si, Linckia Moore, I960, Bulletin of Marine Science of the Gulf and Caribbean I0(4):414-4I6, fig. I. Holo- type: USNM E-8I20, off Guiana, South America, 8"15'N, 58"I7'W, 72 to 91 m. (=Lmckia nodosa Perrier, 1S7.5, fide Downey, 1968) (Echinodermata, Asteroidea, Ophichasteridae ) . calcicola, Ntictda Moore, 1977, The Nautilus 91 (4): 120- 121, figures 1-3. Holotype: USNM 7.585.3.5, Chancanab [Chankanaab] Lagoon, Cozumel, Quiutana Roo, Mexico, 2 m; All paratypes from type locafity: USNM 758537, 7 paired specimens -I- 3 single valves; MCZ 288445, 3 paired specimens + 4 single valves, from tvpe localit\'; UMML 28.2812, 3 p;ured specimens + 4 single valves, from type locality; AMNH 183857, 10 single valves; ANSP 344387, 5 paired specimens + 4 single valves; MNHN unnumbered, 5 paired specimens. 2 left valves, 1 right valve; DMNH 120581, 6 paired specimens + 1 single valve; BMNH 1977089, 3 paired specimens + 3 single valves; NNM 55302, 4 paired specimens + 4 sin- gle valves, all dry. Nucididae. colombiana, Alvania (Alvania) Romer and Moore, 1988, The Nautilus 102(4): 131-133, figures 1-5. Holotvpe USNM 859339, off west Florida, 29°35'N, 87°20'06'"W. 107.3 m; Paratvpes: USNM 859340, 1 parat\pe, t\pe lo- cality; FSBC 3311.3-114, 2 paratvpes, off west Florida, 28°24'N, 8.5°15'06"W. 164.6 m; MCZ 297220, 1 para- type, off Puerto Rico, 17°53'24"N, 66°35'10"\\' 221 m; MCZ 297219 and UMML 30.8349, 2 parat\pes, off Mi- ami, Florida, 25°47'N, 80°01'30"W, 137 'm; UMML .30.8350 and CNMS ICN-MHN(MO) .523, 2 paratvpes, off Miami, Florida, 2.5°46'.30"N, 80°00'08"W, 76.81 m; CNMS ICN-MHN(MO) 522, 1 parat%pe, off Caribbean coast of Colombia, 09°52'.3.5"N, 75°47'25"W, 72 m. Ris- condijlum, Caecum Moore, 1969. Bulletin ol Marine Sci- ence 83(l):26-28, figure 1. Holotype: USNM 679.348, Pavardi Island, near Colon, Atlantic coast of Panama. Parat\pes: USNM 679348, 1 paratvpe. East Flower Gar- den Bank, Gulf of Mexico, 27°54'30"N, 93°35'45"W. cor- al debris, 1 8.. 3-23. 8 m. Caecidae. cynthiac, Condylonucida Moore, 1977, The Nautilus 91(4): 12.3-124, figures 6-7. Holotype: USNM 7.58.534, about 800 m west of outer reet (lagoon), Courtown Cays (Cayos del E.S.E.), off Nicaragua, western Caribbean Sea, 7.5 m. Paratvpes: USNM 7.58.538, 5 paired speci- mens + 1 broken right valve + 1 fragment, Courtown Cavs, off Nicaragua, 1.5 m; ANSP 344388, I p;iired spec- imen + 1 right valve, Courtown Cays, off Nicaragua, 1.5 m; DMNH 120580, 1 paired specimen + 1 left vidve; MCZ 288444, 1 piiired specimen + 2 left valves, Cour- town Cavs, north end, 2 m; UMML 28.2811, 1 paired specimen + 1 left v;ilve, Courtown Cavs, outer reef, 1 m, Nuculidae. rsperanza, Thala Leal and Moore, 1993, The Nautilus 107(2):58-62, figures 1-6. Holotype: USNM 860280, Playa Esperanza, 40 km west of San Juan. Municip;Uity of Manati, northern Puerto Rico .\11 parahpes from tvpe localit%-: USNM 860281, parat^pes 1-4; UF 193.382-.i83, paratvpes .5-6; ANSP 391938, paratype 7; AMNH 232313, paratvpe 8; MCZ .302588, paratvpe 9; MNHN unnumbered, paratype 10; FMNH 2233S8, paratype 11; UMML 30.8375, paratype 12; USNM 860262, paratypes 1.3-16. Costellariidae. fioridana, Macromphalina Moore, 1965, The Nautilus '78(3):7.5-76, plate 7, figure 1-3. Holotype: USNM 6.36610, Madeira Beach at 150"' Avenue, Saint Peters- burg, Floritla; paratvpes: ANSP 295621, 1 shell, type lo- cality; UMML 302773, 1 shell, east of Sokfier Key Bis- cavne Bay, Florida, I m. Vitrinellidae. iiidiscrcta. Rissoiua Leal and Moore, 1989, Bulletin of Marine Science 45(I):I39-147, figures 1-12. Holotype: MNRJ 5760. Ponta Verde Beach, Maceio, Alagoas State, J. H. Leal, 1998 Page 37 Brazil, 09°38'S, 35°44'W, beach at low tide. Paratxpes: MNHSL 6025, paratopes 1-4, t\pe Iocalit\-; MORG 4164, paratvpe.s 5-7, t\pe locality; MNHN unnumbered, paratopes 8-11, t\pe localit\'; USNM 859335-338. para- t\pes 12-15, t\pe loc;iliK; MNRJ 5761, paratxpes 16-18, type localitv-: BMNH l'9S8.()43, paratvpes 19-22, type localit\-; MORG 25463, parat\pe.s 23-24, Atol das Rocas, Brazil', ()3°52'S, 33°4y'W; MORG 25464, parat\pe,s 25- 28, Abrolhos Reef Complex, B;ihia, Brazil, ■l7°58'S, 38°42'W; UMML 8347-48, parat\pes 29-30, Jaragua Beach, Maceio, Alagoas State, Brazil, 09°38'S, 35°44'W. Rissoidae. insiilaniin. Caecum Moore, 1970, Bulletin of Marine Science 20(2):368-373, figures lA-B, Holot>pe: USNM 679178, open sandy bottom at head of Lesser Lameshur Bav, St. John, Virgin Lslands, 1.5 m. All paratypes from type localitv: USNM 679179, 12 paratypes; ANSP 315477. 10 ■ parat\pes; MCZ 271.305. 11 paratxpes; AMNH 147965, 10 paratypes; UMML 30.3203, 14 para- types; "more than a hundred paratypes ... kept bv the author". Caecidae. mai/a. Condi/lonitcula Moore, 1977, The Nautilus 91(4): 124-126. figures 4-5. Holot\pe: USNM 758536, Chan- canab [Chankanaab] Lagoon, Cozumel, Quintana Roo, Mexico. 2m. All paratypes from type locaht)': USNM 758536, 7 paired specimens + 1 right valye, type local- ity; MCZ 288443, 2 p;ured specimens -I- 1 right yalye; ANSP 344389, 3 paired specimens + 1 left yalye; AMNH 183858, 2 paired specimens -I- 1 left yalye; DMNH 120579, 3 paired specimens + 2 single valves; FMNH 198080, 1 paired specimen. Nuculidae. semipuncttis, Solariorhis Moore, 1965, The Nautilus 7S(3):77-78, figures 1-3. Holot>pe: USNM 636309, northwest Campeche Bank, Mexico, 18 m, muddy bot- tom. Paratype: UMML 30.2774, 1 paratype, Baie de Aquin, Haiti, beach drift. Vitrinellidae. starcki, DumjicUa Moore, 1969, Journal de Conchvliol- ogie 107(4): 138-139, one plate with two figures. Holo- tvpe: USNM 678363. complete shell with txvo larxal specimens in brood chamber, Cozumel Island, Mexico. All paratypes from type locality: USNM 678364, 1 paired specimen + 1 detached upper valve; ANSP 312218, 1 paired specimen -I- detached upper valve; BMNH 1967.6171, 1 paired specimen + another with broken lower valve; MNHN 1967.226, 1 paired specimen -I- de- tached upper valve; MCZ 257727-257726, 1 paired specimen with brood chamber in upper valve + de- tached upper \alve; UMML 28.784, 1 paired specimen + another with both v;ilves broken + three small de- tached upper valves. Dimvidae. tcxana. Vitrinclla Moore, 1965, The Nautilus 78(3);76- 77, plate 7, figures 4-6. Holotype: USNM 636311, Mus- tang Island, near Port Aransas, Texas. Parat\pes: ANSP 295622, 1 paratope, t>pe locality; USNM 636312, 1 para- type, type localitv'; UT 1015, t\vo shells, Cline's Point, Port Aransas, Texas; UT 1016, 1 paratvpe, Ferr\' Land- ing, Port Aransas, Te.xas; UMML 30.2775, 1 paratype. Port Aransas. Texas; UMML .30.2776. 1 paratvpe. Port Aransas, Texas. VitrinelUdae. GRADUATE THESES Moore, D. R 1960. Tlie marine and brackish water MoIIusca of the Mississippi Gulf Coast. ,Master of Science. Univer- sity of Sontheni Mississippi. 116 pages. Moore, D. R, 1964. The faniilv N'itnnellidae in South Flonda ant! the Gulf ot Mexico. Doctor ot Philosophy, Uni\'ersity of Miami, xi -I- 235 pages, 35 figures, -I- vitae. [7 June] PUBLICATIONS This hsting includes peer-re\iewed articles as well as Don's reports and contributions to several scientific or- ganizations. Also included are the extended meeting ab- stracts pubUshed up to 1980 bv the American Malaco- logical Union (AMU) in its Annual Reports and Bulletin. References are hsted in chronological order. Moore. D, R 1956. Observations of predation on echinoderms h\ three species of Cassididae. The Nautilus 69(3);7.3-76. [il Fehniarv] Moore, D. R. 1957. A note on Cuna daUi. The Nautilus 70(4): 12;3-125, pi. 8, figs. .5-4. [29 April] Moore, D. R, 1958. Additions to Texas marine MoIIusca. The Nautilus 71(4):124-129. [24 April] Moore, D. R. 1959. Notes on Blanquilla Reef the most north- erlv coral reef in the western Gulf of Mexico. Publications of the Institute of Manne Science 5:151-1.55 ["Decem- ber. 195S-] Moore, D. R. and H. R. Bullis. 1960. A deep-water coral reef in the Gulf of Mexico. Bulletin of Marine Science of the Gulf and Caribbean 10(1): 125-128. figures 1-2. [March] Moore. D. R. 1960. Linckia hnlhsi, a new asteroid from the northeast coast of South America. Bulletin of Marine Sci- ence of the Gulf and Caribbean 10(4):414-416, fig 1. [December] Moore, D R 1961a. The manne and brackish water mollusca of the state of Mississippi. Gulf Research Reports [Ocean Springs, Mississippi] 1(1): 1-58. [April] Thomas, LP., D. R. Moore and R. C. Work. 1961. Effects of hurricane Doniia on the turtle grass beds of Biscayne Bay, Florida. Bulletin of Marine Science of the Gulf and Ca- nbbean 11(2): 191-197, figs. 1-2 [June] Moore, D. R. 1961b. The occurrence of Stephanoscijphuscor- nifonnis Komai (Sc\phozoa) in the western Atlantic. Bul- letin of Marine Science of the Gulf and Caribbean 11(2): 319-.320. [June] Moore, D. R. 1961c. The molluscan sea grass communities in Biscayne Bav. American Malacological Union Annual Re- ports for 1961 [Bulletin 28], page 20, The American Mal- acological Union Twenty-eighth Annual Meeting, Wash- ington, DC, June 19-23, 1961. [Abstract] [1 December] Moore, D. R. 1962a. Notes on the distribution of the spinv lobster Fanuhnis in Florida and the Gulf of Mexico. Crus- taceana 3(4):3L8-319. [March] Moore, D. R and G Gunter 1962. Notes on the Pliocene molluscan fauna from one site in the western Florida Ev- erglades. Bulletin of Marine Science ot the Gulf and Ca- ribbean 12(l):66-72. [March]. Olsson, A. A. and D. R. Moore. 1962. A neglected west At- Page 38 THE NAUTILUS, Vol. 112. No. 1 lantic stronib. The Nautilus 75(4):127-128, pi. 11, figs. 1- 3. [9 April). Moore, D. R. 1962b. Occurreuce and distribution o( Netnopsis hachei Agassiz (Hvdrozoa) in the nortlieni Gulf of Mexico. Bulletin of Marine Science of the Gulf and Caribbean 12(3):399-402. [September]. Moore, D. R. 1962c. The systematic position of the famiK- Caecidae. American Malacological Union Annual Report for 1962 [Bulletin 29], page 5, Twenty-ninth Annual Meet- ing of the American Malacological Union, St. Petersburg, Florida, 31 ]uly-.3-August 1962. [Abstract] [1 December"]. Moore, D. R. 1962d. The systematic position of the family Caecidae (MoUusca: Gastropoda). Bulletin of Marine Sci- ence of the Gulf and Caribbean 12(4):69.5-7()1, figs. 1-3. [December] Moore, D. R. 1962e. An ophiurian shoal on the Mississippi coast. Quarterly Journal of the Florida Academy of Sci- ences 25(l):70-72. [23 August] Moore, D. R. 1963a. Turtle grass in the deep sea. Science 1.39(.3.560): 12.34-12.3.5. [22 March] Moore, D. R, 1963b. Mecoliotia. a gastropod genus new to Florida. Bulletin of Marine Science of the Gulf and Ca- ribbean 1.3(1): 7:3-76, fig. 1. [March] Moore, D. R. 1963c. Distribution of the sea grass, Thalassia, in the United States. Bulletin of Marine Science of the Gulf and Caribbean 13(2):.329-.342. [June] Moore, D. R. 1964. The evolution of the Mesogastropoda, American Malacological Union Annual Reports for 1964 [Bulletin 31], pages 17-18, Thirtieth Annual Meeting of the American Malacological Union, New Orleans, Loui- siana, 21-24 July 1964. [Abstract] Hoffmeister, J. E., J, 1. Jones, J. D. Milliman, D. R. Moore and H. G. Multer. 1964. Field guidebook to the hving and fossil reef types of South Florida Miami Geological Society, Coral Gables, 2S pages, [Noxember] Moore. D, R. 1965a. New species of Vitrinellidae from Gulf of Mexico and adjacent waters. The Nautilus 78(3):7.3-7S, plate 7, figs. 1-6; plate 8, figs. 1-3. [25 Januar\'] Moore, D. R. 1965b. The Marquis de Folin and his work on the Caecidae. American Malacological Union Annual Re- ports for 196.5 [Bulletin .32], pages .3.5-36, Thiity-first Meeting of the American Malacological Union, Staten Is- land. New York, 20-23 JuK. 1965.'[Abstract| [1 Decem- ber] Moore, D. R 1966a. A note on Pseudoparastrophia. Journal de Conciiyliologie 105(2):57-61, figs. 1-2. [1 Marclil Moore, D. R. and K. J. Boss. 1966. Records for I'liralxu-nid squiUimi. The Nautilus 80(1 ):.34-.35. [6 Julyj Moore. D. R. 1966b. The Cyclostremellidae, a new lamiK ot prosobraiicli mollusks. Bulletin of Marine Science 16(3): 480-484, figs. 1-6. [September] Moore, D. R 1966c. Systematics and zoogeography of the Ctiloceratidae. American Malacological Union Annual Report for 1966 [Bulletin 33], page 40, Thirty-.second Annual Meeting of the American Malacological Union. Chapel Hill, North Carolina, 22-27 August 1966. [1 December] Boss, K. J. and D. R. Moore. 1967. Notes on Malleus {Pari- rruillexis) amdeanus (d'Orbignv) (Mollusca, Bi\'aKia). Bul- letin of Marine Science 17(1 1:8.5-94, 2 figs. [Marcii] Bock, W. D. and D. R. Moore. 1968. A commensal relation- ship between a foraininifer and a mollusk. Gulf Research Reports [Ocean Springs, Missi,ssippi] 2(3):27.3-279, plate 1. [No\ember| Moore, D. R. 1968. Obser\'ations on the Caecidae. American Malacological Union Annual Report for 1968 [Bulletin 35, page 39, Thirty-fourth Annual Meeting of the American Malacological Union, Corpus Christi, Te.xas, 15-19 July 1968. [Abstract] [27 December] Moore, D R 1969a. A new Caecum from the tropical western Adantic. Bulletin of Marine Science 83(1):26-2S, fig. 1. [July] Bock, W. D., D. R. Moore, A. C. Newmann and P. R Supko. (organizers). 1969. Late Pleistocene geology in an urban area. Third annual field trip of the Miami Geological So- ciety. Miami Geological Societ\'. Miami, 26 pages. [July] Moore, D. R. and P R. Supko. 1969. Introduction. In: Bock. W. D., D. R. Moore, A. C. Newmann and P R. Supko. (organizers). Late Pleistocene geolog\' in an urban area, pages 1-2. Third annual field trip of the Miami Geological Society. Miami Geological Society, Miami. [Jul\'] Moore, D. R. 1969b. Cijclostrema miranda Bartsch, a syn- onym of Tormis subcarinatus (Montagu). The Vehger 12(2): 169-170. figs. 1-3. [1 October] Moore, D R. 1969c. Systematics, distribution, and abundance of the West Indian micromollusk Rissoina catesbijana d'Orbigny. Nineteenth Meeting of the Gulf Coast Asso- ciation of Geological Societies, October 29-November 1, 1969. Transactions [of the] Gulf Coast Association of Geo- logical Societies [Tallahassee, Florida] 19: 42.5-426. [Oc- tober] Moore, D. R. 1969d. A new genus and species of Dinnidae from the Caribbean coast of Mexico, Journal de Concliv- liologie 107(4):137-141, 1 plate. [30 December] Moore, D. R. 1970a. The subgenus Armata de Folin, of the genus Caecum. Journal de Conchvliologie 108( 1 ):.3-.5. [31 March] Moore, D, R. 1970b. A new Caecum from Puerto Rico and the Virgin Islaiuls. Bulletin ot Marine Science 20(2):.36S- .373, figs, 1-2, [June] Moore, D. R, 1971a. Cochliolcpis parasitica, a non-parasitic marine gastropod. American Malacological Union Annual Reports for 1970 [Bulletin 37], page 80, Thirty-sixth An- nual Meeting of the American Malacological Union, Key West, Florida, 16-20 July 1970. [18 Febniarv] Moore. D. R. 1971b. A deep water Omalofii/ra in the western Atlantic, The Nautilus 84(4): 11.3-1 17, fig, 1. [26 .\pril] Bock, W, D, and D R. Moore. 1971. The Foraminifera and micromollusks of Ilogstv Reef and Serrana Bank and their paleoecological significance. Transactions of the Fifth Ca- ribbean Geological Conference, Geological Bulletin [(Queens College Press, Flushing, New York] No. 5, pages 14.3-146, 2 figs., 2 tables. [May] Meeder, J. F, and D R Moore 1971. The extension of range of Beiilulinia carih}>ea Edmunds to Brazil and Panama (Molliisa [sic]; Gastropoda). Caribbean Journal of Science 11(.3-1):1.59-I61, figs. A-B. [September-De- cember]. Moore. D R. 1971c. Disccners' of the Carolina marsh clam, Poli/mesoda caroliniana (Rose), a supposeil Florida dis- junct species, in Everglades National Park. Florida. Gulf Research Reports [(.)cean Springs, Mississippi! 3(2): 26,5- 277, figs. 1-.3. table 1. [December] Moore. D. R. 1972a. Ecological and .SN.stematic notes on Cae- cidae collected at St. Croix, Virgin Islands. Bulletin of the American Malacological Union for 1971, page 11, Thirt\- se\'enth Annual Meeting of the American Malacological Union, Cocoa Be;ich. Florida, 15-19 |ul\ 1971 [Febman] J. H. Leal, 1998 Pase 39 Moore. D. R. 1972b. Cochliolepis parasitica, a nonparasitic marine gastropod and its place in the V'itrinellidae. Bul- letin of Marine Science 2iZ(l): 100-1 12. figs. 1-6. [March] Moore. D. R. 1972c. Coral reefs, new and old. Pages 1-6, Field trip. 2-3 December 1972, American Quateniarv' As- sociation, [2 December] Moore, D. R 1972d. Ecological and s\stematic notes on Cae- cidae from St. Croix, U.S. Virgin Islands. Bulletin ot Ma- rine Science 22(4):881-.899, figs. 1-12. [with abstract in Spanish] [December]. Moore, D. R. 1973. Mollusks from a small landlocked Mexican lagoon. Bulletin of the American Malacological Union tor 19V2. pages 5-6. Thiit\ -eighth Annual Meeting of the American Malacological Union, GaKeston, Texas, 9-14 July 1972. [Abstract] [March]. Moore, D. R. 1974. Remarks on the molluscan tauna ot north- em South America. Bulletin of the American Malacolog- ical Union for 197.3, page 17, Thirt\'-ninth .\nnual Meeting of the American Malacological Union, Newark an Green- ville, Delaware, 24-28 June 1973. [Abstract] [22 Mav] Moore. D. R. 1975. Philobr^idae in the Northern Hemi- sphere. Bulletin of the American Malacological Union for 1974. pages .34— .35. Fortieth Annual Meeting ot the Amer- ican Malacological Union. Spnngfield, Massachusetts, 4- 8 August 1974. [Abstract] [May] Moore, D. R. 1976a. Is Meioceras living in the Indo-Pacif- ic? (Gastropoda: Caecidae). Bulletin ot the American Malacological Union for 1975, pages 19-20, Joint Meet- ing of the Western Society of .Malacologists and Amer- ican Malacological Union [Fort\-first .\nnual .Meeting]. San Diego, California. 22-26 June 1975. [Abstract] [30 Januar)] Moore, D. R. 1976b. .Vlicromolluscs ot the continental shell, northeastern Gulf of Mexico. Bulletin of the American Malacological Union for 1976, page 49, ForU-second An- nual Meeting of the American Malacological Union, Co- lumbus, Ohio, 2-6 August 1976. [Abstract] 130 Decem- ber] Moore, D. R. 1977. Small species of Nuculidae (Bi\al\ia) trom the tropical western Atlantic. The Nautilus 91{4);119-128. figs. 1-7. J27 October]. Moore, D. R 1978a. The western Atlantic Dim\idae. Bulletin of the American .Malacolf)gical Union tor 1976, pages 41- 43, Forts-third .\nnual Meeting of the American Mala- cological Union, Naples, Flonda, 10-15 July 1977, ]Ab- stract] [April] Moore, D. R. 1978b. From the first cniise of the 'Blake" to the present day-one hundred years of progress in the study of Gulf and Caribbean marine mollusks. Bulletin of the American Malacological Union, Inc., tor 1977, pages 63-68, Fortrv-third Annual Meeting of the American Mal- acological Union, Naples, Flonda, l0-15 July 1977. jApril] Moore, d" R. 197Sc. A.xel A, Olsson— An obituary The Nau- tilus 92(2);.59-6]. 1 portrait. [27 April] Moore, D. R. 1978d. The Caecidae of Brazil. Bulletin of the American Malacological Union for 1978, page 56, Forty- fourth Annual Meeting of the American Malacological Union, Wilmington, North Carohna, 16-21 July 1978, [Abstract] Moore, D, R, 1979. On proper citations. The Mollusk JGreater Miami Shell Club. Inc.] 17(2.):4, [2 Febnian ] Moore. D. R, and M, F Miller 1979. Discovery ot living bi- valved gastropods in the Florida Keys. The Nautilus 93(2- 3): 106. [23 April] Moore. D. R 1980a. Molluscan exploration and research in and around the Gulf of Mexico. Bulletin of the Amer- ican Malacological Union, Inc. for 1979, pages 1-5. Forty-fifth Annual Meeting of the American Malacolog- ical Union, Corpus Christi, Texas. 5-11 August 1979. [24 Marcli] Moore, D R 1980b. Distnbution of some small micromol- liisks in the northeastern Gulf of Me.xico. Bulletin of the American .Malacological Union tor 1979, page 64, Forty- fifth Annual Meeting of the American Nlalacological Union, Corpus Christi, Texas, 5-11 August 1979. [Ab- stract] [24 March] Moore, D. R 1980c. The sliallow water fauna of Sanibel and its relationship to upper Cenozoic fossils in South Florida. In: Gleason, P, J, (ed,) Water, oil, and the geology of Col- lier, Lee, and Hendry counties, pages 57-59, .Miami Geo- logical Societv', Miami. ]May] Moore, D, R. 1980d. A long lost (extinct?) mollusc of the Ohio River. Bulletin of the American Malacological Union for 1980, page 70, Forty-sLxth Annual Meeting of the Amer- ican Malacological Union, Louisville. Kentiick-v, 19-25 July 1980, Meeder, J, F,, D. R, Moore and P, Harlem, (organizers). 1981a. Survey of central Florida geology, Miami Geolog- ical Society 1981 Field Tnp, Tnp! 24-26 April 1981. Mi- ami Geological Society, Miami, 44 pages. [24 .'\pnl] Meeder. J. F, D. R. Moore and P Harlem. 1981b. Road log and narrative. In: Meeder, J. F., D. R. Moore and P. Harlem, (organizers). Survey of central Florida ge- ology, pages 6-15, Miami Geological Society 1981 Field Trip, Trip, 24-26 April 1981, Miami Geological Society, Miami. [24 April] Moore, D, R, 1983. The little bivalve Plnnktomija un- masked. Anales del Instituto de Investigaciones Mari- nas de Punta de Betin [Santa Marta. Colombia] 13:123- 132, figs. 1-9, Romer, N. S. and D. R. Moore. 1988. A new species of Al- lania (Rissoidae) from the West Indian region. The Nau- tilus 102(4);131-1.33. 5 figs. [21 December] Leal. J. H, and D, R. Moore, 1989. Ri.ssoina indiscrcta. a new rissoid species from the tropical southwestern Atlantic with Indo-Pacific affinities. Bulletin ot Marine Science 45(1): 139-147. figures 1-12, table 1, [March] Leal, J H. and D, R, Moore, 1993. Thnia esperanza. a new Costellariidae (Mollusca: Gastropoda) from northern Puerto Rico. The Nautilus 107(2):58-62. figs. 1-11. ]28 July] Moore, D, R, 1997. Close encounters with the nurse shark. In: Feitschrift for C Richard Robins in honor of his re- tirement trom the University of Miami. Bulletin of Marine Science 60(3): 101.5-1016, JMay] ACKNOWLEDGMENTS I am indebted to Cynthia Moore, Nancv Voss, Kav Hale, Avis Miller, Helen All^ertson (all from RSMAS, Univer- sity of Miami), and M.G. Harasewych (USNM) for pro- viding materials and information about Donald R. Moore, and/or comments on the manuscript. The fol- lowing curators/museum specialists are gratefullv ac- knowledged for assistance with type material: .^dam Bal- dinger (MCZ), Phillipe Bouchet (MNHN), M.G. Hara- sevvvch, C\mthia .^hem. Rave Germon (USNM), Paula Page 40 THE NAUTILUS, Vol. 112, No. 1 Mikkelsen (AMNH), Tim Pearce (DMNH), Gar\- Ro- senberg (ANSP), Edmund Gittenherger and Jeroen Goud (NNM), John Tavlor and Joan Pickering (BMNH), Nancy Voss (RSMASAJMML). LITERATURE CITED AnoiiMiious. 1955. Beachcomber. Shell News 2.3(2):4-6. [pub- lished b\ Shell Oil Coinpanv. New York] Downey, .VI. E, 196S. A note on the Atlantic species of the starfish genus Linckia. Proceedings of the Biological So- ciety' of Washington 81{l):41-44. Ponder, W. F. and A. Waren. 1988. Classification of the Caen- ogastropoda and Heterostropha - a list of family-group names and higher taxa. Malacological Review, Supplement 4:2SSU32S. Robertson, R. 197.3. Cijclostrcinclla. a planispiral pyramidellid. The Nautilus 87(,3):88. THE NAUTILUS 112(1):41, 1998 Pase 41 Notices THE R. T ABBOTT VISITING CURATORSHIP The Bailey- Matthews Shell Museum is pleased to invite applications for the 1999 R. T. Abbott Visiting Curatorship. The curatorship, estabhshed originally in accordance with the wishes of the late Dr R. T Abbott. Founding Director of the Shell Museum, is awarded annually to enable mollusk systematists to visit the museum for a period of one to tsvo weeks. Abbott Fellows will be expected, by performing collection-based research, to assist with the curation oi the portions of the museum's Mollusk Collection and to pro\ide one evening talk for the general pubhc. The museum collection consists of marine, freshwater, and land shells, with emphasis on SW Florida. A large percentage of the records has been catalogued through a computerized database management system. A substantial portion of the time will be available for research in the museum collection, but field work in SW Florida can be arranged. The R. T Abbott Visiting Curatorship is accompanied by a stipend of $1,500. Interested malacologists are invited to send a copy of their curriculum vitae togeather with a letter detailing their areas of ta.\onomic e.x'pertise and research objectives, and to provide a tentative title for their talk. Send materials to: Dr Jose H. Leal, Director The Btiilev- Matthews Shell Museum P O. Box' 1580 Sanibel, FL 33957 Applications for the 1999 Visiting Curatorship should be sent no later than Januar\- 31, 1999. The award will be announced b>- Februars' 28, 1999. Questions about the R. T Abbott Visiting Curatorship should be directed to Dr. Leal at: (941) 395-2233: fax (941) 395-6706, or e-m;ul: le;d@water.net 7/i Memoham Harold ("Hal") Le\vi,s 1927-1998 INSTRUCTIONS TO AUTHORS THE NAUTILUS publishes papers on all aspects of the biologN' and svstematics of mollusks. Manuscripts describing original, unpublished research as well as review articles will be considered. 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Wise Houston Museum of Natural Science Houston, TX 77030-1799 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 $15.00 for air mail. All orders should be accompanied bv pavment and sent to: THE NAUTILUS,' RO. Box 1580, Sanibel, FL 3.3957, 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 The Biiiley- Matthews Shell Museum, 3075 Sanibel-Claptiva Road, Sanibel, FL 33975. Periodicals postage paid at Sanibel, FL, and additional mailing offices. POSTMASTER: Send address changes to: THE NAUTILUS PO. Box 1580 Sanibel, FL 33957 TH Et^NAUTI LUS CONTENTS Volume 112, Number 2 September 22, 1998 ISSN 0028-1344 E. Suiirez-Morales R. Casca Theeosome pteropod (Gastropoda) assemblages of the Mexican Carilihean Sea (1991) 43 BiTice A. Marshall A new deep-sea limpet of the genus Prctiiunloiitd Dall, 18S2 from New Zealand, and new distribution records for P. aupoiiria antl P Diorioiia Marshall, 1985 (Gastropoda; Acmaeidae) 52 Rediscoven' oi Discus l)ninsoni Bern', 1955 and Orniliclix alpina (Eirod, 1901) in the Mission Mountains, Montana, with comments on Oivohclix clrodi (Pilsbrv, 1900) 58 EpHoiiitiDi fahriziiu (Gastropoda: Epitoniidae), a new species from Patagonia, Argentina 63 Two new species oi Pciiplomo (Bivalvia; Anomalodesmata: Periplomatidae) from the southern Caribbean 69 Paul Hendricks Guido Pastorino Pablo Penchaszadeh Nestor A. Ardila Juan M. Diaz THE NAUTILUS 112(2):43-51, 1998 Paee 43 Thecosome Pteropod (Gastropoda) Assemblages of the Mexican Caribbean Sea (1991) E. Suarez-Morales El Colegio de la Frontera Sur (ECOSUR), Uiiidad Chetiinuil, Apartado Postal 424, Chetmnal, Quintana Roo 77000, MEXICO esuarez®'nicte-lia.ecosur-qroo,nix R. Gasca El Colegio de la Frontera Sur (ECOSUR). Unidad Chetmnal, Apartado Postal 424, Chetiniial, Quintana Roo 77000, MEXICO, ABSTRACT The composition, distnbution, and abundance of the theco- some pteropods collected off the Mexican coast of the Carib- bean Sea were anal\-zed. Samples were collected during four months (Febniar\-, March, May, and August) of 1991, Highest mean thecosome pteropod density was recorded in Ma\ (329 org./IO'm"), while lowest mean densit)' occun'ed in March (71 organism.s/IO'm'). Seventeen species and subspecific tonus were identified. Creseis acicula aciaila. Limacina troclufonnis. L inflata, Clio pijramidata pijramidata and Cavolinia longi- rostris longirostris were the more abundant species, accounting for 84% of the overall thecosome pteropod counts. The first two species represent the influence of Caribbean surface wa- ters in the sur\e\ed area. The local thecosome pteropod fauna also shows some affinits with that from the southeni Gulf of Mexico, Cluster analysis oi the stations re\ealed two tvpes of assemblages of low and high densities, with distributions that showed month-to-month \ariations. Circadian \anations were well defined for two of the more abundant species; the other two species were nearly equally abundant in day and night samples. The thecosome pteropod communitv' along this oce- anic area seems to be represented b\' a homogeneous complex dominated by neritic-oceanic species with a strong influence of C aciaila acicula even in fully oceanic areas, Intermi.xing of these assemblages and the co-occurrence of neritic and oceanic species near the coast could be attributed to the narrow witltli of the continental shelf and to local circulation patterns. INTRODUCTION Thecosome pteropod molluscs are one of the most abun- tlant holoplaiiktonic groups in neritic and oceanic envi- ronments. As a group, thev show an adaptive trend to- ward shell reduction that results in less densih' and in- creased ability for vertical displacement in the water col- umn (Van der Spoel, 1996). The Thecosomata is one of the two holoplaiiktonic orders of Opisthobranchia (the other one is Gyinnosomata) that were lormerK' united under the name Pteropoda. Todav we know that these two orders are not closely related (Lalli and Gilmer, 1989). The term "pteropod" is used herein niidnlv as an easy reference name based on widespread, common practice. Thecosome pteropods are mainly herbivorous, and feed upon microplankton and cUatoms using drifting mucous webs to capture food (Lalh and Gilmer, 19S9). Most species inhabit surface waters, but some e.xhibit large migration patterns from depths of 1000-2000 m. Some species are useful as hydrological, geological and ecological indicators (Biekart, 1989). In the northwestern Atlantic Ocean thecosome pter- opods have been surveyed mainh' in the Gulf of Mexico and along the East Coast of the United States (Wor- melle. 1962; Chen and Hillman. 1970; Matsubara, 1975; Michel and Michel, 1991; Suarez-Morales and Gasca, 1992). Some studies ha\e been made in the Caribbean Sea about general aspects of their abundance and their horizontal and vertical distribution (Wells, 1975, 1976; Haagensen, 1976; Gasca and Suarez-Morales, 1992). A general review of the thecosome pteropod records in the Gulf of Mexico and adjacent areas is presented by Sua- rez-Morales (1994). However, knowledge ol the com- position, distribution and abundance of the thecosome pteropod fauna in the Caribbean Sea off Me.xico, the westernmost portion of the Caribbean Sea, is practically limited to the innermost reaches of the coast (Gasca and Suarez-Morales, 1990). In the present work, the com- position, distribution and abundance of thecosomes pteropods collected in the oceanic area off the Mexican coast of the Caribbean during four oceanographic cruis- es are anal\zed. Sti'dv Are.\ The surveyed area hes behveen 18° and 21°30'N and 86°20' and 87°40'W off the eastern coast of the Yucatan Peninsula, Mexico (Figure 1). The area is under the in- fluence of the northward flow of Caribbean surface wa- ter on its way into the Gulf of Me.xico through the Yu- catan Channel. The continental shelf is narrow along this section of the coast and depth drops \'ery quickly off- shore (Merino and Otero, 1991), H\drographic condi- Pai^e 44 THE NAUTILUS, Vol. 112. No. 2 CAR/BE l-IV Figure 1. Siineved area with zooplankton sampling stations off the Mexican coast of the Caribbean .Sea (1991). tions along the Caribbean coa.st of Me.vito are deter- mined bv the main northward flow of the Yucatan Cur- rent and by a southward-flowing coastal countercurrent (Merino, 1986). The Yucatan Current shows seasonal variation in strength; its miwimum and mininumi inten- sities occur respectively during spring and winter (Me- rino 1992). Three main climatic periods can be distin- guished: drv (March-|une), rainv (July-October), and the "nortes" perifxl, with northern domiTiant winds, from November to Februaiy. MATERIALS AND METHODS Four oceanographic cruises were carried out ilurmg February, March, May, and August 1991, on board dif- ferent "Dragaminas" (minesweeper) ships (jf the Mexi- can Secretan'a dc Marina (Februar): D-12\ March: D- 04; May and August: D-OS). The sampling plan mchuled 22 stations (Figure 1). Zooplankton was coUectetl bv surface ()l)li(|uc hauls (0-10 m) using a sijuare-mouth (0.45 m side) standard plankton net (0..3 nun mesh size). A digital flowiueter was attached to the net Tnoutli to estimate the amount of water filtered i)\ the net (mean volume: 160 m' per haul). Zooplankton samples were fi.xed and preserved in a buffered 4% formalin solution (Smith and Richardson, 1979). Thecosome pti'r()i)ods wi'ri' sorted from the en- tire samples, identified (Van der Spoel, i9Si), and counted. Voucher material of the Mexican Caribbean thecososme pteropods is deposited in the zooplankton collection of El Colegio de la Frontera Sur (ECOSUR) under catalog numbers ECO-CH-Z0073 through ECO- CH-Z0090. Densih,' data were integrated into four data sets, one for each cruise. Shannon's Index of Di\'ersity was calculated (in bits/indi\idual, which ex-presses the average degree of uncertaintv in predicting what species an individual chosen at random will belong to; Lndwig and Reynolds, 1988). The Index of Importance Value (IIV) was used as a dominance measurement. The IIV accounts for relative (percentual) density and frequency of the species. In one standard data matrix, IIV would have a maximum value of 200 ( 1009c oi densitx' plus 100% of frequency) (Brower and Zar, 1977). The IIV has been used more often by botanists, but the index can also be used in faunal studies (De la Cruz, 1994). The Bra\-Curtis Similaritx Index was used to identifv' station clusters and anaKze the thecosome pteropod communitA stnicture within indi\idual cruises. RESULTS Mean temperature dunng the four-month perioil was 28°C, with a minimum of 23°C in Febmary and maxi- mum of 29°C in August. Salinitx' averaged .'35.2 %c. with a maximum of .3.5.6 %c in Mav and a minimum of .34.5 %o in Febniarv. Space distribution of thecosome pteropod densities in the surveyed area showed monthly variations (Table 1). Highest total mean densities were recorded during May (296.25 organism.s/10'm'), followed by Februaiy (158 org./lO^m'), August (141.79 org./10'm'), and March (71 org./10'm'). During February, thecosome pteropods showed txvo high-densitv zones, one in the northern por- tion of the area, and the other in the central zone. No high-density zones were detected during March. In May, thecosome pteropods showed high densities along the entire coast, and in August two groups of stations with high densities were obseiyed, one in the north and the other in the central portion. Most high-densitx- stations were sampled at night (Figure 2). A total of 17 species and subspecific forms were col- lected in the suneved area (Table 1). Four species, Cic- sci.s (Uivula acinila (Rang, 1828), Liniaciua inflata (d'Orbignv, 1836), L. trorhifonnis (d'Orbigny, 1836), and Clio piirainidala j)i/raini(latti Linnaeus, 1767 were re- cordeil as the douunant thecosome pteropods in the area throughout the surveyed peiioil. Together they consti- tuted 86.6% of the total overall pteropod catch (83% in FebmaiA', 81% in March; 95.6% during May; 74.6% in .August). The relati\e abundance (%) and mean density (org./lOin') of each species during the four cruises is shown in Table 1 . Overall thecosome pteropod densities were 2.2 times higher during the night (mean density 298.75 org./ lO'nr) than during da\time (mean den.sitx' 133 org./ lO'm). During the nigtit samplings, theco.some ptei'o- pods were most abundant during May (499 org./10'm'). E. Siuirez-Morales ami R. Gasca. 19yS Paiie 45 Table 1. Mean deiisih (ofg./lO'i and felati\e abundance i7c) ot pteropod Mexitan Canhbean Sea 11991' species dniin2 eacli iit tonr CARIBE cnii.ses in tin CARIBE I CARIBE II CARIBE III CARIBE I\ Total FebmarA Marc li Ma\ August Mean Densit\ ^f Deiisit) '7r DensitN 7, Densit^ % Densit\- Creseis ncicuin acicula (Rang, 3800 24 13 4.44 6.23 160.30 54.11 55.28 38.90 64.50 1S2S) Creseis acinila tlina (Rang, 4.31 2.74 8.10 2.73 2..52 1.77 3.73 1S2S) Creseis viroula eoniea Escli- 2.7,5 1.75 12.96 18.11 0.35 0.12 2.77 1.95 4.70 scholtz, 1S29 Clio pijramiduta pijramidatn 14.63 9.29 19.18 26.93 20.40 6.90 3.31 2.33 14.37 Linnaeus, 1767 Clio pi/rnmidatn lanceolata 0.44 0.62 Oil (Lesneur. 1S13) Diaeria qiiadridentata ide 0,70 024 0 17 Blaimnlle, 1S2H Cavolinia lon 53), followed by C. pijramidata pijramidata (IIV = 24). Although diversity was variable (0.6-1.4), on average it was moderately high (1.22 bits/ind.). Clustering revealed tsvo station groups (Figure 5e). The first one included stations with less than 45 org./lO'm', and contained onlv 11.7%' of the the- cosome pteropod numbers in this month. In most of these stations Creseis aeicula acicula was the onl\' spe- Page 46 THE NAUTILUS, Vol. 112, No. 2 CAR/BE/ (February, 1991 CAR/BE// (March. 1991) CARIBEIII (May, 1991) CARIBE IV (August, 1 991) Figure 2. Monthly deiisitie.'i (org./lO'ni') of thecosome pteropods for each sampling; station dnrincr the snneyed period Me,\ican Caribbean Sea, Black circles indicate night samples. the cies present, but with densities below 30 org,/10'ml The group was distributed in the fully oceanic zone and in the shelf border area. The second assemblage showed the same general distribution as the first one; it included stations with the highest density' values (most over 100 org,/10'm^) (Figure 5a), Liinacina inflata was character- istic of this group, which comprised 100% of the occur- rence of the species. In March, Limacina trochifonnis represented up to 35,6% of the total thecosome pteropod numbers, fol- lowed by C, ptjramidata pyramidata (27%) and by C, virgithi conica (18%), Clio piiramidafa ptjramidata and Crcscis acicula acicula occurred in more than (S0% ot the stations, the former with higher densities (Figures 3b, 4f), Both species oi Limacina showed a limited dis- tribution, and were more frequent in night samples (Fig- ures 3f, 4b), Nighttime densit\- represented 47%- ot the total catch (mean densit)': 76 (day) us-, 69 (night) org./ lO^m'), Clio pyramidata pyramidata was dominant (IIV = 56,3), followed by L, trocliiformi.s (49) and C. vir^ula conica (36), Diversity was moderate (mean 1,14), slightly lower than in February. Two station clusters were re- vealed (Figure 5f), with a distribution similar to that oi February (Figure 5b), The first group was characterized by low to moderate thecosome pteropod di'usities (2S- 128 org,/10'm'), and mcludetl only 38% ol' the total the- cosome pteropod density. Up to 75% of C. pyramidata pyramidata and 96% of L. inflata numbers were includ- ed in this group, A second group showed low and high densities (5-344 org,/10'm'), and was characterized In' the highest densities of L, trochifonnis; this group in- cluded up to 84% of the total number of C, vir^ula conica. In May the most abuuilant species was Crcscis acic- ula acicula, representing about 54%- ol the total the- cosome pteropod catch; it was followed by L. injlata (30%) and by L. trochifonnis (4,4%), Only C, acicula acicula occurred at all stations, with medium and high densities (Figure 3c); C, pyramidata pyramidata was collected at most stations with medium densities (Fig- in"e 4g), and L trocliiformis was collected at a few sta- tions (Figure 4c). Limacina inflata was abundant in nighttime samples on the southern portion of the area (Figure 3g). Nighttime overall densit\' represented 65% of the total catch and was 1.8 times higher than davtime density (499 vs. 264 org./10'm'). The IIV showed the dominance of C, acicula acicula (80) and of L, injlata (42,1) in this month. Mean diversit\ was moderate (1.10 bit,s/ind.), but slightly lower than in the tvvo pre- vious months. Two clusters were defined (Figure 5 g). The first one included only four stations with the lowest densities (9-29 org,/10'm'). Only 0,1% of the theco- some pteropod numbers occurred in this cluster; these stations were made in the oceanic area and near the coast (Figure 5c), Crcscis acicula acicula and Clio pyr- amidata pyramidata were present in this group. The second cluster showed the highest total densities (over 98 and up to 1578 org,/10in'), with more than 98% ot C, acicula acicula and 1009( of L. inflata numbers. This group showed a variable (h\ersity. In August, Crcscis acicula acicula was again the most abundant species (39% relati\e abundance), followed by Caioliuia low^irostris limhata (19. 3%), by L. inflata ( i6,9%) and by L. trochifonnis ( 16,3%), Crcscis acicula acicula occurred at all stations with medium and high densities (Figure 3 d), only both species of Limacina occurred at more than 50% with medium density \alues (Figures 3 h, 4 d), while the remaining species each occurred at less than 20% of the sampling sites, Clio pyramidata pipamidata was scarce in this cruise (Fig- ure 4 h). Nearly 75%' of the total densitv' was collected in nighttime samples; mean density was about 3 times higlier duruig night collections (295 vs. 97 org,/10''m'), C'rcscis acicula acicula sliov\ed the highest dominance E. Suarez-Morales and R. Gasca, 1998 Pa^e 47 — -1P-V.O a p p 4 f CARIBE 1 J. Creseis acicula acicula CARIBE I Limacina inflata Figure 3. Distribution and densit^ of the most ubumlant species during the sunesed period, a-d. Crc: Liiiiticiiui inflatd. Black circles indicate night samples. ,s'('(.s acicula acicula. e-h. (IIV = 61.6), while L inflata. C. lonfiirostris limhata were less dominant. Di\ersit:v fluctuated (0.67-1. 28) and was in average (1.16 bits/ind.) the second highest in the survey. Again, t\vo clusters were defined, as shown in Figure 5h. The first assemblage, with stations distributed at the middle and southern portions of the area, was characterized by the lowest total thecosome pteropod densities (below 20 org./10'm') and contained onh' 1.49( of the total thecosome pteropod counts. In this four-station cluster, C. acicula acicula was either absent or showed its lowest densities. Diversity was low. The second assemblage was distributed all along the surveyed area and showed the highest total densities (40-450 org./10'm') (Figure 5d). More than 98% of C. acicula acicula. 100% of L trcchifonnifi and of L. in- flata occurred in this assemblage. Diversity was vari- able. DISCUSSION Previous records ot thecosome pteropods from the tropical northwestern Atlantic report 51 species and subspecific forms (Wells, 1975, 1976; Haagensen, 1976: Suarez-Morales and Gasca, 1990; Michel and Michel, 1991; Suarez-Morales, 1994). Most of these records are from the Gulf of Mexico and the central and eastern portions of the Caribbean Sea. All the species identified here have been reported pre\iouslv from adjacent areas of the Northwestern Atlantic (Suarez-Morales, 1994). In a coastal system adjacent to the surveyed area, only 6 species have been recorded (Gasca and Suarez-Mo- rales, 1990). However the present study shows for the first time faunal records of thecosome pteropods in the oceanic area of the Caribbean Sea off Mexico. More- over, occurrence of Limacina Ic.sucuri. Creseis acicula Page 48 THE NAUTILUS, Vol. 112. No. 2 Figure 4. Distnliiitioii and ilt'iLsih ot the nuxst aluiiKlaiit .species duniig the suneved period, a-d. Liinacina trochijoniiis: e-h. Clio ]njniini(lafn jii/rdiiiidata Black circles indicate niijht sanijiles. clava and Ci/iiihiilia pcroni in the Caribbean Sea is herein recorded for the first time. These species were pre\i()uslv known onl\ from the central and sonthern portions of the Gulf of Me.xico (Suarez-Morales, 1994) The main branch of the Yucatan Current, which trans- ports Caribliean surface waters northward, iiffects offsliort^ conditions in the Mexican C>'aribliean. The current repre- sents an important factor that influences the fauuiil tlistii- butions and the dynamics of the entire ecosystem. The pteropod fauna of tropical surface waters of tlie C;iribbean is dominatefl bv a group of sevenil species (Haaiienseu, 1976), wluch include Cirsci.s (iciailu, Liiiwrina tiDcliiJor- mis, Creseis virffila and CavoUnia lon^roatris. Only part of this group of species w;ls found to be dominant in tlic surface layer of the Caribbean Sea off Me.xico during the present shidv. The lormer two .species indicate the prinian influence of die (Caribbean suriace water oii the study ;uea. The low abundance of C. vArfftla conira and C. langirosth-s might result from specific, unknown local conditions. The niiiin species group of the Mexican (Caribbean is similar to that reported bv Suarez-Moniles (1992> for the southern Gulf of Mexico (C. aciaila aciaila, L. injiata. L. trochijor- ini-s, C. lonjS,ir('stri.s lonprostris). The near-shore and continent;il shelf oceaiiographic dv- uamics along the (Caribbean coast ot Mexico are related to the strength of a coast;il coimtercurrent moxing soudi- wards (Merino, 1986) from the northernmost edge of the (Caribbean coast, which is iilso the easternmost portion of the (^ampi'che Bank. Its influence would explain, at least partialK, a certain affinitx of the local thecosome pteropod iaiuia with that of the southern (iulf of Mexico, with at least 60% of species in conunou behveiMi the 2 areas (Le;il, 1965; Matsubara, 1975; Suarez-Moniles and Gasca, 1992). The species composition of the loc;il thecosome pteropod fauna seems to represent a mi.xed faunal com- plex of Caribbean and Gulf of Mexico ;iffinities, E. Suarez-Morales and R. Gasca, 1998 Pas^e 49 CARIBE I (February) fl 14 7 24 23 10 21 15 6 5 20 12 19 11 t^ CARIBE II |_ (March) U H- r20 •-19 16 ■15 13 21 10 ^ 14 CARIBE IV (August) Figure 5. Distribution of pteropod assemblages a-d. Obtained from the Bray-Curtis Index dunuij; the drogrums resultinsl from the Bra\-Curtis Index during the tour months suneved. tour monttis ths e-h. Den- In thf surveyed area, the oceanic thecosome pteropod Limocina hulimoidcs was collected rjuite close to the coast, as most of the species (either neritic-oceanic or neritic) recorded here. In Bahia de Ascension, a coastal- estuarine s\steni on the central coast ot the Mexican Caribbean (Figure 1), species such as C. acicnia acicnla. L. trocliifonui.s and C. lougiro.stiif; loiiaimstiis have been recorded well inside this system (Suarez-Morales and Gasca, 1990). The occurrence of neritic/oceanic zoo- plankton within the estuarine systems on this coast has been studied by Zamponi and Suarez-Morales (1991) and by Suarez-Morales and Gasca (199fi). According to local studies of coastal suriace currents (Merino, 1986), the plankton carried northward by the western edge ot the Yucatan Current tend to drift inshore. This niiglit explain the presence of oceanic species yen near the coast, over the narrow shelf, antl in the estuarine sys- tems. The inshore drift might also promote the hydro- graphic homogeneity represented bv the salinity and temperature values recorded during the sur\e\' period It is generally accepted that the low variability of these factors does not have a pronounced effect on the zoo- plankton distribution in the Caribbean (Hubbard et al., 1991). Another possible consecjuence of the narrow shelf present in this area is the presence of deep-sea species near the shelf break (Segura-Puertas and Ordonez- Lopez, 1994). Howexer, no deep-sea thecosome ptero- pods were recoriled during this sunex'. Most species have been reported as common in ne- ritic ami oceanic waters, antl represent more than 85% of the recorded species; C. acicnla acicida. the most abundant thecosome in the area, has been considered to be the most aiumdant thecosome in Caribbean neritic- coastal waters. (Haagensen, 1976). The influence of ne- ritic yvaters on these otherwise mainly oceanic assem- blages is represented bv the occurrence of high densities of C. acicnla acicnla. The study area clearly shows a strong oceanic affinitx' with a variable neritic influence. Neritic-oceanic species were mmiericallv dominant or subdominant durinsi; the four-month survey and can be Paee 50 THE NAUTILUS. Vol. 112. No. 2 regarded as the m;iiii group ol repre.sentative local pter- opod.s. Thi.s is probably a result of local mixing processes within the boundaries of the main current and the coast- al countercurrent. In a survey of fish larvae, Sanchez- Velasco and Flores-Coto (1994) found that the Carib- bean Sea off Mexico is characterized mainly by oceanic species intermixed with neridc-coastal species. The Brav-Curtis Index clusters show a similar pattern throughout the area during the tour sui"veved periods, with overlapping distributions and irregular z(jnation patterns. Clusters were sorted m;iinly by whether sta- tions showed low or high densities. However, day-night results were partialK' affected bv the migratory circadian behavior of the most abundant species. This was evident for Limacina inflata and C. pi/ranudata pi/ramklata. known to be present during the dav below the 10()-2()() m, moving up to the surface layer at night (Haagensen, 1976; Wormuth, 1981; Gilmer and Harbison, 1986). In the sur(.eved area, up to 85% of the local densitv ot L. inflata and over 70% of C. pijramidata pi/ramidata den- sit)' was recorded in night trawls. Clearly, the high-densit\' assemblages were deter- mined mostly by the night samples; two (C acicula acic- ida and L. trochiforims) out of four dominant species were represented bv similar densities during both day and night samples, as recorded bv Haagensen (1976) in the Caribbean. This, and the fact that overall dav-night differences were not significant, would buffer the effect of circadian variations in our cluster patterns. It appears that, even with the migration effect, the species com- position did not varv significantlv, which suggests that the local thecosome pteropod community is more or less homogeneous throughout the surveyed area. High and low densitv assemblages overlapped in several areas din- ing the four cruises, probably as a result of strong and continuous mixing processes between the oceanic and the neritic environments. Monthly variations in theco- some pteropod density and distribution may be related to the influence of the Yucatan Current, which increases in the northern portion of the suivey area during late spring upwelhng (May). During winter and early spring (March-Februarv), the current is weak and the environ- ment becomes oligotrophic (Merino, 1992). This could cau.se a slight decrease in density (as obsened iluring March), but the community' stnicture remains othenvise relatively stable throughout the four-month period. The studied area represents a nearly homogeneous mixing zone in which oceanic and neritic-oceanic species were represented tluring the survey periods by variable and mixed cbstributional patterns of distriliution. This could be related to the presence of different environ- ments (coastal, neritic, oceanic! within a relatively small area, and to the dispersal and transport caused the local hydrologic dviiamics (mixing processes, eddies). ACKNOWLEDGEMENTS We received financial support from Mexico's (X3NAC\T (Projects D112-9()452() and 1189-N9203). We gratefully acknowledge the authorities of the Mexico's 11th Naval Zone and of the Secretaria de Marina (Depannent of the Navy) for access to the cruises of the "Dragaminas' ships. Rosa Maria Hernandez Flores and I. Castellanos participated actively in the project and in the laboraton' work. LITERATURE CITED Biekart, ]. W. 19S9. Eiitliecosomatoiis pteropods as paleo-hv- drological antl paleoecological indicators in the Tvrrenian deep-sea core. Palaeogeograpliv, Palaeocliniatologv, Palaeoecologv' 71:205-224. Brower, J. E. and J. H. Zar 1977. Field and Laboratory' Meth- ods for General Ecology. William C. Brown Co., Iowa, 194 pp. Chen, C. and N. Hiilman. 1970. Shell bearing pteropods as indicators of water masses off Cape Hatferas, North Car- olma. Bulletin of Marine Science of the Gulf and Carib- bean 20:350-367. De la Cruz, G. 1994. ANACOM. Sistema para el analisis de comunidades. Version 3.0, Manual del usuario. CINMilS- TA\-IPN, Merida. 99 pp. Gasca, R. v E. Suarez-Morales. 1992. Pteropodos (Mollusca: Gastropoda: Thecosomata) de la Bahi'a de la Ascension, Resen'a de la Biosfera de Sian Ka'an. Quintana Roo, Mex- ico. In: D. Navarro v E. Suarez-Morales (eds. ) Diversidad Biologica en la Reserva de la Biosfera de Sian Ka'an, Qmntana Roo, Mexico. II. Centro de Investigaciones de Qmntana Roo/SEDESOL. Mexico. 115-122. Gilmer R. W. and R. G. Harbi.son. 1986. Moq:)hologv and field behavior of pteropod molluscs: feeding methods in the families Cavoliniidae, Limacinidae and Peraclididae (Gas- tropoda: Thecosomata). Marine Biology 91(1 ):47-57. Haagensen, D. A. 1976. Caribbean zooplankton. Part 11, The- cosomata. Office of Naval Research, Department of the Navy. Washington, pp. 551-712. Hubbard, R. H., ). B. R. Agard and S. N. Gajbhiye. 1991. Groups of zooplankion along environmental gradients in the Caribbean. Caribbean M^irine Studies 2(r-2):54-59. Lalli. C. M. and R. W. Gilmer 19S9. Pelagic snails. The Biologv' of holoplanktonic gastropod molluscs. Stanford tlniversiU' Press, Stanford, 259 pp. Leal, D. G. 1965, Distnhucion de los pteropodos de \'eracnr/, \'er Anales del Instituto de Biologia de la Universidad Nacional Autonoma de Mexico 36:249-251. Ludwig, J. A. and J. F. Reynolds. 1988. Statistical ecology. A primer on methods and computing. John Wiley & Sons, New York, 337 pp, Matsubara, ]. A. 1975, Sistematica, distribucion, ahundancia y relaciones ambientales de los pteropodos tecosomados de la Bahia de Campeche. Mexico. Tesis |irotesional, Fac. de Ciencias, Univi-rsidatl Nacional Autonoma de Mexico, Mexico, 52 pp. Menuo. M 1986. Aspectos de la circulacion costera superficial del Caribe Mexicano con base en obsenaciones utilizando larjetas de deriva. Anales del Instituto de Ciencias del Mar \ Limnologia 13:31—46 .Vleriiio, M, 1992, Afloramiento en la plal.itorma di- Yucatan: estnictura v fertili/aiion, Tesis doctoral, Instituto de Cien- cias del Mar \ Linmologia. lhii\ersida(l Nacional .Auton- oma de Mexico, Mexico, 255 pp. Menno. M, and I„ Otero, 1991. Atlas amhu-ntal costero de Puerto Morelos Centro de Investigac ioenes de Quintana E. Suarez-Morules ant! R. Giisca, 1998 Page 51 Roo /Institiito lie Ciencias del Mar \ Limnologia, Univ- ersidad Nacional Autononia de Mexico, Mexico. 80 pp. Michel, H. B. and J. F Michel, 1991. Heteropod and theco- sonie (Molln.sca: Gastropoda) niacroplankton in the Flor- ida Straits, Bulletin of Manne Science 49:563-574, SanchezA'elasco, L, and C, Flores-Coto. 1994. Larval fish as- semblages at the Yucatan Shelf and in the Mexican Carib- bean Sea during the upwelling period (spring, 1985). Scientia Marina 58:289-297. Segura-Puertas, L, and U. Ordoiiez- Lopez. 1994. Analisi.s de la coinunidad de medusas (Cnidaria) de la region oriental del Banco de Campeche v el Caribe Mexicano, Caribbean Journal of Science 30(1-2):104-115. Smith, P. E. and S, L, Richardson, 1979. Tecnicas modelo para prospecciones de huexos v larvas de peces pelagicos. F.A.O. Docuniientos Tecnicos de Pesca 175:1-107. Suarez-Morales, E. 1994. Distribucion de los pteropodos (Gas- tropoda: Thecosomata v Pseudothecosomata) del Golfo de Mexico V zonas advacentes. Revdsta de Biologia Tropical 42(3):523-530. Suarez-Morales, E, and R, Gasca, 1992, Pteropodos (Gastrop- oda: Thecosomata y Pseudothecosomata) del estrato su- perficial (0-50 m) del sur del Golfo de Mexico. Anales del Institute de Ciencias del Mar y Limnologia 19:199-207. Suarez-Morales, E. and R, Gasca, 1996, Planktonic copepods of Bahia de la Ascension, Caribbean coast of Mexico, a seasonal survey, Cnistaceana 69(2): 162-174. \'an der Spoel, S. 1996. Pteropoda. In: R. Gasca and E, Suarez- ,M()rales (eds,), Introduccion al Estudio del Zooplancton Manno. El Colecj;io de la Frontera Sur/CONAC^T, Mex- ico, pp, 459-.52S'. Wells, F E. 1975. Comparison of euthecosoniatous pteropods in the plankton and sediments oft Barbados, West Indies, Proceedings of the Malacological Society of London 41: 50:3-509. Wells, F E, 1976. Seasonal patterns of abundance and repro- duction of eutheco.somatoiis pteropods off Barbados, West Indies. The Vehger 18:241-248. Wonuelle, R. L. 1962. A survey of the standing crop of plank- ton of the Florida Current. \T. A study of the distribution of the pteropods of the Florida Current. Bulletin of Ma- nne Science of the Gulf and Caribbean 12:95-136. Worniuth, ]. H. 1981. Vertical distributions and diel migrations ot Euthecosomata in the northwest Sargasso Sea. Deep- Sea Research 28:149.3-1575. Zamponi, M, O, and E. Suarez-Morales. 1991. Algunas me- dusas del Mar Caribe Mexicano con la descripcion de Te- traotoporpa siankaanensis gen. et sp nov. (Narcomedusae: Aeginidae). Spheniscus 9:41—46. THE NAUTILUS 112(2):52-57. 1998 Paw 52 A New Deep-sea Limpet of the Genus Pectinodonta Dall, 1882 from New Zealand, and New Distribution Records for P. aupouria and P. mohoria Marshall, 1985 (Gastropoda: Acmaeidae) Bruce A. Marshall Museum of New Zealand Te Papa Tongarewa P.O. Box 467 Wellington, NEW ZEALAND bnic-em@tepapa.go\t,nz ABSTRACT Pectiiuxltitilii iiuiriiioiichi n. sp, is ile.senbetl antl illustrated. and new distrihution data are pro\ided tor P. aupouria Mar- shall, 19S5 and P moriorui Marshall. 1985. Kei/ wonts: Mollusca, Acmaeidae, Pccthuxhrnta. tieep-sea, new species. New Zealand INTRODUCTION Since the (li.sc()ver\' of two Recent Prctinixlonta species in the New Zealand region {Marshall, 1985), a third spe- cies has been recognized, and considerable additional material of the other two species has come to hand. The opportunit\' is taken to record this additional material and to plot the distributions. Note that New Zealand Pectinodonta species have onlv been found in areas that have been extensivelv worked by commercial trawlers, so their tnie distributions are likelv to be more extensive than implied bv the cbstribution maps presented here (Figures 6-8). Acronyms: NMNZ, Museum of New Zea- land, Welhngt(m; NZ'OI, National Institute of Water and Atmosphi'ric Research, Wellington. SYSTEMATICS Superfamily Acmaeoidea Forbes. I S5() Family Acmaeidae Forbes, 1850 Subfamily Pectinodontinae Pilsbn. 1891 Genus Pcdinoilonta Dall, 1882 Pectinodonta Dall. 18S2: 409. T\])e species (In onginal desig- natioii); Pectinodonta arcnata Dall, 18cS2; Recent. C!:aril)- bean Rcmark.s: Liudberg (198fi) rcdclincd .\cniaeidae, re- stricted it to include Aciiuieti iniira f-Jathke, 18.3.3 (Ac- maeinae) and Pcctinodoida species (Pectinodontinae) alone, and referred most other ta\a hitherto referred there to Lottiidae. More recently Okiitani ct aJ. (1992) have introduced two new genera (Scrradonta and Beith- i/avniaca) for pectinodontines with highly chstincti\e rad- ulae that live on vestimentiferan tubes and Calijptogena \'aK'es from a bathval cold seep in Sagami Bay Japan. Worldwide Recent and Tertian' Pectinodonta species were listed by Marshall (1985). Liudberg and Hedegaard (1996) discussed pectinodontine shell stnicture, reciiiit- ment patterns and ph\logenetic relationships. Pectinodonta maiinovichi new species (Figures 1-6) Pectinodonta aupouria Marsliall. 1985: 277 (in part: largest measnreil parahpe). Description: Shell (Figures 1, 2) up to 25.4 mm long, strongK arched, of moderate thickness, white, exterior dull, interior porcellanous. Anterior end occupying 20- ■5] % of shell length; anterior slope weakly concave in most specimens, in others almost flat or weakly con\-ex; posterior slope rather strongK convex: lateral slopes wt'akly convex. Aperture elliptical, shallowlv or \er\' shal- lowly concave from side to side, sides broadly rounded, anterior end more broadK rounded than posterior. With growth, shell gradually t^\isting clockA\ise about 10° be- tween ju\enile and adult. Teleoconch at 1.6-1.8 mm length translucent and glossy, smooth apart from \ery fine growth lines: fine radial hues show through outer shell layer but do not resolve as surface features. Sub- siMjueiit teleoconch chalk-\- white, with low, rounded ra- dial ribs, interspaces wider than each rib: and stronger, rounded or roundK augulate concentric ridges. Fine, crowded, concentric growth lines throughout. Myostrac- um clearly defined. Animal with eyes lacking. Foot and ui:inlle edge cream wliiti'. Large, tapered ctenidium ex- teuilmg antero-lateralK from left to right behmd head. Mantle edge weakK and irregularly serrate. Muzzle lap- B, A. Marshall, 1998 Paee 53 Figures 1-5. Pcctiiuxlcntd inuriiiovichi new species 1, 2. Holohpe, ott Cape Kidnappers, 1100 in. length 25.4 nun. 3-5. Radula of holot)pe. 3. Inner side of single tooth (length 370 |jLm). 4. Part of ratlnlar rihhon showing teeth in situ attached to basal plates (width .3.30 (jLni)- 5. Part ot radnlar ribbon showing basal plates from which teeth ha\e been renio\ed (shrinkage is minimal) (width 370 fjLm). pets thin, broad. Cephalic tentacles small, circular in sec- tion, tapered, tips rountled. Radular nhbon about 409^ longer than shell. Radular formula 0+3 + 0+3 + 0, each tooth (as defined here) comprising 3 (used laterals, in- dividual units ot which are defined bv fine gro(j\'es (Fig- ures 3-5). Teeth arranged in a posteriorK' tli\erging \'- shape; each with 13 or 14 conical cusps, innermost cusp largest, next 3 cusps smaller, outer cusps yet smaller and similar. Type data: Holobvpe, NMNZ M. 127085 (length 25.4 mm, width 19.4 mm. height 15.0 mm) and 3 paratopes, NMNZ M. 127023: Off Cape Kidnappers, 39° 50' S.177° 39' E, ;ilive on wood with P. aupouria, 1100 m, 10 Au- gust 1995, F.V. Petersen, coll. M. Marino\ich. Other material examined: Off White Island, Bav of PlentA-, New Zealand, 37° 23.7' S, 177° 39.5' E, alive on wood together with P aupouria. 1075-1 10() m, 23 No- vember 1981, coll. O.M. Moore and P.J. McMillan, F.V. Kalinovo (stn. KOl/19/81) (3, NMNZ M. 126299); off East Cape. 37° 32.6' S. 179° 19.3' E, ali\-e on wood with Page 54 THE NAUTILUS, Vol. 112, No. 2 Figure 6. Map of New Zealand showing distribution of Pectinodonta nmrinovichi new species: 200 and 1000 ni isobaths indicated. P. aupotiria, 1280-1215 ni, 4 August 1974, R.V. Taiijia- roa (2, NZOI, stii. KS73/2); N of Cape Runaway, 37° 19.46' S, 178° 04.16' E, alive on wood with P. aupouria, 1124-1169 m, 14 Mav 1995, F.R.V. Tangaroa (stn. 9506/ 42) (1, NMNZ M. 126152); off Ea.st Cape, 37° 47.95' S, 179° 04.35' E, alive on wood, 1289-1364 m, 23 March 1993, F.R.V. Tangaroa (stn. 9.303/57) (33, NMNZ M. 118799); off Gi.sbome, 38° 49.14' S, 178° 34.81' E, alive on wood, 736-760 m, 25 March 1993, FR.V Tan- garoa (stn. 9.30.3/69) (6, NMNZ M. 117958); off Cape Egmont, 38° 58.5' S, 172° 10.2' E, alive on wood, 1045- 1055 m, 3 June 1986, FV Wanaka (stn. WK5/17/86) (1, NMNZ M.86819); off Mahia Peninsula, 39° 25.7' S, 178° 25.3' E, alive on wood, 921-939 m, 24 June 1986, FV. Otago CallianI (stn. (JC2/31/86) (20, NMNZ M. 126569); off Hokitika, 43° 27' S, 168° 47' E, alive on wood, 910-987 m, 17 May 1985 (3, NMNZ M. 118017). Distribution (figure 6): Off White Island to off Hok- itika. New Zealand, lixing and ieethng on wood, 736- 1364 m. Remarks: Pectinodonta nmiinovichi attains larger size than P. aupouria and smaller size than P. morioria (length up to 25.4 mm against 18.2 mm and 14.7 mm respecti\e]\) It differs further from lioth these species in that the radial ribs are nnich narrower, and the inter- spaces are broader than each rib instead of much nar- rower. The anterior slope is weakly concave in most of specimens of P. marinoiichi, but convex in most speci- mens of /' (ntponria The nuintle edge is more weakly B. A. Marshall 1998 Page 55 Figure 7. Map of New Zealand showing distiilnition oi Pcctinoclonta aiipoiiria Murshall, 19.S5: 200 and 1000 ni isoliatlis indicated. serrate than in P aiipoiiria. Tlie railiila resembles that of F aupouiia and has more numerous cusps than that of P. morioria. P. imirinovichi has been taken twite li\ing together on the same piece of wood as P. aiipoiiria. Three specimens of P. marinovichi were found among the 1232 parat\pes of P. aiipoiiria Marshall, 1985, the largest of which (length 20.2 mm) was imtortunatelv in- cluded in a table of shell measurements lor that species (Marshall, 1985, Table 1). Etymology: After Miro Marino\ich, F.V. Petersen (Si- munovich Fisheries, Auckland), who collected the ho- lotype and who has acquired tremendous quantities of bvcatch for the Museum of New Zealantl. Pectimnlonta anpoiiria Marshall. 1985 (Figure 7) Pcctinoclonta tjiipouria Marsliall, 198.5: 277. fig. 1. 3F-I. 4A, D (in part; largest paratope = P. marinovichi n. sp.; spec- imen identified h\ Powell (1979) as Maoricrntcr cxplorata = P morioria). Type data: Holotspe, M. 76089 and 1229 paratvpes, NMNZ; Off White Island, Bav of Plenty-, New Zealand, 37° 23.7' S, 177° 39.5' E, alive on wood together with P. marinovichi n. sp., 1075-1100 m, 23 Noxember 1981, coll. O.M. Moore and P.J. McMillan, F.V. Kalinovo (stn. KO 1/19/81). Other material examined: N of Cape Runaway, 37° Page 56 THE NAUTILUS, Vol. 112. No. 2 Figure 8. .\I;i]-) of New Zealand showing ilistnhiition uf Pcctiiiixloiitti inorior'ui Marsiiall. 1985: 2(10 ami lOOO ni isohatlis indicated. 19.46' S, 17(S° 04.16' E, alive on wood witli /' marinov- ichi, 1124-1169 m, 14 Mav 1995. F.R.V. Tanoowa (stn. 9506/42) (1, NMNZ M. 126151 ); off East Cape, 37° 32.6' S, 179° 19.3' E, alhe on wood with P. iniraiuiiiclu n. sp., 1280-1215 m, 4 Angnst 1974. R.V. Taiipe data: Hok>t^pe, NMNZ M. 76090 and mam paratopes, NMNZ NI, 75107: NE of Menioo Bank, Chat- ham Rise, New Zealand, 42° 43,9' S, 176° 08' E, alive on wood, 800-810 m, 28 September 1982, F.V. Krdtan (KTN/152/82). Parat^pes (4, NMNZ M.75242): Off Ti- maru, 44° 33.2' S. 17.3° 42.3' E, alive on wood, 750-738 m, 8 June 1984, F.R.V. James Cook (stn. JlO/4/84). Other material examined: Oft Aldermen Islands, 366—176 m, on wood (1, Auckland Institute and Muse- um AK131610); off Aldermen Islands, alive on wood, 400 m, April 1988, pres. D. Gibbs (2, NMNZ M.95302); off Cape Runawav, 37° 29.9' S, 177° 47' E. alive on wood, 450-481 m,'8 December 1985, FV. Wanoka (stn. \\'K3/19/85) (ca. 1000, NMNZ M.84252); off East Cape, 37° 47.64' S, 179° 04.45' E, ahve on wood, 1289-1342 m. 24 March 1992, FR.V. Taiii^ama (stn. 9203/133) (27. NMNZ M. 117898); off Maliia Peninsula, 39° 05' S, 178° 10' E, ahve on wood with P. atipoiiria. 900-1000 m, June 1994, coll. M. Manno\ich, FV. Petersen (18, NMNZ M. 118321); off Cape Kidnappers, 39° 42.91' S. 178° 10.15' E, alive on wood with P. aiipoiiria. 818-952 m, 8 June 1993, coll. M. Clark, FR.V. Tan^aiva (stn. 9306/ 210) (.several hundred, NMNZ M. 117996); off Hokitika, 42° 37.6' S, 170° 02.8' E, alive on wood, 490-439 m, 13 October 1988, F.V. Diashin Mam 22 (stn. 269/24) (sev- eral hundred, NMNZ M. 92467); slope of Mernoo Bank, Chatham Ri.se, 42° 58.06' S. 175° 01.03' E, N alue on wood, 545 m, 15 Januarv 1992, FR.V. Tanzania (stn. 9106/91) (manv hundreds, NMNZ M. 126570); E of Mernoo Bank, Chatham Rise, 42° 53'- 43° 08' S, 176° 04'-177° 00' E, alive on wood, 370-420 m, December 1994, coll. M. Marinovich, FV Petersen (50, NMNZ M. 119153); off Puv.segur Point, 46° 29' S, 166° 14' E, alive on wood, 560 m, 20 October 1987, F.V. Cdiii/o Mant 5 (.52, NMNZ M.90196). Distribution (figure 8): Aldermen Islands to off Puv- segur Point, New Zealand, h\ing and feethng on wood, 366-1342 m. Remarks: Pcctinodonta niorioria proves to attain a shell length of 14.7 mm, which is considerably larger than the type material (length of largest paratype 8.30 mm). Apart from smaller size, P. morioria differs from P. anpoiiria in having more numerous radial ribs, rountl- ed instead of adapicallv shelved concentric ribs, finer, more numerous cusps on the radular teeth, and a smooth instead of serrate mantle edge. P. rnin-ioria and P. aitpouria have been taken li\ing together on the same piece of wood on two occasions. ACKNOWLEDGEMENTS I am grateful to the numerous fisheries scientists and obseners (MAFFish, Welhngton) and particularK' Miro Marinovich (Simuno\ich Fisheries, Auckland), who pro- vided the material. Thanks to Bmce Hawvard (Auckland Institute and Museum) for the loan of specimens, to James McLean (Los Angeles Countv Museum of Natu- ral Histoi-y) for comments on the manuscript, to Norman Heke (Museum of New Zealand, WelUngton) for the photographv and to Wendv St George (Institute of Geo- logical and Nuclear Sciences, Lower Hutt) for scanning electron microscope facihties. LITERATURE CITED Dall, \\'.H. lSiS2. On certain limpets and chitons from the deep waters off the eastern coast of the United States. Pro- ceedings of the United States National M\iseimi 4:400- 414. Lindberg, D,R, 1986, Name changes in the "Acmaeidae". The Veliger 29(2): 142-148. Lindberg. D.R, and C, Hedegaard, 1996, A deep water patel- logastropod from Oligocene water-logged wood of Wash- ington State, USA (Acniaeoidea; Pcctinodonta). Jonnial of Molluscan Stndies 62(3):299-314, Marshall, B.A, 1985. Recent and Tertiarv' deep-sea Hnipets of the genus Pcctinodonta Dall (Mollusca: Gastropoda) from New Zealand and New South Wales. New Zealand Journal of Zoolog)- 12: 27.3-282. Okutani. T. E. Tsuchida, and K. F'tijikura, 1992. Fi\e hatl]\al gastropods li\ing within or near the Cali/ptof^cna-com- niunit\ of the Hats\ishima Islet, Sagami Bd\. \'enus 51(3): 137-148. Powell, A.W.B. 1979. New Zealand Mollusca: Marine, land and freshwater shells. Collins, Auckland, .500 pp. THE NAUTILUS 112(2):58-62, 199S Page 58 Rediscovery of Discus hninsoni Beriy, 1955 and Oreohelix alpina (Elrod, 1901) in the Mission Mountains, Montana, with Comments on Oreohelix elrocli (Pilsbry, 1900) Paul Hendricks Montana Natural Heritage Program 1515 East Sixth A\enue Helena, MT 596:20 phen(lricks(S'nris.nit.gov ABSTRACT During sunniier 1997, extant populations ot the endemic land snails Discus brtinsoni and Oreohelix alpina were rediscovered near the type localities in the Mission Mountains, Montana; last known collections were made 33 and 48 years ago, re- specti\elv. Discus hniusttiii and Oreohelix elrodi (also endemic to the Mission Mountains and nearhv Swan Range) were pre- viously reported to be svnipatric in subalpine limestone talus. Both .species were found in 1997 in diorite talus; the latter species also occupied argillite talus with little, if any, limestone present at any site. Live D. hninsoni [n = S) were foinid near the talus surface only on bare rock or foliose lichen on rock during cool, wet conditions. Live O. elrodi were found near the talus surface on rock (n = 9) or organic litter accunuilations {n = 55) under all conditions, although in reduced number during warmer and drier weather Above treelme, live O. al- pina were found exclusively in exposed locations under lime- stone talus, sometimes attached to rock surfaces {n = 7) but more often present in organic detritus {n = 10). Lack of col- lecting records of D. hninsoni and O. alpina in recent decades probablv resulted from 1) extremely restricted distributions, 2) limited knowledge of habitat requirements, and 3) .scarcitv of active collectors in the region. Key ivords: Discus hninsoni, Oreotntix nlpina, Oreohelix el- rodi. Montana, distribution. ecolog\'. INTRODUCTION The Mission Mountains of northwestern Montana are recognized as an area of significant snail endemicitv in the Interior Columhia Ri\-er Basin (Frest and Joluinnes. 1995). Tile inoimtains extend about SO km north to south and 19 km east to west, rising ahruptK from a ha.se at 1036 m in the Swan and Flathead valleys to a crest of nearly .3()I 15 mm diameter. At Sites 2 and 3 the respective ninnbers of live snails in this category were 6 (33.3%) of 18, and 2 (28.6%) of 7. Sample,s\m- doubtedlv contained members of more than one cohort The presence of inchviduals < 7.5 mm diameter in each sample suggests reproduction is occurring at each site. Oiu'.ouiLix \i.ri\-\ Presence at different sites and .substrate selection: Weather on East St. Mans Peak was clear and cool (12.5"(; at the smnmit) on 27 August. I touuil no snails in 30 min (12:15-12:45) at Site 1 on the .southea.st ridge. At Site 2 I found 16 live O. (dpina in 45 min (13:30- 14:15) in three areas searched along 300 m of ridge. All live snails were imder limestone blocks about 20 X 30 ciri scpiare and 4-8 cm thick. Six snails were attached to the undersiiles ol linicstone Iragmenls or atop iiare rock beneath (deriving blocks, one group of 10 li\e snails was founil in leaf litter (area = 9 cm-) accumulated near the base of snow cinquefoil. Some soil development was present at each site where li\e suiiils were found. Dead shells were found with little effort on open ground. I searched the summit area (Site 3) for 15 min (14: 30-14:45) and found one live snail at 2865 m near the top of the southwest ridge. This individual was on bare rock under a limestone block near a patch of snow cinquetoil. Shell diameter: Diameter of h\e shells (mean ± SD, n = 16) was 5.7 ±1.9 mm; range was 2.5-8.5 mm. The smaller individuals indicate that reproduction is probably occurring at this location. Mean diameter of dead shells (n =9) was 8.9 ± 0.5 mm; range was 8.0-9.6 nun. DISCUSSION Extant populations of Discus bninsoni and Oreohclix al- pina were located in 1997 at or near the t\pe localities 33 and 48 vears, respectively, after the last docinnented collections (R. B. Brunson, pers. comm.; T J. Frest, pers. comm.). The range in shell size of live individuals at each site (including O. clrodi sites) indicates the pres- ence of multiple cohorts and likelihood of continuing reproduction. The sites where O. alpina was found on the southeast ridge of East St. Marvs Peak represent a slight range e.xpansion. (The type locality is on the south- west ridge.) I have foimd no evidence that O. alpina has been collected on East St. Mans Peak since 1900, when Elrod discovered the species. The last documented col- lection of O. alpina (Bninson specimen cat;ilog) was made on McDonald Peak on 2 August 1949. Correspon- dence from Stillman Bern (21 .\ugust 1951: Bnm.son pers. comm.) indicates that another collection mav have been UKide on McDonald Peak in 1950 or 1951. but tliere is no evidence of this in Bnmson's c;it;il()g Several intenicting factors prob;iblv contributed to the failure to find Discus bninsoni and Oreohclix alpina dur- ing the last several decades. First, I know of no active resident collectors of terrestrial mollusks. Non-resident collectors passing through the area m;iy luive visited sites when I'onditions were not espei i;illv tavonible tor finchng these siiccies ( D bninsoni in particular) near the surface p. Hendricks. 1998 Paee 61 of tains slopes. The nearlv halt-centim' span liehveen first tliscoveries of O cirodi and D hniiisoui at tlie same site supports tliis contention. Differences in lialiitat se- lection mav have contributed to this lapse, as will he discussed below. Furthermore, it seems unlikely that \er\' nianv non-resident collectors would attempt to xdsit the relati\elv inaccessible localities of O. ulphia. Second, the known ihstributions of both species are (|uite re- stricted. It would be easv for a collector to o\erlook them without knowledge of the geographical area. For example, one cannot identifv "Sin\aleamin Mountain" or find the exact location of O. aljiina on this moimtain or on McDouiild Peak without a c(jpv of Elrod's (1902) de- scription of his collecting trip. Third, habitat retjuire- ments, especiallv for D hniiisiyni. mav be narrower than pre\iouslv appreciated. Information gleaned from Beny (15^55) and Brunson (1956) suggests that D. bninsoiii and O. cirocli are ,s\nnpatric in talus slopes on the north side of McDonald Lake. Howexer. it ma\ be that both species co-occur in onlv a verx' restricted zone defined bv the narrower habitat preference of D Ijnnisoui Ev- idence to date indicates that O. el rod i is significantK more widespread, even in the McDcjnald Lake cinjue. All three sn;iil species favor exposed talus habitats. The t\pe of talus in which each species is found, how- ever, appears to differ among species. Orcohclix alpina is found exclusivelv above treeline in shallow limestone talus on mount;un ridges. The descriptions proxided by Elrod (1901; 1902; i903b) and Frest and Joliannes (1995) generallv coincide with my 1997 findings. Elrod (1902) commented that the snails were found among and imder rocks with little \egetation nearbv. All five imlixiduals I located were under the protection of stones in areas of scant \egetation cover, Init usnalK with some soil and litter accumulation nearby. Ten (62.5%) of 16 li\e indi\iduals that I could measure (one shell was bro- ken during handling) were sm;iller than the minimum diameter (7 mm) described for this species (Elrod, 1903b; Pilsbr)', 1939). However, size range of empty shells found mostly on open ground closelv matched the published range and mean. Perhaps adults are more Uke- 1\' to be caught awav from refuges in aiKerse conditions, with dead adult shells washing onto open ground Discus hninsoui and O. cirodi are found well below the treeline in t;ilus slopes surrounded bv closed- and open-canopv forest (Biimson, 1956), but the rock txpes comprising the talus inhabited hv the snails are infre- (juentK mentioned in pubfished reports. Berrv' (1955) and Frest and Johannes (1995) identified the talus as limestone; Elrod (1901; 1902; 1903a) and Bmnson (1956) failed to mention rock composition. I did not de- tect limestone at the three sites where I foimd one or both species. Discus bninsmii was found exclusivelv in talus where diorite boulders predominate. Orcoliclix ci- rodi was found at this site as well as in smaller-sized talus of argillite (Table 1). Discus bninsoni may have narrower habitat recjuirements than expected, associating only with a particular subset of a\ailable rock types. The biologv and ecological requirements of Discus hninsoui and Orcohclix cirodi remain largelv unknowii, antl the following explanations for patterns of presence or absence near the surface of talus slopes are specula- ti\e. Orcohclix cirodi was more abundant (4-6 fold dur- ing mv searches) than D hruusoni near the surface of talus in wet and cool conditions, and a few indi\iduals could still be found when it was warmer and drier. Sev- eral factors could contnbute to this pattern. First, ab- solute population size of O. cirodi at Site 1 may be great- er than that of D, hruusoni. Ratios of each species near the talus surface could be representative for all depths in talus. Second, I found significant tlifferences in sub- strate selection between the hvo species. Some D. hnin- soui were found on foliose lichen (tentatively identified as Arcfopunncliii suhccntrifu^a) that grows on the dio- rite, but most uuli\iduals were found on bare rock. In contrast, O. cirodi were most often found on organic litter and \egetation. Preference by O. cirodi for organic litter mav keep them nearer the surface of talus slopes where fitter accinnulations are larger and apparentlv more numerous. Thnil. larger shell size of O cirodi might reduce its rate of desiccation bv decreasing the surface are;i/\'olume ratio (see Goodfriend. 1986), allow- ing larger individuals to remain nearer the drier talus surface for longer periods than D. hninsoui and small O. cirodi. The four live O. cirodi found on 10 June in warm and diT conditions measured 18-21 mm diameter and were estivating at that time. Fourth, timing of m\ searches did not allow for detection of this pattern, but D. hntihsoni may tend to be nocturnal or crepuscular and make \'ertical migrations to the talus surface during pe- riods of acti\its' (Bnmson, 1956). ACKNOW'LEDGM ENTS Field work was generonsK supported bv a Canon Ex- ploration Grant aihnmistered by The Nature Consenan- cy and Canon U.S.A., Inc. I benefited greatly from con- versations with T. J. Frest and especiallv R. B. Bnmson, whose knowledge and documentation of the snails of the Mission Mountains were enthusiasticallv shared when- ever requested. J. S. Marks and L. M. Hendricks partic- ipated in the hunt for Discus hninsoui; T Gignoux iden- tified the rocks at the D. hninsoui site. T Shreve \isited the tvpe localiU of Orcohclix alpino with me. An earlier draft of the manuscript benefited greatlv frf)m the com- ments of Uvo anonvmous reviewers. This paper is decfi- cated to the memor)' of the late J. R. Reichel of the Montana Natural Heritage Program, who supported my efforts even before this project was conceived. LITERATURE CITED Alden, W. C. 1953. Phvsiographv and glacial geoIog\- of western Montana and adjacent regions. U. S. Geological Survey Professional Paper 231:1-200. Ausden. M. 1996. Invertebrates. In: .Sutherland, W. J. (ed.) Ecological census techniques, a liandbook. Cambridge Universitv Press. Caniliridsie. pp. l.'59-177. Page 62 THE NAUTILUS, Vol. 112, \o. 2 Berrv', S. S. 1955. An important new land-snail from the Mis- sion Range, Montana. Bnlletin ot the Sontheni Calitoniia Acadeniv of Sciences 54:17-19. Brunson, R. B. 1956. The nivsterv of Di.sais hninsoni. The Nautilus 70(1):16-21. Elrod, M. J. 1901. Montana shells. Rocky Mountain Magazine 2:691-697. Elrod, M. J. 1902. A biological reconnoissance in the \icinit\- of Flathead Lake. Uni\ersitv of Montana Bulletin No. 10, Biological Series No. 3. Elrod, M. J. 190.3a. Notes on Pi/raniirluhi clrodi Pils. The Nau- tilus 16(10):109-1 12. Elrod, M. J. 190,3b. Montana shells — Pi/nniiidiilri strii^^osa. The Nautilus 17(1): 1-6. Fairbanks, H. L. 1984. A new .species ot Onohclix (Gastrop- oda: Puhnonata: Oreohelicidae) from the Se\en De\-ils Mountains, Idaho. Proceedings of the Biological Societ\' of Washington 97:179-185. Frest, T |., and E. J. Johannes. 1995. Interior Cohunbia Basin mollusk species ot special concern. Final Report to Inte- rior Columbia Basin Ecosystem Management Project. Dei.\is Consultants, Seattle, 274 pp. Goodfriend, G. A. 1986. Variation in land-snail shell fonn and size and its can.ses: a resiew. S\stematic Zoolog\' .35:204- 223. Pilsbry, H. A. 1939. Land moUnsca o\' North Amenca (north of Mexico), volume 1 part 1. Monographs of the Academx' of Natural Sciences of Philadelphia (3), Philadelphia, 57.3 pp. Sokal, R. R., and F J. Rohlf 1981. BiometrA. Second Edition. VV. H. Freeman, San Francisco, 859 pp. THE NAUTILUS 112(2):63-68, 199S Page 63 Epitonium fabrizioi (Gastropoda: Epitoniidae), a New Species from Patagonia, Argentina Cuido Pastorino' Departaniento Paleozoologi'a Invertebrados Museo de La Plata Paseo del Bosque s/ii 1900 La Plata ARGENTINA i"vpastor@criba. edu. ar Pablo Penchaszadeh INTECMAR Universidad Simon Bolivar 1080 Caracas VENEZUELA and Museo Argentino de Ciencias Naturales "Beniardino Rivada\ia" Angel Gallardo 470 1405 Buenos Aires ARGENTINA ABSTRACT Epitonium fabrizioi, a new species of gastropod mollusk be- longing to the family Epitoniidae, is described from shallow waters of Puerto Piraniides, Chubut Pro\ince, Argentina. This new species is similar to E. gcorg,fttiniim from the same area, which was known only from its shell. It can be distinguished from E, ^eoracttimim by its smaller size, more straight profile antl, b\' its sharjier, most numerous ribs. In addition, the pro- toconch of the new species consists of 4.25 whorls. Its radula has only marginal teeth, each with three cusps. The outermost cusp is larger and hook-like, the central and inner cusps are shorter anil similar in size. An additional, ob.solete, cusp is pres- ent near the base ot the teeth. The egg capsules ha\e a mean diameter of 76 \).m. The new species is compared with £. ^cor- ^ettinuui. the species with which it co-occurs, and £. albidnm. INTRODUCTION Most recent papers about the taiinlv Epitoniidae deal with .species from the northern hemisphere (CJarihliean: Robertson, 1983a, 1983b, 1994a, 1994b; northeastern Pacific: DuShane, 1974, 1979; northeastern Atlantic: Bouchet and Waren, 1986). An e.xception is the work of Kiiburn (1985) who studied representatives ot the taniilv from South Africa and Mozambicjue, and provided a good account of the subgenera living in that area. In their classic and comprehensive work. Clench and Tur- ner (1951, 1952) reported most of the knowii western Atlantic species. Rios (1994) included a complete list of the approximately 30 species of Epitoniiilae knowm from northern South America, together with illustrations and distribution ranges. However, the material examined in ' Current Address: Department of Invertebrate Zoolog)', Di- vision of Mollusks, National Museum of Natural History, Smithsonian Institution. W'ashinsTton, DC 20560, USA. his work was primarily from Brazil, and only margmally included representative species from Uruguay and northern Argentina. The Magellanic area is poorly represented in mollus- can faunal reports, and almost never represented in pub- hshed revisions of Epitoniidae. Strebel (1905) was the last author to publish a systematic account of the Mag- ellanic epitoniid fauna. He described one species and one variety from the Strait of Magellan. In this paper we describe a new species of Epitonium, and include illustrations of its radula, operculum and protoconch. Data on eggs and egg capsules are also in- cluded. MATERIALS AND METHODS Specimens of Epitouiitin fabrizioi were collected from tidepools near Puerto Piramide, Chubut, Argentina (42°34'S, 64°17'W) in November, 1995, during .spring (lower low water) tides, and near Punta Loma (42°49'S, 64°53'W) in February, 1996 (see Map 1). This species was found only in the intertidal zone around the pedal discs of sea-anemones (genus Biinndoctis?) on which they probably feed. In several cases, more than one specimen was found on the same sea-anemone. Several animals were dissected and the radulae were prepared for the SEM obsei'vation. The protoconch whorls were counted folltming the method of Jung (1986). Institu- tional abbreviations used are: ANSP, Academy of Natural Sciences of Philadelphia; MACN, Museo Argentino de Ciencias Naturales "Bernardino Rivadavda", Buenos Ai- res, Argentina; MHNC, Museum d'Histoire naturelle de Geneve, Switzerland; MLP, Departaniento Zoologia In- vertebrados, Museo de La Plata, La Plata, Argentina; USNM, National Museum of Natural Histoiy, Smithson- ian Institution. Page 64 THE NAUTILUS, Vol. 112. No. 2 Figures 1-9. Epitoninin jahrizioi new .specie.s. 1-3. Holotvpe, MLP 54():i, .Scale bar = 5 nnii. 4. Apical \ie\v, MLP 5.3.3.3. Scale bar = 200 (xiii 5. Protocoiicli, MLP .5.3.33. Scale bar = 2.50 |j.rri. 6. Operculum of holoUpe, Scale bar = 5 mm. 7. Radula of liolotype in '.^eiieral \ie\v. Scale bar = 20 |xm. 8. Detail of luar'Tinal teeth. Scale bar = 10 jjim 9. Enn capsules. Scale bar = 1 nun. SY.STEMATICS KaiTiilv Epitoniiilac BcriA', 1910 Cenu.s Epitoninin K()cliiii», 1798 Epitoniiini fahrizioi new specie.s (Figures 1-9) Epitoninin alhidum (d'OrbifiUN. bS42,). — Ciencli .mil 'runiei, 1951: pi. 114. fi" 3 only. ' Epitonmni '^rorui'ttiiia (Kiener. 1S.39). — Scarabuio, H)7T:1.S3, pi. 2, fig. 8. Epitoninm '^eori^fttinai' (Kiener, lS.39i, — Paslonno, 1995:S, pi. 2. fig. 10, Dcseriptioii: Shell sinali, iij) to 1.3 nun leniitii, clialkT, thill, opaque, Frotoeoneli iWoni in all but oni' spei-inien) eon.sisting of 4.25 whorls and nie;isuring 4S5 length X ,390 fjLm width. Protoeoneh whorls smootli imder SEM, tran- sition to teleoeoneh not vei"V' shaip. Teleoeoneh with S gentK' convex, smooth whorls. Spire angle 40°. profile strmght. Suture crossed li\ tenuination of a\i;il ribs (fenes- trate). A.\ial ornamentation cousi.sting of vei"v shiup ribs, up to IS on last wiiorl, hut usuiillv l.'3-14 (Tal)le 1). A.\i;il ribs slighlK o!)li(ine in relation to shell a\is, with iiTegularlv de- fined edges near sntine, and ri'mnants from breakage in the r("st. .\.\ial ribs gentlv ri'fleeted on last whorl. y\ll ribs perteitK iiligned witli overlapping ribs from preceding whorl; attachment erect. Ajiei+nrt' o\al, penstome with one la\'er, forming a getitle basal expansion (aunculate). Colu- mell.ir callus weakK' deNcloiied. L'mbilicns i-lo.si'd. G. Pastorino and P. Penchaszadeh, 199S Page 65 Puerto Madrvtf*-mf Lonia -42 S P Ninfas to 5 0 10 ' ■ ' Map 1. Records o( Epitoniuin fabrizioi (® = t\pe localit)') Operculum oval, paucispiral, thin, translucent, hrown- i.sh, entire surface co\'ered b\' growth lines. Radula lacldiig central and lateral teeth, but with nu- merous tricuspid marginal teeth, identical in shape. Out- ermost cusps large and hook-hke, central and inner cusps similar in size and shoiter Secondar\' cusp present centrally on teeth near base. Egg masses always found near (but not attached to) adults. Egg masses composed of 180-320 egg capsules attached to each other b\- tough elastic string as in other Table 1. Epitnitiiiin fabrizioi Shell measurements and number of axial ribs per whorl. Numbered column headings refer to whorl munber, Duslies indicate worn nbs on tliat wliorl. Shell No Lens \N'idt!i Teleo- concli whorls 1 10,92 5.30 5.5 2 11.7 5.9 6.5 3 15.01 6.92 6.5 4 fi.S9 3.32 5.5 5 1.3.22 6.07 6.5 6 12.10 5.85 6.5 7 8.68 4.45 6 8 13.05 6.54 6.5 9 7.89 4.1 6 10 12.91 6.14 6 11 12.27 5.56 6 12 12.57 6.15 6.5 13 9, IS 4.49 6.5 14 10.66 5.30 7 15 9.72 4.92 6.5 16 11.78 5.76 5.5 17 10,94 5.14 5.5 18 6.:57 3.20 6.5 19 9.10 4.77 5.5 20 10.52 5.01 6.5 21 10.17 4.95 6.5 22 9.23 4.54 6.5 23 8.48 4.57 5.5 24 11.50 5.36 6,5 25 9.96 5.08 i — 16 15 15 13 13 13 13 14 — 14 14 14 15 13 13 13 14 13 13 13 13 15 13 13 13 13 14 — 14 14 14 14 — 12 12 13 15 — 12 12 13 14 — 14 14 13 15 13 13 13 13 14 14 12 14 14 15 13 12 13 13 14 — — 15 15 15 13 14 14 14 15 — 14 13 14 — 14 14 15 — — 14 13 14 14 14 14 15 — 14 14 14 16 — — 14 14 16 13 13 13 13 14 — 15 15 16 — — 15 15 16 — — 14 14 17 16 IS Page 66 THE NAUTILUS, Vol. 112, No. 2 Figures 1()-16, 17-21. EjnUiminn ficor<^cttiiiiiiii (Kit-iu-r, 1S.391, 10-12. A[ii-rtunil, basal, ami a]iic.il \it-\\s. MLP 54(14. Puerto Piranuiles. Scale bar = 5 niiii. 13. Operculum of a 24.72 mm length specimen. Scale bar = 10 nnii 14. Scalaria ^corgeftina, paralectohpe, MHNG 98.3/111/2, "Ocean Atlantique". Scale bar = 10 mm. 15. Raduia of the specimen shown in F'igiires 10-12. Scale bar = 20 (xni. 16. Detail of the railula. Scale bar = 10 jjim. 17-21. Epihmhim albidum (d'Orbigiiy, 1842). 17^ 18. USNM 4.39556, Baie Anglai.se, Haiti. 17. Apertural \iew. Scale bar = 1 nnn 18. Detail of the shell surface. Scale bar = 10 (jLm. 19. Detail of the same area under higher magnification. Scale bar = 2 fim 20. Protoconch. Scale bar = 40 jjim. 21. Apical view. Scale bar = 40 |xm. species of sanK' gi'inis. Largest egg capsules iwaminci! measured 1..35X1.1 nmi (ii = 2(); x= 1 .19X0.94; Sn = 0.()S()X().()69). Egg capsules pyramidal or polyhedral in .shape and covered with sand grain.s. Nmnher of eggs per capsule ranged 69-141 (n=l(); x= 123.9; SD = 2().12). Uncleaved eggs measured 76 \i.m in diameti'r. Type locality: Puerto I'inimides. (-'huhut Pro\iuci-, Argentina (42'°34'S, 64°]7'VV). T\pc material: nolot\pe. MLP 5102; 10 paratopes, USNM SS0252; I parat\pi>, MLP 5.33.3; 4 parahpes. ANSP A1S86(); 10 paratypes MLP 5403; all from the t\pe loc;ilit\-, (;. Pastorino coll., November 1995; MLP 5333, 1 specimen (protoconch), from near Punta Lonia, Chubut Province. Argentina, 42°49'S, 64°53'\V, J. Mer- moz coll., Februarv 1996. Other material e.xamined: Five specinien.s. MACN muuunbered, trom near Punta Loma. Chubut Province, Argentina, 42°49'S, 64°53'W, J. Mermoz coll.. Feb. 199(i. Etviiioloijy : Dedicated to Fabiizio Scar;ibino, voung niiihii'ologrst ;uul Iriend, who called our attention to the new siH'cies. G. Pastorino and P. Pencha.szadeh, 199.S Page 67 Table 2. Epitoniiii}! acorvcttiiiiim (Kieiiei). Shell measurenient.s and number ot axial nbs per whoil. Nuniberetl tohuini headings refer to whorl rnniibei' Dashes indicate uoni ribs oir that uhorl. Teleo- Shell conch No Length Width whorls 1 2 ■i 4 5 6 / 8 9 1 31.13 2 24.75 1 26.73 4 20.81 5 18.02 6 14.79 10.64 8.90 9.72 8.33 6.73 5.86 9 8.5 9.5 9 9 8 DISCUSSION — 16 16 16 14 13 13 14 14 13 12 13 12 13 14 — 13 15 13 13 12 12 13 13 13 12 13 12 12 11 12 — 12 12 12 11 10 11 11 — 13 1 '^ 13 12 12 12 Epitoiiitim gcorgetfiuuin (Kiener, 1839) is the most sim- ilar species to E. fabrizioi. It also occurs in the httoral zone in the Puerto Pinimides area, where it lives near sea anemones on hard sulistrate, but, unlike E. jahhzioi. it can also be found on sandv bottom. The type specimens of Epifoniitm gcorgcttiiuiin (MNHG' 983/11 1/1-3) were illustrated bv Clench and Turner (1951, Pi. 117, fig. 1). Figure 14 herein repre- sents one of the paralectotvpes (MHNG 98.3/111/2). The main conchological differences between E. gcorgettiiunn and the new species is the profile of the shell, which is straighter in £ gcorgcttiniiiii. and the whorls, which are more con\ex and detached in this latter species. The rilis in E. fabrizioi are sharper, thinner, and increase in num- ber with growth (Tables 1, 2). Epitonium gcorgcttiniim has a multispiral, thick, and opaque operculum (Figure 13), whereas that of E. fabrizioi is paucispiral, thin, and translucent (Figure 6). The radula of E. georgcttinitm is also distinct. It has one sharp terminal cusp per marginal tooth, and two lilimt cusps, one almost obsolete (Figures 15-16). The eggs and egg capsules ol these two species are quite similar (Pastorino and Penchaszadeh, in press). However, based on the material available, £. fabrizioi has smaller egg capsules, each containing fewer eggs al- though egg diameter is nearh' identical to that in E gcorgctiiniiiu Bell (1985) has indicated the lar\al fife- span and size in E (//;/ Pilsbrv, 1921 (26 days; 3 whorls, 390 |jLm). The larger number of protoconch whorls and size oi Epitonium fabrizioi (4.25 whorls, 485X390 |xm) suggests that E. fabrizioi spends more time than E. (//(/ in the planktotrophic lanal stage. In comparison with other known species of Epitonium, the new species has the largest protoconch (Table 3). Epitonium albidum (d'(Jrliigny, 1842) is suuilar in shell shape, liut its protoconch shows a deficate (visible under SEM) ornamentation that contrasts with that in the new species (Figure 20), In adcfition, the entire sur- face of the teleoconch in E. albidum shows a character- istic pitted microsculpture (visible under SEM; Figures 18, 19). This character is not t)pical of the genus Epi- tonium and may support a new generic allocation for E. gcorgettiiunn, as suggested bv Bouchet and Waren (1986) for E. albidum^ In contrast, E. fabrizioi lacks this h-pe of microsculpture. Robertson (I983ii) pointed out that Clench and Turner's (1951) Argentinean record of E. all'idum from Bahia San Bias was zoogeographicallv anomalous. Their illustration (Clench and Turner, 1951; Plate 114, fig. 3) represents a specimen of E. fabrizioi. Kilburn (1985) proposed a subgeneric arrangement for the South African .species of Epitoniidae. He did not consider, however, that radular morpholog)' could pro- \ide rehable taxonomic characters to his arrangement. If we followed Kilbuni's classification, E. fabrizioi could be allocated in the subgenus Hirioscala Monterosato, 1890. However, E. fabrizioi does nf)t have corcjnate lamellae and a duplicate peristome that are characteristic of that subgenus. Including E. fabrizioi. there are 5 named species of Epitoniidae living in the Magellanic prn\ince. Keen (1971) illustrated 76 Epitoniidae from the tropical east- em Pacific; Kilburn (1985) cited 80 species from south- em Africa and Mozamlii(|ue; Rios (1994) recorded 30 Table 3. Measurements of protoconchs of Epitonium species in relation to the adult size (in mm' Epitoniiiiii speci Protoconcl Whorls Aduli Protoconch size leirgth-width Source E. millccostation 2.7 9.7 0.5 Robertson, 1981 E cijiiinaticostd 3.5-5 9.5 0.9-0.3 Robei-tson, 1983a & E albidum 4 8-15 0.4 Robertson, 198.3b E. phipnanthi 3.2-;3.4 16.9 0.42-0.5 Robertson, 1993 E uorsfohli 4.1^.2 25.3 0.52-0.54 Robertson, 1993 E. iilii 3 13 0.39 X 0.30 Bell, 1985 E. fabrizioi 4.25 13 0.48 X 0..39 this paper 19941) Page 68 THE NAUTILUS. Vol. 112. No. 2 from Brazil, while Diaz-Merlano and Pu\ana-Hegedii,s (1994* listed 22 from the Colombian Carihliean. Based on pubhshed records ot the known distribution of Epi- toniidae this famiK' is more speciose in tropical than in temperate and cold regions. ACKNOWLEDGMENTS Special thanks are due to f. Mermoz, wht) pr()\ided specimens of Epitoninin from Punta Loma. Yves Finet, Museum d'Histoire naturelle de Geneve, Switzerland, kindly sent photographs of the paralectotype of Epiton- ium ^corin'ffinum. M. G. Harasewvch, Nation;il Museum of Natural HistoiT, Smithsonian Institution, gave good ad\ace on earlv stages of the manuscript. Two rcNdewers improved considerablv the original manuscript. This work was made possible b\' an external fellowship grant- ed bv the Consejo Nacional de Investigaciones Cienti- ficas'y Tecnicas (CONICET), Argentina, to G. P and a grant from Funilacion Antorchas, Argentina to P. P. LITERATURE CITED Bell, ]. L, 1985. Lanai growth and metanioqihosis of a pro- sobranch gastropod associated with a solitary coral. Pro- ceedings of the Fittli Inteniational Coral Reef Congress 5:159-163. Bouchet. P and A. Waren, A. 1986. Re\ision of the Northeast Atlantic bathyal and abyssal Aclididae, Euiimidae, Epito- niidae (Mollusca, Gastropoda), Boilettino Malacologico, Supplemento 2:498-523. Clench, W. J. and R. D. Turner. 1951. The genus Epitonium in the western Atlantic (Part 1). Johnsonia 2(301:249-288. Clench, W. J. and R. D. Turner. 1952. The genus Epitonium. (Part II), Depressisc/ila. Ci/lindriscala. Ni/stiella and So- hiti.sailii in the western Atlantic. Johnsonia 2(31 ):289-35fi. Dfaz-Merlano, J. VI. and M. Pnvana-Hegediis. 1994. Molusc'aribbean coast of Colombia. Cosel (1978, 1986) and Diaz (1985, 1990), who studied the molluscan fauna from ibfferent localities of the Colombian coast, first noticed the occurrence of these unnamed Pcriploina. Finther material of one of these species has become availalile to the authors in re- cent vears through intensive collecting near Santa Marta. Thcv are herewith described as new species. Institution- al abbreviations are: BMSM, The Baile\-Matthews Shell Museum, Sanibel, Florida; INVEMAR MOL, Collection of Mollusks, Instituto de Investigaciones Marinas y Cos- teras, Santa Marta, Colombia; SMF, Senckenberg Mu- seum Frankfurt, Cermanv. SYSTEM ATICS Class Bivahia Linnaeus, 17.58 Subclass Anomalodesmata Dall, 1889 Superfamily Thracioidea E.A. Smith, 1885 Familv Periplomatidae Dall, 1895 Genus Paiploma Schumacher, 1817 Periploma [Pcriploina) coseli new species (Figures I, 2, 5) Periploma spec. — Cosel, 1978:161. pi, 4, figs. 12,1.3. Pcriploina spec, II. — Diaz, 198.5:84, erroneously figured in pi. 12, fig.l. Periploma sp".— Cosel, 1986: 199, fig. 11.3 Periploma sp, 1.— Diaz & Puyana, 1994:104, pl.:31, fig. :30.3. Description: Shell merhum-sized (length up to 29.3 mm), subcircular-oval, ven- thin and fragile, moderately convex, antero-ventral margin evenlv circular posterior margin scjmewliat projected upward and subtnmcate. Subecjuilateral, anterior end slightly larger, beaks with characteristic tran.sversal slit. Outer surface white, with irregular growth lines and numerous, extremely fine, ir- regular radial striae more apparent in the central area. Periostracum diity gray, with comarginal rows of micro- scopic granules. Inner surface weaklv nacreous, smooth but with weak impressions of growth lines. Palfial sinus broad and short, reaching to about 1/3 of shell length. Ligament internal; resiliter conspicuous, spoon-shaped, thrected vertically toward center of valve and weakly re- inforced at its basis b\ cui-vetl buttress pointing to pos- terior muscle scar Type material: Ilolohpe, SMF 311857, complete shell (left valve slightly broken), 29.3x25.2 mm (lengthxheight), trawleil h\ RA' Ancdn at tspe locality, C.P Arango coll. September 1995. Paratvpes: INVE- MAR MOL-1151, 1 complete shell (left valve fractured near midline), 24.8X20.0 mm, Golfo de Salamanca, Co- lombia, ir07'N, 74°20'W, trawled hv WW Ancdn, 59 m, muddv sand, C.P Arango coll. September 1995; INVE- MAR MOL 1152, 1 complete shell (left valve damaged), 10.5X8.5 mm and BMSM 2361, 10.9X8.8 mm, 1 com- plete shell (left valve damaged), off mouth of Toribio Page 70 THE NAUTILUS. Vol. 112. No. 2 ^ / Figures 1—1. 1-2, Pcriploma roscli. Hol()t)pe, riglit \al\e, 29.3 mm length, 25.2 mm lielglit, 3—4, Frriphtuia sanrtmimiihncnsis. HoloUpf. left \alve. 21.9 mm length, 17.5 nnii height. river. ir04'N, 74°15'W, taken by bottom grab, 6 in. mud, N. Ardila coll. Octolier 1996;' BMSM 23fi(). 1 com- plete .shell, 16.8X14.2 mm. off mouth of Toiihio Ri\er, 1I°()4'N, 74°15'\V, bottom grab, fi m. imid. N. Ardila coll. Jamiar)' 1998. Type locality: Colfo dc .Salamanca, (>'olond>ia. ini'N, 74°15'W, 59 m, muddy .sand. Etymology: Named altci- I-tiido xon Cose), who fiist noticed the existence ol this species in his comprehen- sive study of the molhiscan fauna of the ('aril)bean coast of Colombia. Remarks: PcripUiinu roseli new species is similar to F. discus Stearns, 1890, from the eastern Pacific, and to P. camerunensis Cosel, 1995, from off west Africa. The postero-dorsal margin of P. discus is almost straiglit (somev\hat projected dorsalK' in Pcoscli). and its posterior margin more angulale ibroadK rounded in the new species) (compare figures 5 and 7). Pcri- ploma caincnincnsis exhibits a somewhat more elon- gatetl shell, with the umbones situated posteriorh' to the vertical midline. In addition, the resilifer in P. ca- incnincnsis is projecteil obliquelv. in relation to the lunge line and toward the antero-\entral margin, whereas it is almost perpendicular in relation to the hinge line in P coscli (compare Figures 5 and 8). These 3 species may be regarded as members of a group ol similar taxa that show Recent disjunct distri- bution. Isolation ol eastern Pacific and eastern Atlan- tic anci'Stral populations could ha\e occurred respec- ti\('K' alter establishment of a continuous Isthmus of Panama and a lull-Hedged mid-Atlantic oceanic bar- rier. Absence ol planktotrophie de\elopment in the genus Pciiploina (see (ioodsell ct al. 198.3) ma\' ha\e laeilitated isolation and subse(juent speciation. P CDipicHiic .\ltena. 19tiS Irom the iiortlu'in and N. E. Ardila and J. M. Diaz, 1998 Page 71 Figures 5-11. Diagrams ot'tlie inner surfaces of left \;il\es of several species o( Pciiploinu 5. f ((iscli Holohpe, 6. P snmiaiiuiiihiwnsis:. Holohpe^ 7. P disriis Steams, 1890. retlrawii from Keen (lOThfig. 750). 8. P caiiirnincmis Cosel. 1995. Holohpe reth-awii from Cosel (1995:fig. 144). 9. P coqucttac Altena. 1967. Holohpe redrawn from Altena (1967:fig-147a). 10. P filnnUiscuhnn ,Sowerb>, 18:34, retb'awi from Keen (1971;fig, 7.52). 11. P hii^aiHUa Olsson, 1961. Holohpe reclrawii from cils.son (196!:pl. 82, fig. .5h). northeastern coa.st.s of South America (off Suriname and Colombia), i.s another vaguely similar specie.s, hut it has definitely a more elongated and inequilateral shell (compare Figures 5 and 9). P. coscli seems to he \\idel\' tlistrihuted ;dong the Carih- bean coast of Cok)mhia. Cosel (1986) recorded diis species as "Pcriploum spec." from off Punta Broijueles (approxi- mateh- 9°2()'N, 76°1.5'W) and die Santa Mmta Baw and Diaz (1990) found it iJ.so ;ls north as Poitete Ba\- (12°18'N. 71°55'W) Page 72 THE NAUTILUS, Vol. 112. No. 2 Periplonta (Pcriploina) sanctanuuihacims new specie.s {Figures 3, 4, 6) Periplomn spec. I. — Di'az, 1985:84, pi. 12, fig. 2. Periplomti .sp. 2.— Diaz & Puvana, 1994:104, pl.32, fig. .304. Description: Shell niedimn-.sized (length to 22 mm), .semicircular-ov;il, equilateral, convex, very fragile; an- tero-ventral margin semicircular; posterior margin sub- truncate, postero-dorsal margin strtiight. Beaks with a characteristic tran.s\ersal slit. Outer surface milk\' white, with irregular growth lines, some of which appear as shallow grooves under magnification. Radial ridge nms from beaks to postero-ventral margin. Periostracum vel- lowish-gray, with numerous microscopic granules which are arranged comarginalK- and are more aliuntlant within growth lines and posterior slope. Inner surface dirts- white and weakl\' nacreous, showing impressions of ir- regularities on outer surface. Pallial sinus broadK trian- gular in outline and rather deep, reaching to about Vi of shell length. Ligament internal, in well-defined, scjuarish resililer. Resiliter oblique in relation to hinge line, ori- ented toward ventral margin. Type material: Holot>pe, SMF 311S58, 21.9X17.5 mm (lengthxheight), 1 left valve, from t\pe locality, SCUBA, J.M. Diaz coll. Febman 1983; Parat\pe, IN- VEMAR MOL 1153, 13.4X11.3 mm, 1 right valve (bro- ken in two pieces after measurements), from type local- itv-, SCUBA, J.M. Diaz coll. Februaiy 1983. T^pe localit>': Bahia Nenguange. Ta\Tona Natural Na- tional Park. Colombia, ir20'N'," 74°()5'VV, 7 m, coarse coralline sand with broken shells. Etymology: The species is named after the citv oi Santa Marta. near which the Tavrona National Natural Park, the type locality, is located. Remarks: The shell of Periplomn saiictamaiihaciisi.s differs from that of P. coseli in being smaller and more inflated. In addition, the outline of the antero-\entral margin is gentlv rounded, has a strmght postero-dorsal margin that does is not projected dorsallv. and a railial ridge nnming obliquel)' along the postero-ventral slope. Additional differences between these species do exist in the outline of the pallial sinus and the shape of the re- silifer (compare Figures 5 and 6). Pcriploma planiiisciilutn Sowerbv. 1834. from the eastern Pacific, has a larger and more elongated shell with a short, rounded pallial sinus (broadly triangular in P. saiirtaiiuu-tliaciiMis) (compare Figures fi and 10). Pir- iploina laoarlilla Olsson, 1961, from the Pacific coast of Panama, is also more elongated, and exhibits a rather pointed posterior margin and a short and narrow pallial sinus (compare Figures 6 and 1 1 ). P. stmrtaiiwiiluiciisis new species is so iai' known oni\ from the t>pc locality. ACKNOWLEDGMENTS We wish to express our gratitude to Claudia P. Arango for providing the type material of Pcriploma coseli new species, and to Dr. Eugene V'. Coan for his kind help with the literature search. The personnel and facilities of the Reference Collection at INVEMAR, Santa Marta, made possible the assemblv and improvement of pho- tographs and drawings. Two anonvinous reviewers con- tributed to the improvement of the maiuiscript. We ac- knowledge the economic support of COLCIENCIAS — BID (grant No. 2105-13-079-97). This is contribution No. 601 of INVEMAR. LITERATURE CITED Alteiia, C. O. van Regteren. 1968. The holocene and recent marine bivalve Mollusca of Surinam. Studies on the Fauna of Suriname and other Guyanas 10(42):15.3-179. Bernard. F.R. 1989. Living Periplomatidae of the Pacific and Indo-Pacific regions, \emis 48(1): 1-11. Cosel, R. \on. 1978, Die .Mollusken der Cieuaga Grande de Santa Marta (Kolumhien) uud ihre Lebensgemeuischaften im Wechsel der Jahreszeiten. Ph.D. Dissertation. Univer- sity of Giessen, West Germany. 348 pp. Cosel, R. von. 1986. Moluscos de la region de la Cienaga Gran- de de Santa Marta (costa del Caribe de Colombia). Anales del Institute) de Investigaciones Marinas de Punta de Be- ti'n (15-16):79-^370. Cosel, R. von. 1995. Fiftv-one new species of marine bivalves from tropical West Africa. Ibenis 13(1): 1-1 15. Diaz, J.M. 1985. Mollusken iind ihre Gemeinschaften in der Bahi'a de Nenguange (Karibik. Kolumbieni. Ph.D. Disser- tation, Uui\'ersitv of Giessen, West Germany. 305 pp., 29 pis. I^iaz. |,M. 1990. Malacofauna subfosil v reciente de la Bahi'a tie Portete, Caribe colombiauo, con iiotas sobre algunos fosiles del Terciaiio. Boletin Ecotropica 23:1-22. Diaz, J. M. y M. Puvaua. 1994. Moluscos del Caribe Colom- biano, un catalogo ilustrado. COLCIENCI.^S-Fuudacion Natura-IN\'EMAR. Santafe de Bogota. 291 pp., 74 pis. Goodsell, J.G.. R A, Lutz. M Castagna and J. Kraeuter 198.3. Nonplanktotropliic development of two species of conti- ueiitnl shell bi\alves Journal of Shellfish Research 3(1): 91 Keen, M. 1971. Sea shells of Tropical West .Vmeiica. Stanford University Press, Staufoixl. 1074 pp. Olssou. A. A. 1961. Mollusks of the eastern tropical Pacific, particularly from the soutlieni halt of the Pauamic-Pacific laiinal province (Panama to Peru). Pananiic-Pacific Pele- cvpoda. Paleontological Research Institution, Ithaca. 574 pp., 86 pis, Rosewater |. 1968. Notes on Periplomatidae iPelecvpoda: .\ii- omalodesmata), vritli a geographical checklist. Annual Re- ]icirt 111 llic .\mi-ncaii .Malacologic.il t'niou tor 1968: 37- 39. INSTRUCTIONS TO AUTHORS THE NAUTILUS publishes papers on all aspects of the biologv- and systeinatics of mollusks. Manuscripts describing original, unpublished research as well as review articles will be considered. Brief articles, not exceeding 1000 words, will be published as notes and do not require an abstract. No- tices of meetings and other items of interest to malacolo- gists will appear in a news and notices section. Manuscripts: Each original manuscript and accompanying illustrations should be submitted in triplicate. 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Box 1580, Sanibel, FL 3.3957, USA. @ This paper meets the requirements of ANSI/NISO Z39.48-1992 (Permanence of Paper). THE NAUTILUS Volume 112, Number 3 October 15, 1998 ISSN 0028-1344 A quarterly devoted to malacology. oLi zz ]998 EDITOR-IN-CHIEF Dr. Jos^ H. Leal The Bailev-Matthews Shell Museum 3075 Sanibel-Captiva Road Sanibel, FL 33957 EDITOR EMERITUS Dr. M. G. Harasewych Department of Invertebrate Zoolog)' National Museum of Natural Histor)' Smithsonian Institution Washington, DC 20560 CONSULTING EDITORS Dr. Riidiger Bieler Department of Invertebrates Field Museum of Natural History Chicago, IL 60605 Dr. Arthur E. Bogan North Carolina State Museum of Natural Sciences Raleigh, NC 27626 Dr. Robert T Dillon, Jr. Department of Biologv' College of Charleston Charleston, SC 29424 Dr William K. Emerson Department of Living Invertebrates The American Museum of Natural History New York, NY 10024 Dr Eileen H. Jokinen Institute of Water Resources University of Connecticut Storrs, Ct 06269-4018 Mr. Richard I. Johnson Department of Mollusks Museum of Comparative Zoology- Harvard Universit)' Cambridge, MA 02138 Dr. Douglas S. Jones Florida Museum of Natural Histor\' University of Florida Gainesville, FL .32611-20.35 Dr. James H. McLean Department of Malacology Los Angeles Count)' Museum of Natural Histon' 900 Exposition Boulevard Los Angeles, CA 90007 Dr Arthur S. Merrill % Department of Mollusks Museum of Comparative Zoolog) Harvard University- Cambridge, MA 02138 Dr. Paula M. Mikkelsen Department of Living Invertebrates The American Museum of Natural History New York, NY 10024 Dr. Gustav Paulay Marine Laboratory Universit)' of Guam Mangilao, Guam 96923 Mr. Richard E. Petit RO. Box 30 North MyrtJe Beach, SC 29582 Dr. Edward J. Petuch Department of Geology- Florida Atlantic Universih- Boca Raton, FL 33431 Dr. Gar)- Rosenberg Department of Mollusks The Academy of Natural Sciences 1900 Benjamin Franklin Parkway Philadelphia, PA 19103 Dr Ruth D. Turner Department of Mollusks Museum of Comparative Zoology Harvard University Cambridge, MA 02138 Dr. Geerat J. Vermeij Department of Geolog)' Univ-ersitv' of California at Davis Davis, CA 95616 Dr. G. Thomas Watters Aquatic Ecologv' Laborator)' 1314 Kinnear Road Columbus, OH 4.3212-1194 Dr. John B. Wise Houston Museum of Natural Science Houston, TX 770.30-1799 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 $15.00 for air mail. All orders should be accompanied bv payment and sent to: THE NAUTILUS,' PO. Box 1580, Sanibel, FL 33957, 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 The Bailey- Matthews Shell Museum, 3075 Sanibel-Captiva R(i:h1 Sanibel. FL 33975. Periodicals postage paid at Sanibel, FL, and additional mailing offices. POSTMASTER: Send address changes to: THE NAUTILUS RO. Box 1580 Sanibel, FL 33957 TH EfyNAUTI LUS CONTENTS Vohimi' 112, Number 3 October 15, 1998 ISSN 0028-1344 Noah Feinstein Stephen D. Cairns Learning from the collector: a .sur\e\' of azooxanthellate corals affixed hx Xcnopluva (Gastropoda: Xenophoridae), with an anaK'sis and discussion of attachment patterns 73 Shell microstructure of mvtilids ( Bivalvia) from deep-sea hvdrothermal vent and cold-water sulfide/methane seep environments 84 Molluscan taxa and bibliography of Henry van der Schalie 90 A new Favdiiia (Miurxiclla) from the Panamic Province (Gastropoda: Muricidae) and designation of a lectotvpe for F. (M.) cxiffLia (Broderip, 1833) . ." ' 95 Michael J. Kennish Antonieto S. Tan Richard A. Lutz Mark E. Gordon Carole M. Hertz Barbara W. Myers THE NAUTILUS 112(3):73-83, 1998 Page 73 Learning from the Collector: A Sui-vey of Azooxanthellate Corals Affixed by Xenopliora (Gastropoda: Xenophoridae), with an Analysis and Discussion of Attachment Patterns Noah Feinstein Hanard Lliiiveisih' Cambridge, MA 02138, USA and National Musenni of Natural Historv Research Training Program Smithsonian Institution Washington, DC 20560 USA Stephen D. Cairns' Department of Invertebrate Zoolog\' National Museum ot Natural Historv Smith.sonian Institution Washington, DC 20560 USA caims.stephen@nmnh.si.edu ABSTRACT AU species of die gastropod familv Xenophoridae affix foreign ob- jects to the upper shell surface. Affixed objects ma\' include bi- valve shells, smaller gastropod shells, sheO fragments, and conJ skeletons, as well as a wide array of inorganic materiid. In deep water, Xcnophom im\ ser\'e as useful prox\' collectors ot bentluc organisms. Since 1S42, coral skeletons have been noted among the attached objects, but this association has never been studied in detail. Tliis paper survevs 227 Xenophora shells, comprising 8 species from 69 stations, for affixed azooxandieUate corals. Fi\e hundred and eightv-one coralla were foimd, representing 74 coral species, 2 of wliich remain undescribed. Twenh-tour of the affixed coralla were alive at tlie time die Xcnophom hosts were collected; Xenophora not onlv collect live conils, but corals can remain iJive long after being iiffixed. Corals were found at 6 sites where the\' had pre\iousK' been unknown, iind the geographic riuiges of 29 species of coral were exjiiuided as a result of specimens found on Xenophora This paper luis hvo sections. In the first. quantitati\e obsena- tions were made on die orientation of affixed conils; statistical analvsis of these obsenations revealed non-random patterns of attachment, based on orientation of bodi the long axis of die coral iuid die coral cahce. Qiuditative obsenations suggest diat species of Xenophora favor corals ot paiticnku- shapes. In tlie second sec- tion, die speculations ot previous authors regiii-ding the ecological basis for attachment behavior are summarized and new dieories are discussed. Four of diese explanations suggest defensive ad- aptations, and die remaining 3 are functional support adaptations. Three of these hvpodieses (annor, tactile camouflage, and snow- shoeing) are proposed tor the first time in tlus paper Key iLords: Carrier shells, Mollusca, gastropods, aliermat\pic corals. INTRODUCTION According to Ponder (1983), the nionotxpic gastropod familv Xenophoridae contains 25 Recent species of AV/i- ' Corresponding author. ophorei. marine gastropods that inhabit the continental shell and slope regions of tropical and temperate oceans. Xenophora have dra\Mi the attention of naturalists and svstematists since the earK' 1800s. This was caused pri- mariK' because of a peculiar behavioral pattern: all spe- cies affix objects to the upper surface of the shell throughout some or all of its growth (see Shank, 1969 for a detailed description of the affixing procedure in X. conchi/Iiophora (Bom, 1780)). Although objects are onlv affixed at the growing edge of the whorl, older attach- ments remain on the perimeter of earlier whorls as the shell increases in size. Older attachments often become secondariK' affixed to the \oimger whorl that is ft)rmed beneath them. One of the traits used to distinguish spe- cies of Xenophora is the degree to which the shell sur- face is obscured by attachments (Ponder, 1983). A great diversih' of material has been found affixed to Xenophora: among the objects we obserxed in the course of this studv were coral skeletons, bivalve and other niollusk shells, brachiopod shells, echinoderm spines and skeletal fragments, brvozoans, sponges, sharks' teeth, and a wide assortment of inorganic frag- ments. Although corals tvpicalK' comprise less than 10% of the affixed objects, their presence on Xenophora shells was first recorded a centurv and a half ago (Reeve, 1842). Pourtales (1871) was the first to report a deep- sea (azooxanthellate) coral, Can/oplii/Uia cornufonnis Pourtales, affixed to a Xenophora shell, collected from the Straits of Florida at 433—454 m. In this case, the coral remained alive after attachment. Morton (1958) Listed living, solitarv flabellid corals affixed to A', corru- gata (Reeve, 1843) from New Zealand, and Kawase (1996) identified 4 species of azooxanthellate corals on X. paUidula (Reeve, 1842) from Japanese waters. Cairns (in press) reported 19 azooxanthellate species affi.xed to xenopliorid shells collected from the slope region of Page 74 THE NAUTILUS. Vol. 112. No. 3 Vanuatu ( = New Hebrides) (Figure 6), prompting the research presented here. In summary, there have been sever;il pubUshed observations of deep-water corals af- fixed to Xcnopliora, but no comprehensive review of the association. Xeiiophora are not highly mobile (Berg, 1975). It is therefore possible for one to learn something about the benthic fauna of a particular region b\' examining the local Xcnophi)ra. Although some species are found only in shiillow water, most Xcnophora inhabit regions well below the penetration depth of photosyntheticalK' active radiation, and specimens have been dredged from depths exceeding lOOO m (Ponder, 1983). Because rel- ativelv little is known about the deep-water bentlios. deep-dwelling Xcnophora are potentialK' useful as proxT collectors. In the process of examining Xcnophora shells for af- fixed conJs, we noticed that there appeared to be some regularity in orientation of affixed objects with respect to the shell. These apparent trends, in the conte.xt of the pre\iousK documented observation that Xcnophora at- tach lamellibranch valves with the concave side up (Pon- der, 1983; Morton, 1958; Linsle)- and Yochelson, 1973; Shank, 1969). led us to look for statistical patterns in the manner and orientation of coral attachment. The results of that analysis are presented in this report. MATERIAL AND METHODS Two hundred and twentv-seven coral-bearing A'riio/j/iora shells were examined: 145 of these are from the collec- tions of the National Museum of Natural History (USNM), Washington, DC; 42 from the Museum na- tional d'Histoire naturelle (MNHN), Paris; 38 from the Delaware Museum of Natural Histon, (DMNH), Wil- mington; and 2 from the Museum of Comparative Zo- ology' (MCZ), Cambridge. Appendix 1 lists the stations at which coral-bearing Xcnophora were collected, the station data, and the coral species collected at those sta- tions. The study material includes shells collected throughout the Indo-West Pacific, Hawaiian Islands, and the Gulf of Guinea. Eight coral-bearing species of Xcn- ophora were examined, a large majorih of the speci- mens (166/227, including all shells borrowed from the MNHN and DMNH) belonging to the species X. palli- dula. The other 7 species, in order of abundance were: X. Japonica Kuroda and Habe, 1971, X pcroniana kon- cloi Ponder. 1983, X cornifSfita, X. ccrca (Ree\e, 1845), X. ncozcahmka Snter, X. cti.spa (KiJuig, 1831), and X. granulosa Ponder, 1983. The 145 USNM specimens in- cluded all coral-bearing shells in the USNM collections, with the exception of those belonging to X. conchi/lio- phora, which is tyjiicallv found in shallow water (Ponder. 1983), and certain lots of X. pcroniana and X. japonica. which preliminarv' examination showed to be redimdant in terms of sites sampled and corals collected. When possible, corals were identified to tiic species level; when onlv a coral fragment or badlv eroded or damaged coralluiu was present, it was identified to the Figure 1. Diagram of the apical view ot a Xcnophora shell, illustrating the various tvpes of orientatious in which a solitary coral may be affixed: radial (up, down), lateral (out, in), and svinmetrical (facing up, facing down). Note: Number of apical whorls underrepresented in figure. lowest possible taxonomic level (tvpically genus). Un- described species were designated bv a letter. If a well- presened but unidentifiable phenotvpe appeared re- peatedly, it was given the designation n. sp. Anv corallum that contained remnants of dried coral tissue or was e.x- ceptionallv well preserved was considered to have been alive when the Xcnopliora was collected. Everv coral found was classified as a primarv or sec- ondary attachment. Primary attachments were those that were embedded in the shell and had clearlv been affixed bv the Xcnophora: secondarv attachments included those conds growing on a substrate which was snbse- (juentlv affi.xed to the Xcnophora and those that settled upon the surface through no action of the Xcnophora. The length of the long a.xis (maxinuun dimension, re- gardless of the orientation of nioqihological features) of each coral skeleton was recorded. The orientation of the long axis was classified as radial, meaning roughly per- pendicular to the arc of the whorl: lateral, meaning roughlv parallel to the arc of the whorl: or svmmetrical, meaning that all axes of the coral v\ere eclluin sp. (1), TnincatoflahcUnm sp. (1). Marinduque, Philippines, depth not knowii, on X, pallidula, DMNH 189002 and 188943: A. fni\tuin (1), C spiimcarens (4), H. sulcatus (1), F deludcns (1), Tnincatoflabelhnn sp. (1), B coniu (2). Batangas, Philippines, 152-163 m, on X palUdula. DMNH 189854: T. formtmim (3). Batangas, Philippines, depth unknown, on .V. pnllidula. DMNH 190007: C. spinacnrens (1), H stdcatus (1). Philippines, depth unknown, on A', pallidula. DMNH 188572: C. spinacarens (7), C. transvcrsali.'i (1). Philippines, depth unknowni, on .V. pallidida. DMNH 188944: C. spinacarens (1), T. pusillum (1), B cornu (1). B iinpcrialis (1). Quezon, Philippines, 596 m, on A', pallidida. DMNH 189043: Tnincatoflahelluni sp. (1). OffZnluland. 293 ni, on A. pallidnla. DMNH 11505-7, 11499, 17902: Madmcis sp. (1). Canjophi/llia sp. (2), FMullum sp. (I ), E paioninum (5), Tnincatoflahcllum sp. (6), T sflrdincii (6), dendroplnlliid (1). Off Zululand, 220-293 ni, on A pallidula. DMNH 20120: F paioninum (4), F sp. (1), T sp. (1), 7" cf. fonnosum (1), I g,ardineri (10). Off Natal, South Africa, 293-329 in, on X. pallidula. DMNH 186438: Flabcllum sp. (1), F pauon/UHHi (1), I gardincri (2). off Japan, 110 in, on X. pallidula, DMNH 6180: C quadragcnaria (1), F dentiformis (2), F paioninum (1), Endopachi/s graiji (1). Sagaini Bay, Japan, depth unknown, on X. pallidula. DMNH 55071: F palifenis (1), H. sulcatus (1), Tmncatoftahcllnm sp. (1). off Kii, Japan, 183 m, on X. pallidula. DMNH 55070: £. grai// (1), P dentiformis (2). MCZ Auckland, New Zealand, depth unknown), on .V, neozcnlandica. MCZ 160266-67: M nibnni} (4). USNM Albatross 3810 (.soudieast of OiJui, Hawaiian Islands, 386—463 in), on A, pcroniana. USNM M749844: A, pacifica (1), £. grai/' (!)• Albatross 3838 (southeast coast of Molokai, Hawaiian Islands, 168-.388 in), on X. peroniana, USNM M335067: E grat/i (5). Albatross 4079 (Puniawa Point, Maui, Hawaiian Islands. 262-326 in), on X. peroniana. USNM M.3.35063: A macrolobata (3). Albatro.-is 5117 (13°52'22"N. 120°46'22"E. 216 in), on A. pallidula. USNM M243416: C, spinigcr (1), Flabcllum sp. (1). Albatross 5131 (off Panabutan Point, Philippines. 49 in), on .\'. pallidula. USNM M243391: C.'secta (1), H. .mlcatus (1). Albatross 5212 (12°04'15"N, 124°04'36"E, 198 m), on X pallidula. USN.M M277576: F lamellulosum (2), T cf gardincri (1), B. coniu (3), B. imperialis (1). Albatross 5213 (12°15'00"N, 123°57'.30"E, 146 ni), on A. pallidula. USNM M243398: Tmncatoflabcllum sp. (1). Albatross 5265 (13°4I'15"N, 120°00'50"E, 247 m). on X. pallidula, USNM M243347-9, -353^ -356-357, -.361-2. -.366, -.368, -370, -372, -373, -375, -376, -426, -881: Anthcmiphijllia sp. (I), C. spinnrflredA (1), C. .swta (2). C. graiji (1). T. t/rgnffw (1), H. sulcatus (1), D. philippinensis (1), C. /j(7e(j.s (I), Flabcllum sp. (3), F pavoninum (I), F lamcllulo.wm (I), T. fonnosum (12), T f/ms (1), I pusillum (1), I sp. (1), B. co;7i!/ (I), B. imperialis (2). £ grai/i (D. W roWi/ra (1). Albatross 5273 (13°58'45"N. 120°21'35"E, 209 in), on X. pallidula. USNM M237519: Balanophi/llia sp. (I). Albatross 527H (I4°00'1()"N, 12(n7'1.5"E. 187 in), on X pallidula. USNM M 24,3411: C. octonaria (1), H. .si//rafi/.v (3). H. altcrnatus (1), A. nibesccns (1), Tropidocijathus sp. (1). Albatross 5289 (13°41'.50"N, 120°58'30"E, 315 in), on .\. pallidula. USNM M237585: 7" fonnosum (1). I (;irn/.sto?i/»i (1). Albatross 5312 (21°30'00"N. 116°32'00"E. 256 in), on A. japonica. USNM M243420: I cf gardincri (1). Albatross 5391 (12°13'15"N, 124°05'03"E, 216 in), on X. japonica. USNM M238138: H. .siJraffl (1), F po/ifi/»! (14). T candcanum (1), Balanophijllia sp. (2). Albatross 5392 (i2°13'35"N, 124°02'48"E, 247 in), on A pallidula. USNM M238192, 243394: F paioninum (2). F /A)/(f!/»i (7), F sp. (10). B. ronii/ (4), Albatross 5392 (see above) on X. japonica. USNM M238192: F politum (50). F paioninum (1), F sp. (13), B. foni!/ (6), B. sp. (5). Albatross 5394 (12°00'.30"N. 124°05'36"E. 280 m), on A pallidula. USNM M24.34I5: T candcanum (1). Albatross .5395 (ir56'40"N. 124°14'E. 2.56 in), on .Y, pallidula. USNM M240513: B wr/ii; (1). Albatross 5405 (10°49'20"N, 124°24'23"E. 479 m). on X pallidula. USNM M238257: Co/iofroc/u/.y sp. (2). Albatross 5408 (10°40'I5"N. 124°15'00"E. 291 ni). on A. pallidula. USNM M238277: Fimgiacijathus sp. (I). Madrepora oculata (2), H. .s(//cn?i/.s (4). Albatross 5416 (I0°11'30"N, 12,3°53'30"E. 274 in), on A pallidula. USNM M238372: A. fnistum (2). //, .si/Zcrtfi/.s (5). Flabcllum sp. (3). T angustum (1). Albatross .5417 (10°10'00"N, 12.3°53'I5"E. 302 in), on .V pallidula. USNM M24.3381: Deltocijathus sp. (I). Flabcllum sp. (2). F paioninum (2). Albatross 5418 (I0°08'5t)"N. 12:3°52':30"E, 291 in), on X pallidula. USNM M243377: .\/. ori/Z^ffl (1), D rt(u/«/(m/i/c».v (1), Flabcllum sp. (I), r angustum (1). Albatross 5592 (4°12'44"N, 118°27'44"E. 558 in), on A. pallidula. USNM .M229317: H. altcrnatus (1). Albatross 5661 (5°49'40"S, 120°24'30"E, 329 in), on A. pallidula. USNM M239497: Canjophi/llia sp. (1). Dauirf Starr Jordan, stn TC32-2 (21°21'54"N. 158°I2'24'\V. 119-291 m). on A, pcroniana. USN.M M8076,59: Anthcmiphi/llia sp. (1), £. grflyi (2). Placotrochides n. sp. (1). Dnr/f/ Starr Jordan, stn TC3:3-9 (2r00'06"N. I56°45'42"W. 227-234 in), on .V peroniana. USNM M807662: A, pflf(/ir« (1), Flabcllum sp. (2). F paioninum (1), Placotrochides n. sp. (3). Page 82 THE NAUTILUS, Vol. 112, No. 3 David Starr Jordan, stn TC40-8 (21°09'42"N, 157°24'42"W, 183 ni), on X. penmiana. USNM MS076.31: BalnnH( (llll'.HM M.-VeNT MvTII.IDS The microstructures encountered within the shells of the hvdrothermal-vent mytilids examined in this stud\' ;u-e hpical of calcified structures pre\ionsI\ described in oth- er members of the tainiK Mstilidae, including the vent mussel Baflii/iuodiolus thcniiophilus. From the vacuo- lated periostracum inwards, the calcified lavers of B. tliermopliilus consist of (I) fibrous prismatic calcite; (2) nacre (aragonite): (3) irregular prismatic aragonite (pal- lial mvostracuin); and (4) nacre (aragt)nite) (Lutz and Hhoads, 1980). A relatively thick organic periostracum appears to effectiveK' prevent substantial dissolution of M. J. Kennish et al., 1998 Page 87 Figures 13-21. Microstmctures of deep-sea mytilids. 13-15. Species A. ANSP 400787. 13. Angular view of periostracum (P) (ventral margin), coarse grained homogeneous (CGH) (comarginal fracture), and fibrous prismatic (FP) stnictures. Autero-posterior shell axis from top to bottom of micrograph. Horizontal field width (HF\V) = 240 (xm. Preparation 1. 14. E.xterior \ie\v of periostracum infested with branching filamentous organisms. Same orientation as Figure 1. HFVV = .320 jim. Preparation 1. 15. Exterior wew of an oblique fracture of the coarse grained homogeneous (CGH) and fibrous prismatic (FP) structiu'es. Antero- posterior shell axis from top left to bottom right of micrograph. X'entral shell margin to right. HFVV = 40.5 p-m. Preparation 1, 16- 17. Species B, ANSP 400788, 16. Coarse grained homogeneous (CGH) and fibrous prismatic (FP) structures. Double fracture: (1) radial fracture towards bottom left of micrograph; (2) horizontal (tangential) fracture towards top right. Autero-posterior shell axis perpendicular to plane of micrograph. HF\V = 70 \x.m. Preparation 1. 17. Transverse fracture of predominantlv lath-type fibrous prismatic structure. Outer shell surface towards bottom of micrograph. Antero-posterior sliell axis from left to right of micrograph. HFW = 35 ^x.m. Preparation I. 18. Species C, ANSP 400789, transverse fracture of predominantlv rod-hpe fibrous prismaHe structure. Outer shell surface towards bottom of micrograph. ."Vntero-posterior shell axis from left to right of micrograph. HFW = 35 \Lm. Preparation 1. 19. Species D, ANSP 400790. Radial fracture showing conical arrangement of rotl-txpe fibrous prismatic structure. Outer shell surface towards top of micrograph. Outer shell surface towards bottom of micrograph. Antero-posterior shell axis fVom left to right of micrograph. HFVV = .305 (jlui. Preparation 2. 20-21. Species E, ANSP 400791. 20. Radial fracture showing crossed lamellar arrangement of lath-txpe fibrous prismatic structure. Antero-posterior shell axis from bottom left to top right of micrograph. HFW = 70 \xm. Preparation 1 (Same orientation as Figure 7). 21. Radial fracture showing complex crossed lamellar arrangement of lath-t>pe fibrous prismatic structure. Antero-posterior shell axis from bottom left to top right of micrograph. HFW = 170 |a.m. Preparation 1 (Same orientation as Figure 7), the relativei\' thin and fragile shell of B. thcrmopliilus throughout its life (Lutz, 1982). The calcitic fihroiis prismatic laver, a specialized layer observed in the mxlilids, is characterized by prisms with high length/width ratios. As reported by Carter (1990), first-order prisms in fibrous prismatic structures have a non-spherulitic prismatic and non-composite prismatic substnicture, but unlike simple prisms, appear as long fibers. Several investigators have classified fibrous pris- matic structure as an independent entitx' because of its unique stmcture (MacCIintock, 1967; Carter, 1980a, b; Carter and Clark, 1985; Watabe, 1988; Carter, 1990). Nacreous structures in h\'drothermal-vent niNtilids are composed of tabular aragonitic crxstals deposited on an organic matrix. The calcareous tablets, identical to tlie sheet nacre described by Carter and Clark (1985) and Carter (1990), are arranged in broadh' continuous, reg- ular, and mutualK parallel sheets. The\ exhibit similar Page 88 THE NAUTILUS, Vol. 112. No. 3 -^f/^U lS^^^^ i'-.'' ^^ ^^ ' .M-i^^A Figure 22—30. Microstmctiires of deep-.sea nntilitl.s. 22. Specie.s C, ANSP 4007S9. Radiiil Iractuie ot tlie boiiiRlan- between fibrous prismatic (FP) and nacreous (N) structures. Dorsal shell surface towiirds bottom of micrograph, .'^ntero-posterior shell axis from bottom left to top right of micrograph. HFW = 3.5 |xm. Preparation 1. 23. Species A, ANSP 400787. Transverse fracture of the blocky prismatic (BP) sandwiched between the fibrous prismatic (FP) and nacreous (N) structures. Dorsal shell surface towards top of micrograph. Antero-posterior shell axis from left to right of micrograph. HFW = .35 \i.m. Preparation 1. 24. Species B, ANSP 400788. Inner surface view of the nacreous stnictnre, near the boundan,' between fibrovis prismatic and nacreous stnictures. \'entnil shell surface towards left of micrograph. Antero-posterior shell axis Irom top to bottom ot micrograph. HF^\' = 70 jo-m. Preparation 1. 25- 27. Species D, ANSP 400790. 25. Radial section of columnar nacre (CN) underKing the fibrous prismatic (FP) structure, \entral shell surface towards bottom ot micrograph. Antero-posterior shell axis trom left to right of micrograph. HFW = 45 jjim. Preparation 2. 26. Inner surface view oi hexagouiil tablets of sheet nacre. Shell margin towards bottom left ot micrograph. HF\\' = 70 jjim. Preparation 1. 27. Inner surface view ot oval tablets ot sheet nacre. Shell margin towards bottom left of micrograph. HFW = 70 \x.m. Preparation 1. 28. Species D, ANSP 400790. Inner surface view of rhomboidal tablets of stack nacre. Shell margin towards bottom left of micrograph. HFW = 70 \i.m. Preparation I. 29. Species C, ANSP 400789. Radial fracture of adductor nivo.stracum (AM) consisting of simple prismatic stmctme underneath the nacreous laver. Inner shell surface towards bottom left of micrograph. HFW = 70 \i.n\. Preparation 1. 30. Species B, ANSP 400788. Angular view of pallial mvostracum (PM) consisting of simple prismatic stnicture underneath the nacreous layer Inner shell surface towards bottom right ot micrograph. HFW = 70 ^.ni. Preparation 1. complexitv and form as those comprising the inner or middli' shell laver of other nn+ilid species (Carter, 1990). Irregular simple prismatic structure occurs under the areas of muscle attachment: the palliiil and adductor myostraca. The prism cross sections are highly variable along their lengths (Carter, 198()a; 1990). This irregular- ity of the prism cross sections results in a lens-like or wedge-hke prism shape (Carter and Clark. 1985). DISCUSSION SEM examination of fractured and sectioned shells of deep-.sea mussels targeted in this study reveals similar microstructures in distinct arrangements. The shells of mytilitis inhabiting both cold-water sulfiile/methane seep and deep-sea hvdrothermal vent environments are con- sistently composed of outer fibrous prismatic and inner nacreous stnictures. which underlie a periostracal laver. Simple prismatic structure forming the adductor and pallial mvostraca are also present. The microstruitures in these specimens exhibit similar complexity to those of the previously described vent mussel ButhijiiuHliolus tlwriitopliihis. Based on shell microstnicture, Ta\lor et id. (1969) identified 2 groups of species belonging to the Mvtilacea: M. T. Kennish et al.. 1998 Page 89 (1) a t\vo-la\'ered, whoIK- nacreous, \\'hol]\' aragonitic warm-water group (e.g., Pcnw liiidis (Linnaeus, 1758), Choroimjtihis palliopunctatus (Carpenter, 1857), and Pcniinytihis puijniratiis (Larmarck, 1819): and (2) a two- or three-la\ered aragonitic and c;ilcitic, temperate group (e.g., Mijtihts cduUs (Linnaeus, 1758) and Mytilui caUforniamts (Conrad, 1837)). In the latter group, the prismatic lavers consist of calcite, whereas the nacreous layers are composed of aragonite. Bet^\een the outer ciil- citic prismatic and tlie inner nacreous la\ers, a middle aragonitic prismatic \a\ex is present in some mvtilid shells (Blackwell et al.,' 1977; Carter, 1980a). Lutz and Rhoads (1980) and Lutz (1982) recognized 4 distinct las- ers in the vent mussel Bothi/mocliohis thcnnophilus. in- cluding (1) fibrous prismatic calcite, (2) nacre (arago- nite). (3) irregular prismatic aragonite (pallial niNostrac- um), and (4) nacre (aragonite). The shell microstructures in Bathyinocliolii.s tlienno- phihis are most similar to those in species of Mytihis. Idasola, and Modiolus (Carter, 1990). According to Car- ter (1990), the outer shell la\er of B. thcntiophihis is calcitic with outer homogeneous and inner, predomi- nantly fibrous prismatic sublayers. The middle and inner shell layers are largeh' comprised of nacre, with minor irregular simple prismatic structure. Hence, the findings of Carter (1990) corroborate those of Lutz and Rhoads (1980) and Lutz (1982). Carter (1990) noted that the outer shell layer of m\- tilids in general shov\s considerable mineralogical and microstructural yariabilitv. In studies of m\tilids from the Carboniferous to Recent, some species exliibit a cal- citic fibrous outer shell la\er, whereas other species ha\e various, different calcitic and/or aragonitic prismatic, ho- mogeneous, and even minor crossed structures in this layer. Waller (1978) generalized that a calcitic outer shell layer in the M\tiloida, where present, consists of regular fibrous prismatic structure. Carter (1990), however, identified various combinations of aragonitic and/or cal- citic irregular simple prismatic, irregular spherulitic pris- matic, irregular fibrous prismatic, regular fibrous pris- matic, and homogeneous structures in the outer shell lavers of this order. The shell microstructures of the 6 undescribed mxtilid species investigated in this stud\' contain 4 calcified shell layers composed of aragonite and calcite, although dif- ferences in the layers can be significant owing to various combinations of aragonitic and/or calcitic structures present within the shell la\ers. The arrangement of shell layers can also vary appreciably from one part of a mv- tilid shell to another, because of alternating adductor myostraca, dissolution, or other effects. Thus, in the as- sessment of shell microstructure patterns, careful ex- amination of the entire shell is necessary, as conducted in this stutK'. These differences in shell microstructure appear to be of great potential \alue in s\stematic stud- ies of nn+ifids from deep-sea h\drothermal-vent and cold-water sulfide/methane seep environments. .ACKNOWLEDGMENTS This is Contribution No 98-05 of the Institute of Marine and Coastal Sciences, Rutgers Universit\', supported by New Jersey state funds, the Deep-Sea Ecology- and Bio- technologv' Center, and National Science Foundation grants OCE 89-17311, OCE 92-17026, and OCE 96- 02205. We thank Richard Custafson for collecting some of the mxtilid specimens and Alan Poole)' for help with the illustrations. LITERATURE CITED BiackwvU. ]. R, L. F. Gainey. Jr. imd M. J. Greenberg. 1977. Shell uitm.stnictiire iii h\o subspecies of tlie ribbed mussel, Geu- kcu.sia tlciuissn (DilKwi, 1817)), Biologiciil Bulletin 1.52:1- 10. Caiter. J. G. 19S0a. Guide to bi\al\e shell microstnictures. In: Rlioads, D. C. and R. A. Lutz (eds.) Skeletal Groiiih of Aquatic Organisms. Plenum Press, New York, pp. 64.5-673. Garter, J. G. 1980b. Selected mineralogical data for tlie Bi\a]\ia. 7/1 RlioatLs. D. C. imd R. A. Lutz'(Eds.) Skeletal Gnmth of Acjiiatic Organisms- Plenum Press, New York, pp. 627-643. Garter, J. G. 1990. Glossaiv of skeletal biominer;ilizatiou. In: Gar- ter J. G. (ed.) Skeldal Biomineralization Patterns. Processes and Eiolutionan/ Trends. Van Nostnuid Remhold, New York, pp. 609-671. Garter, J. G. and G. R. Glark II. 1985. Glassification and ph\Io- genetic significance of moUusciUi shell ultnistnictiire. In: Bottjer, D. J.. C. S. Hickman and R D. W'luxl (eds.) Mollusks: Notes for a Short Course. Studies in Geology 13, Depajt- ment of Geological Sciences, Universitv of Tennessee. Knox- \iUe, pp. 50-71. Kenk, V. C. and B. R, \\'ils()n. 1985. A new mussel (Bi\iJ\ia, M\tilidae) from Indrothenna] \ents in die Galapagos Rift zone. Malacologia 26:25.3-271. Kemrish, M. J.. A. S. Tan, and R. A. Lutz. 1996. Shell microstnic- tnre of \'esicom\id clams from \arious h\drodieniiiJ \ent luid cold seep emironments. Malacolo0a 37:.36.3-^373. Kennish, M. J., A. S. Tim, and R. A. Lutz. 1998. Deep-sea vesi- comvid clams from h\dr(3tliemi;J \ent luid cold seep emi- ronments: analvsis of shell microstnictiire. The \eliger 41(2): 19.5-200, Lutz. R. A. 1982. Shell microstructure and mineralogy of two species of bivalves from deep-sea Inclrotheniial vents. Anier- iciui MiilacologicJ Bulletin 1:101. Lutz, R. A. iuid D. C., RlioacLs. 1980. Shell structure, mineralogy, iuid niicromoq^hologv of deep-sea dieniial vent bi\;ilves from tlie Galapagos Vtik: ecologiciil implications. Proceedings of die National Shellfislieries Association 70:127, MacClintock, C. 1967, Shell stnicture of patelloid mk\ bellero- phontoid gastropods (Mollusca). Bulletin of die Peabcxfv Museum of Natural Hi.story 22:1-140. Tavlor, J. D., W. J. Kennedy and' A. H;ill. 1969. The shell structure and mineralogv" of die Bivalvia: Introduction. Nuculacea — Tiigonacea. Bulletin of the British Museum (Natural Histo- iv). Zoology Supplement .3:80—85, Waller, T. R. 1978. Moipliologs; moiphocUnes, ;uid a new classi- fication of the Ptenomoipllia I. Mollusca: Bivalvia). PMosopli- ical Transactions of die Roval Sixieh' London Series B 284: .•34.5-.365. Watabe, N. 1988. Shell stnicUire. In: Trueman. E. R. lUid M. R. Glarke (Eds.) The Mollusca. Volume 11. Form and Function. Academic Press, New York, pp. 69-104. THE NAUTILUS 112(3):9()-94, 1998 Page 90 Molluscan Taxa and Bibliography of Henry van der Schalie Mark E. Gordon Zoolog\' Section UniversiU- ol Colorado Museum Campus Box 315, Hvuiter Building Boulder, CO 80.309 USA and New Mexico Museum of Natural Histor\' and Science 1801 Mountain Road NW Albuquerque, NM 87104 USA Henrs van der Schalie was bom in Amsterdam, The Netherlands, on Jamiarv 8, 1907. His family immigrated to the United States in 1909. following the death of his father, and settled in Paterson, New Jersey. Van left New Jersey in 1925 to enroll in Calvin College, Grand Rapids, Michigan. During his undergraduate studies, he devel- oped an interest in parasitology-. After receiving his A.B. in 1929, Van began graduate studies in parasitologN' at the University of Michigan (UM) under George R. LaRue, Chair of the Department of Zoologx'. But before long he accepted an assistantship in and transferred to the Mollusk Division of the Museum of Zoologs at UM. He received an M.S. in 19.31 and Ph.D. in 1934. Thus began a career in malacolog>' at the Museum of Zoolog)- that extended from his initial appointment as a museum assistant in 1929 to his retirement as curator of mollusks and professor of zoolog\' in 1977. A review of activities in the Division of Mollusks during this period is pre- sented in van der Schalie (1981). Van became assistant curator of mollusks in 1934 and was promoted to curator of mollusks in 1944 fcjllowing the retirement of Calvin Goodrich. Initially appointed as an instructor of zoology (1937-1944), he later held as- sistant (1944-19.50). associate (1950-1956), and full pro- fessorships (1957-1977) in the Department of Zoology, UM, and served in visiting or adjunct capacities at other institutions. With the onset of World War II, his research returned to parasitologv. He studied mollusks as vectors for various diseases, particularlv schistosomiasis. Subse- (juently, he ser\ed on numerous national and interna- tional committees and as a consultant to various inter- naticmal organizations involved in medical malacology. Van was widely known as a researcher, tnlucator, and advocate in malacology, parasitology, environmental con- servation, and general education. He received many re- search grants and was member of numerous professional societies. In recognition for his earlv contributions to malacology, the fossil alasniidontiue (Hivalvia: Uiiiono- idea) taxon, Vonder.schalica Modell (1943: 112; tvpe-spe- cies: Unio kolasii Modell, 1931), was named after him. Although retiring as curator emeritus from the Univer- sitv' of .Michigan in 1977, he did not cease his profes- sional interests and continued his research and advcxacy until his death on 15 April, 1986. In adthtion to data in van der Schalie (1981), bio- graphic information has appeared in Kraemer and Berrv' (1986) and Abbott (1987); however, none of these ref- erences listed or reviewed his publications or the taxa that he described. Van der Schalie published 142 manu- scripts during the period of 1932-1986. Several of these works were published initially as a summary or an ab- stract, with the complete version of the paper pubhshed later in a different journal, A few papers were reprinted in other journals; in the following bibliography, the sub- secjuent printing is noted in parentheses after the orig- in;il reference. Between 19.34 and 1939, Van co-authored 3 new species of prosobranch (Hydrobiidae) and 4 pul- monate (1 Ancylidae, 2 Oleacinidae. 1 Sagdidae) gastro- pods from Guatemala with his mentor Calvin Goodrich, and described 3 new species of unionoid bivalv es (2 from the United States, 1 from Canada). MOLLUSCAN TA>L\ Abbreviations: CM = Carnegie Museum. Pittsburg, Pennsylvania; UF = Florida Mu.seum of Natural Histon', University of Florida, Gainesville; UMMZ = Museum of Zoology, University of Michigan, Ann Arbor (i^iiailtic. Fcnis.sin (Laevapex) Goodrich and van der Schalie, 1937 (1.5 March), Miscellaneous Publications, Museum of Zoologv', Universitv- of Michigan 34:23, pi. 1, fig. 7-7a ; type localitv': Aguada de Copo, one mile south of La Lib- ertad, Petc^n, Guatemala. Holotxpe: UM.MZ 6.5583. The generic allocation of this species is luicertain. aharadoi. Spiraxis Goodrich and van der Schalie, 19.37 (15 March). Miscellaneous Publications, .Museum of Zoologv, Uui\ersitv of Michigan 34:23, pi. 1, fig. 1: hpe localitv: limestone knoll five miles north of El Pa.so de los Caballos, Peten, Guatemala. Holohpe: UMMZ 65165. brouksiana, Anodonta van der Schalie, 1938 (4 June), Annals of the Carnegie Museum 27:167, pi. 16; tvpe locality: Spout Pond Ann, Fcrrviand District, Southern Shore, Newfoundland. Holotvpe: CM 61.13137. The taxonomic status of this .species is not clear. {\\. R. Hoeh, personal conumuiication). cicnchi. S()»w/(igi/n/.v (ioodrich and van der Schalie, 1937 (15 March), Miscellaneous Publications, Museum of Zoologv; Univer.sitV' of Michigan 34:.37, pi. 1, fig. 6: tvpe localitv: Ri'o de la Pasicin, S;iyaxclu'', Petc^n, Guatemala, Holotvpe: M. E. Gordon, 1998 Page 91 UMMZ 65371. Aroapi/n^iis ciciichi {fide Hershler and Thompson, 1992). franccsac. Cochliopii Ciootlritli and \an der Schalie, 1937 (15 March), Miscellaneon.s Pnbhcations, Museum ot Zoolog\', Universih' of Michigan 34:3S, pi. 1, fig. 3; Upe localiU': Ri'o de la Pasion, at the mouth of Arrovo Chajchim'c, northwest of Ponenir, Alta Vera Paz, Guatemala. Holo- t\pe: UMMZ 6533S. Cochliopiiui franccsac [fide Hershler and Thompson, 1992). ftinibiis. Spiraxis Goodrich and \an der Schalie, 1937 (15 March), Miscellaneous Publications, Museum of Zoolog\; Universit\' of Michigan 34:23, pi. 1, fig. 2; t\pe localit\': limestone kiioll fi\e miles north of El Paso de los Caballos, Peten, Guatemala. Holot\pe: UMMZ 65166. jonesi, LampsiUs van der Schalie, 1934 (4 Mav), The Nautilus 47(4): 125, pi. 15, fig. la-b, 2, 3a-b; type localit\-: Pea River Preston's Mill, Dale County, Alabama. Tvpe material orig- inally was deposited in the Alabama Museum of Natural Histor)': however the malacological collection of that nni- seum was given to the F"lorida Museum of Natural Histon with se\'eral lots subsequentlv transferred to the Museum of Comparatise Zoolog\-, Harvard Uni\ersit\' (KG, Thompson, personal communication). Lectotvpe: UF 6555S, designated by Johnson (1967; the specimen se- lected was the male figured in van der Schahe. 1934). Pti/chobrnnchtis jonesi (fide Atheam, 1964), mcglaineriac. Medionidiis van der Schalie, 1939 (24 June), Oc- casional Papers, Museum of Zoologw University of Mich- igan 407:1, pi. 1; hpe locality-: Tombigbee River Epes, Sniiipter Count^, Alabama. Holot\pe: UMMZ 130460. pa-iioncnsi.s. Aiiinicold Goodrich and van der Schalie, 1937 (15 March), Miscellaneous Publications, Museum of Zoologw Universitv of Michigan 34:36, pi. 1, fig. 4; tvpe localih': Arroyo Subin, tributarv- of the Ri'o de la Pasion, about two miles above Santa Teresa, Peten, Guatemala. Holotvpe: UMMZ 65357. Aroapijraus pasionensis (fide Hershler and Thompson, 1992). taintori. Xenodi.scuin Goodnch and van der Schalie, 1937 (15 March), Miscellaneous Publications, Museum of Zoolog\, University- of Michigan .34:26, pi. 1, fig. 5, 5a-b; hpe lo- cality; woodland just east of El Paso de los Caballos, Pe- ten, Guatemala. Holotvpe: UMMZ 65177. BIBLIOGRAPHY OF HENRY VAN DER SCHALIE van der Schalie, H. 1932. The station of Diisnoiniii tiicjiutra (Raf ). The Nautilus 45(3):104-105. Goodnch, C. and H. van der Schahe. 1932. I. On an increase in the naiad fauna of Saginaw Bav, Michigan; II. The naiad species of the Great Lakes. Occasional Papers, Museum of Zoology. University of Michigan 238: 1-14. van der Schalie, H. 1933. Notes of the brackish water bivalve, Pohjmesoda caroUniana (Bosc). Occasional Papers, Mu- seum of Zoologv', Universih- of Michigan 258:1-8. van der Schalie. H. 1934. Lampsilis jonesi. a new naiad from southeastern .i^labama. The Nautilus 47(4):12.5-127. van der Schalie, H. 1936a. Transposed teeth of North Amer- ican naiades. The Nautilus 49(3):79-S4. van der Schalie, H. 1936b. Ovovi\iparitv among mollusks. The Nautilus 50(l):lf-^19. van der Schalie, H. 1936c. The naiad fauna of the St. Joseph River drainage in southwestern Michigan. The American Midland Naturalist 17(2);52.3-527. van der Schalie, H 1936d. An unusual naiad fauna of a south- em Michigan lake. The American Midland Naturalist 17(3):626-628. van der Schalie, H. 1937a. A mussel taken from the stomach o{ Rana catesbiana Shaw. The Nautilus 50(3): 104-105, Clench, W.J. and H. van der Schalie. 1937. Lampsilis cariosa (Say), The Nautilus 50(3):105. van der Schalie, H. 1937b. K method lor shippmg freshwater mollusks. Basteria 2(1):4— 6. van der Schalie, H. 1937c. The relationslnp of the gravid pe- riods ot certain nuissels in Michigan to the pearl button industn-. Transactions of the American Fisheries Society' 66:406^10. Goodrich, C, and H. van der Schalie. 1937. .Vlollusca of Peten and north Alta Vera Paz, Guatemala. Miscellaneous Pub- lications, Museuui of Zoologv, Universitv of Michigan 34; 1-50. van der Schalie, H. 1938a. The naiad fauna of the Huron River, in southeasteni ,Michigan. Miscellaneous Publica- tions, .Museum of Zoologv, Universitv of Michigan 40:1- 83. van der Schalie, H. 1938b. On the occurrence of Helix lactea Miiller in North America. The Nautilus 51(4):132-1.34. van der Schalie, H. 1938c. Remarks on some of Dr. Paul Bartschs experiments. The Nautilus 51(4):134-135. van der Schalie, H, 193Sd. Review of Ovster culture and ovs- ter biology-, b\ Orton. Copeia 1938:52. van der Schalie, H. 1938e. Anodonta brooksiana. a new naiad from Newfoundland. .Annals of the Carnegie Museum 27(12): 167-170. van der Schalie, H. 1938f. Hitch-hiking mussels and pearl but- tons. Michigan Conservation 7(Jiine):-l-5. van der Schalie, H. 1938g. Contributing factors in the deple- tion of naiades in the eastern United States. Basteria 3(4): 51-57. van der Schalie, H. 1938h. The naiades (fresh-water mussels) of the Cahaba River in northern .\labama. Occasional Pa- pers, Museum of Zoologv-, Universitv of Michigan 392:1- 29. van der Schalie, H. 1939i. Hendersonia occidta (Sav) in Mich- igan: its distribution, ecology-, and geographical signifi- cance. Occasional Papers. Museum of Zoologv, Universitv of Michigan .399:1-8. van der Schalie, H. 1939j. Mcdionidus nicglantciine. a new naiad from the Tombigbee River with notes on other nai- ads of that drainage. Occasional Papers, Museum of Zo- ology, Universitv of Michigan 407:1-6. van der Schalie, H. 1939k. Distributional studies of the naia- des as related to geomoiphologv. Journal of Geonioq-)hol- o,g\- 2(3);251-257. van der Schalie, H. 19391. Additional notes on the naiades (fresh-water mussels) of the lower Tennessee River The American Midland Naturalist 22(2):452-457. van der Schalie, H. 1939in. Aquatic mollusks of the upper peninsula of Michigan, part II; the naiades (fresh-water mussels). Miscellaneous Publications, Museum ofZoologv. Universih- of Michigan 43:3.5 — 15. van der Schalie, H. 1940a. Larger land snails from pine woods in northern Michigan. Papers of the Michigan .\cadcmv of Science, Arts, and Letters 25:367-370. van der Schalie, H. 1940b. Notes of Mollusca from Alta Vista Paz, Guatemala. Occasional Papers, Museum of Zoologv-, University^ of Michigan 413:1-11. van der Schalie, H. 1940c. The naiad fauna of the Chipola River, in northwestern Florida. Llovdia 3(3);191-208. Page 92 THE NAUTILUS. \'oI. 112. No. 3 van der Schalie. H. 1940d. .^estivation of Li/innacii l/iiicctitii (Gould). The Nautilu,s 53(4):134-135. van der Schalie, H. 1940e. .\e.stivation of fresh-water mussels. The Nautilus 53(4): 137-138. van der Schalie. H. 1941a. Zoogeograph\ of naiades in the Grand and Muskegon ri\ers of Michigan as related to gla- cial histor\'. Papers ot the Michigan Academy ot Science, Arts, and Letters 26;29T-,310. van der Schalie, H. 1941b. The taxonomy of naiades inhabiting a lake environment. Journal of Concholog>'21(8):246-253. van der Schalie, H. and F. Locke. 1941. Hermaphroditism in Anodonta grandis. a fresh-water mussel. Occasional Pa- pers, Museum of Zoologv', Universih^ of Michigan 432:1- 7. van der Schalie, H. 1941c. On collecting fresh-water mussels. Annual Report of the .American Malacological Union tor 1940: 9-14. Clench, W.J. and H. van der Schalie. 1944. Notes on naiades from the Green, Salt, and Tradewater rivers in Kentucky. Papers of the Michigan Acadenu' of Science. Arts, and Letters 29:22.3-228. " Goodrich, C. and H. van der Schalie. 1944. A revision ot the Mollusca of Indiana. The American Midland Naturalist 32(2):257^326. van der Schalie, H. 194.5a. The value ot mussel distribution in tracing stream confluence. Papers of the Michigan Acad- emv of Science, .\rts. and Letters .30:.3.5.5-.373. van der Schalie, H. 1945b. What has happened to the genus lo? Mollusca 1:59-61. van der Schalie, H. 1947. The ecology' of molhisks for the biology teacher American Biology Teacher 9(6): 174-176. van der Schalie. H. 194Sa. The Michigan pearl button indus- try. Annual Reports of the .American .Malacological Union for 1947:8 (repnnted m 1963. Sterkiana 9:29). van der Schalie, H. 1948b. The land and fresh-water mollusks of Puerto Rico. Annual Report of the American Malaco- logical Union for 1947:18 (reprinted in 1963, Sterkiana 9: 29). van der Schalie, H. 1948c. The commerciallv \aluable mussels of the Grand River in Michigan. Miscellaneous Publica- tions of the Michigan Department of Consen ation Insti- tute of Fishery Research 4:1—12. van der Schalie, H. and E.G. Berry. 1948. A(|natic puliiionates from Lake Tahoe. The Nautilus 62(1):.3— 1. van der Schalie, H. 1948d. The land and fresh-water mollusks of Puerto Rico. Mi.scellaneous Publications, Musemri of Zoologv'. Universih of Michigan 70:1-1.34. van der Schalie, H. 1949a. The fresh-water mussels of the Mississippi River from St. Paul to the mouth of the Ohio River. Annual Reports of the American Malacological Union for 1948:16 (reprinted in 1963, Sterkiana 9:29). van der Schalie, H. 1949b. Biographic notes on .\rnold Ed- ward Ortmami as revealed b\ some of his letters. .Annual Reports of the American Malacological Union lor 1948: 11-12 (reprinted in 1963, Sterkiana 9:.3()). van der Schalie, H. 1950. Another techni(]ne for the prepa- ration of radulae. Ainnial Reports of the American Mala- cological Union for 1949:8. Rogick, D. aTid H. van der Schalie. 19,50. Studies on fresh- water Brvozoa. Ohio Journal of Scicnee 50:1.36-146. van der Schalie. H. and A. van der Schalie 1950. The mussels of the Mississippi River The .American Midland Naturalist 44(2):448-466. \an der Schalie. H. 1951a. Arnold Edward Ortmaim as re- vealed by his letters. The Nautilus 64(4):I.34-141. van der Schalie, H. 19.51b. .Arnold Edward Ortmann as re- vealed by his letters-II. Tlie Nautilus 65(l):23-26. van der Schalie. H. 19.52a. .An old problem in naiad nomen- clature. .Annual Reports of the .American Malacological Union for 1951:4-5 (reprinted in 1963, Sterkiana 9:.30), van der Schalie, H. 1952b. ,An old problem in naiad nomen- clature. The Nautilus 65(3);9:3-99. van der Schalie, H. 1953a. Ecology and distribution of Lijitt- nacii (Bidimnca) iiicga.soina in Michigan, .Anniuil Reports of the American Malacological Lhiion. for 1952:14 (re- printed in 1963, Sterkiana 9:31). van der Schalie. H. 19.53b. Interim report: bilharzi;isis control project at Qiil\iib, Egypt. United Nations World Health Organization EM/BILy6:l-41. van der Schalie. H. 1953c. Mollusks from an interglacial de- posit (Sangamon age?) in Mead Counts; Kansas, The Nau- tilus 66(3):80-90. van der Schalie, H. 1953d. Nembutal as a relaxing agent for mollusks. The American Midland Naturalist 50(2):511- 512. van der Schalie, H. 19.54a. Bilharziasis in Eg\pt The Biologist 36(l-4):3;^.34. \'an der Schalie, H, 19.54b. Obser\ations and report on a pro- gram for bilharziasis control in the Gezira irrigation area of the Sudan. United Nations World Health Organization EM/BIL/5:l-5. van der Schalie, H. 19.54c. Eg\pt's fight against bilharziasis. WHO Newsletter 7(1 ):4. van der Schalie, H. 19.54d. Cakin Goodrich, 1874-1954. Re- port to the Director of the Museum of Zoologv, Universih' of Michigan 56:9-14 (reprinted in 1955. The Nautilus 68(4):1.35-I40). van der Schalie, H. 1954e. The problem with snail control in Egypt. Report of the Director of the Museum of Zoology, Universitv of Michigan 56:35-38. van der Schalie. H. 195.5a. Bilharziasis control project at Qal- Mib. United Nations World Health Organization EM/BIIV 6:1-41. \an der Schalie. H. 19.5.5b. Discussion of the paper bv L.J. Olivier. American Journal of Tropical Medicine and Hy- giene 4(3):424-425. \an der Schalie, H. and D. S. Dundee. 1955. The distribution, ecolog\', and life histor\' of Painatiupsis cinciniiaticn.sis (Lea), an amphibious operculate snail. Transactions ot the American Microscopical Socieh' 74(2): 119-133, \'an der Schalie, H. 1955c. Calvin Goodrich (obituary). .Annual Reports of the .American Malacological Union for 1955:1. \an der Schalie. H 195.5d. Bilharziasis in Eg\pt. .Annual Re- ports of the .Amencaii Malacological Union for 1955:23. van der Schalie. H, 19.56a. Facilities for training malacologists and medical zoologists in the Uni\ersit\' of Michigan. United Nations World Health Organization Bilharziasis Conference 2:1-5. van der Schalie, H. and D. S. Dundee. 1956. The moiphology of Pomatioj)si.s cincinnaiiensus (Lea), an amphibious pro- sobranch snail. Occasional Papers. Museum ot Zoologv; Uiii\ersit\ of Michigan .579:1-17, \an der Schalie, H. 19.56b. Human blood fluke (schistosomi- asis) control in Egxpt. Annual Reports ot the .American Malacological Union for 1956:9. van der Schalie, H. 19.56c. Studies on Ponmtiojtsis shells in M. E. Gordon, 1998 Page 93 Michigan. Annual Reports of the American Malacological Union for 1956:22. 1 der Sehalie, H. 1957. Concerning the t'uhire of The Bi- ologist. The Biologist 40(1-2): lT.'"2.S-29. 1 der Sehalie, H. and H. J. Walter 1957. The egg la\ing habits of Pomatiopsis cinciniinticnsis (.Lea). Transactions of the American Microscopical Society- 76(4):4()-l — 122. I der Sehalie. H. 195Sa. Problems in control of intermediate hosts (Oncomclaniii and Pi»natioi>.sis) of the Oriental blood fluke. Annual Report ot the American Malacological Union for 1957:12. der Sehalie, H. 1958b. The effects of thim years of 'prog- ress" on the Huron River in Michigan, The Biologist 41(1):7-10. 1 der Sehalie, H. 1958c. X'ector snail control in QaKiib. Eg\pt. Bulletin of the World Health Organization 19:26.3- 283. I der Sehalie, H. and D. S. Dundee. 1958. Helix pomatia coionv at Jackson, Michigan. The Nautilus 72(1): 16-18. I der Sehalie, H. 1958d. Phil Lewis Marsh, 1891-1957. The Nautilus 72(2):64-66. I der Sehalie, H. and D. S. Dundee. 1959a. Egg la\ing habits of Pomatiopsis lapidaria and P cincinnaticnsis. problems relating to the culture of schistosome interme- diate hosts. Annual Reports of the .'American Malacological Union for 1958:19. 1 der Sehalie, H. and D. S. Dundee. 19.59b. Transect dis- tribution of eggs of Pomatiopsis lapidaria (Sa\), an am- phibious prosobranch snail. Transactions of the American Microscopical Societ\- 78(4):409-420. 1 der Sehalie, H, 1960a. Eg\pts new High Dam — asset or liability-. The Biologist 42(;3-4):6;3-70. I der Sehalie, H, 1960b. Pearls, food, and buttons: practical uses of Michigan mussels. Michigan Department of Con- ser\'ation Fisheries Di\ision Pamphlet 32:1-6. 1 der Sehalie, H. and P. W. Parniiilee. 1960. .\nimal remains from the Etowali site, mound C, Bartow Counts, Georgia. Florida Anthropologv' 13(2-3):37-54. 1 der Sehalie, H. 1961a. Peter Olaus Okkelberg, 1880-1960. The Nautilus 74(3):119. 1 der Sehalie, H. 1961b. Peter Olaus Okkelberg, 1880-1960 [continued]. The Nautilus 74(4):16.3-164. I der Sehalie, H. and L. L. Getz. 1961. Compari.son of adult and young Pomatiopsis cincinnaticnsis (Lea) in respect to moisture requirements. Transactions of the Amencan Mi- croscopical Societv 80(2):21 1-220. 1 der Sehalie, H. 1961c. The naiad (fresh-water nnissel) fau- na of the Great Lakes. LlniversiK' of Michigan Institute of Science and Technology' Great Lakes Research Dnision Publication 7:1.56-157. 1 der Sehalie, H. and L. L. Getz. 1962a. Distnbution and natural histor\- of the snail Pomatiopsis cincinnaticnsis (Lea). The American Midland Naturalist 68(1):20.3-231. 1 der Sehalie, H., L. L. Getz and B. C. Dazo. 1962. Hvbnds between American Pomatiopsis and oriental Oncomclania snails. American Journal ot Tropical Medicine ami H\'- giene 11(3):41 8-420. 1 der Sehalie, H. and L. L. Getz, 1962b. Moq^hology and development of sex organs in the snail Pomatiopsis cin- cinnaticnsis (Lea). Transactions of the American Micro- scopical Society 81(4):332-340. 1 der Sehalie, H. and L. L. Getz. 1962c. Reproductive iso- lation in the snails, Pomatiopsis lapidaria and P cincin- naticnsis The .\merican .Midland .Natur;dist 68(1): 189- 191. van der Sehalie. 1962. Mu.ssel distribution in relation to stream confluence in northern .Michigan. Annual Reports of the American Malacological Union for 1962:3. \an der Sehalie, H. and G. C. Robson. 1963. Bixalve. Five pages in Enc\clopffdia Britannica van der Sehalie, H. 1963a. Mussel distribution in relation to former stream confluence in northern Michigan, U.S.A. Malacologia l(2):227-236. van der Sch;Jie, H. 1963b. People and their snail-bome dis- eases. Michigan Quarterlv Re\iew 2: 106-114. van der Sehalie, H. 1963c. Comparison of temperature and moisture responses of the snail genera Pomatiopsis and Onconulania Ecolog\' 44(l):7.'3-83. \an der Sehalie, H. 1963d. Laboratorv guide and notes for medical malacolog\- bv Emile A. Malek (book review). .\merican Journal of Tropical Medicine and Hygiene 12(3):444-445, \an der Sehalie, H. and A, van der Sehalie, 1963. The distri- bution, ecolog)', and life histors' of the mussel, Arf»io/iflifl.s cllipsifonnis (Conrad), in Michigan, Occasional Papers, Museum of Zoolog\\ Universits' of Michigan 6.33:1-17 (ab- stracted in 1963, Annual Reports of the American Mala- cological Union for 1963:18). van der Sehalie, H. 1963e. Studies of the biolog\- of Poma- tiopsis and Oncomclania. snail intermediate hosts of ori- ental blood fluke (Schistosonta japonica). .Annual Reports of the Amencan Malacological Union for 1963:16. van der Sehalie, H. and G. m! Da\is. 1964. Stunting of On- comclania fonnosa in culture. Annual Reports of the American Malacological Union for 1964:13-14. van der Schahe, H. 1964. Notes on die sex of Campclonm. .\nniial Reports of the .American Malacological Union tor 1964:24-25, van der Sehalie, H, 1965a. Obsenations on the sex of C.am- pcloma (Gastropoda: \'i\iparidae). Occasional Papers, Mu- seum of Zoology, University of Michigan 641:1-15. \an der Sehalie, H. and G. M. Davis. 1965. Growth and stunt- ing in Oncomclania (Gastropoda: Hvdrobiidae). Malaco- logia 3(1 ):81-102. van der Sehalie, H. 1965b. Second European Malacological Congress, .'\nniia! Reports of the American Malacological Union for 1965:67-69. van der Sehalie, H, 1966a. The role of snail intermediate hosts in cultiiring Schistosoma japoniaim. Malacologia 5(1): 17-20. van der Sehalie, H. 1966b. Hermaphroditism among North American freshwater mussels. Malacologia 5(1): 77-78. van der Sehalie, H. and G. Pace. 1968. The freshwater Mol- lusca of Taiwan (Formosa). Annual Reports of the .Amer- ican Malacological Union for 1967:26-27. van der Sehalie, H. and G. M. Davis. 1968. Culturing Onco- mclania snails for studies of oriental schistosomiasis. Malacologia 6(3):321^327. van der Sehalie, H. 1969a. Schistosomiasis: control in Egvpt and the Sudan. Pages 62-65 in Fanar M. T and J. Mil- ton, eds. Special supplement: the unforeseen international ecological boomerang. Natural Historv 78(2):41-72. van der Sehalie, H. 1969b. Two unusual unionid hermaphro- dites. Science 163(3873):1.333-1334. van der Sehalie, H. 1969c. Man meddles with nature — Ha- waiian stvle. The Biologist 51(4):1.36-146. van der Sehalie, H. 1969d. Snail control problems in Hawaii. Page 94 THE NAUTILUS, Vol. 112. No. 3 .\iiiuui! Reports of the American Malacological Union lor 1969;5.5-.56. van der Schalie, H. 1970a. The control of schistosome der- matitis in the Great Lakes region (U.S.A.). Malacologia 9(1):44 (the vohmie pubhcation date for this and tlie fol- lowing reference was misprinted as 1969). van der .Schalie, H. 1970b. .American nnissel resonrces m re- lation to the Japanese pearl industiT. Malacologia 9{ 1):28.5. vail der Schalie, H. 1970c. Book re\ie\v: A guide for the iden- tification ot the snail intermediate hosts of schistosomiasis in the .Americas. Pan American Health Organization Sci- entific Publication 168, October, 1968. Journal of Parasi- tolog>-56(l):18.5-186. van der Schalie. H. 1970d. Hermaphroditism among North American freshwater nuissels. Malacologia 10(1):9.'3-112. van der Schalie. H. 1970e. .Mussels in the Huron River above Ann Arbor in 1969. Sterknana .39: 17-22. van der Schalie. H. 1971. Review: Terrestrial slugs by N.W. Riinham and PJ. Hunter. Science 17.3(4000):904. ' van der Schalie, H. 1972a. World Health Organization Project Egypt 10: a case histoiv of a schistosomiasis control pro- ject. In: Fai"var. M.T. and J. P. Mitton (eds.) The careless technologv': ecologv' and inteniational development. The Natural Historv Press, Garden Citv, pp. 116-1.36. van der Schalie, H. 1972b. Dam(n) large rivers. . .then what? Michigan .Alumnus (April):10-12 (reprinted in 1973, The Biologist .55(2):29-35). van der Schalie, H. 1972c. Problems in culturing snail inter- mediate hosts. Malacological Review .5(1): 10. van der Schalie, H. and E. G. Berrv. 1973a. The effects of temperature on growth and reproduction in aijuatic snails. Malacological Review 6(1):60. Liang, Y.-S., H. van der Schalie, and E. G. Beny. 1973. Trans- mission of ostracods in snails. Malacologiciil Review 6(1);66. van der Schalie, H. and E. G. Berry. 1973b. Effects of tem- perature on growth and reproduction of aquatic snails. Environmental Protection Agency Research Series, U.S. Government Printing Office, Washington, 164 pp. (reprinted in 1974, Sterkiana ,50:1-92; abstracted in 1974. Bulletin of the American Malacological Union for 1973:4.8 — 19). van der Schalie, H. 1973. Molhisks of the Duck River drainage in central Tennessee. Sterkiana 52:45-.55. van der Schalie, H. 1974a. Snail-related public health prob- lems in the Mekong. Bulletin of the American Malacolog- ical Union for 1973:.32. van der Schalie, H. 1974b. Tlie niollnsks of the Duck River drainage in central Tennessee. Bulletin of the Anu'ncan Malacological Union for 1973:49: van der Schalie. H. 1974c. Aswan Dam revisited. Environment 16(9): 18-20, 2.5-26. Liang, Y.-S. and H. van der Schalie. 1975a. Cultivating iJlh- <>-61(5):915-919. Liang, Y.-S. and H. van der Schalie, 1975b. (;ulti\'ating Fii.s- sarid huliinuidfs (Lea), a snail host lor the liver Hnke. Ftis- ciolfi luptitica. Malacological Revii-w 8(1-2): 123. van der Schalie, H. 1975. An ecological approach to rare and endangered species in the Great Lakes region. Michigan Academician 8(l):7-22. van der Schalie, H. and II. D. Blankespoor. 1976. Attachment and penetration of miracidia obsened bv scanning elec- tron Tiiicroscop)'. Science 191 (4224):291 -1293. viui der Schalie, H. 1977. Malacolog)- and [)urasitolog\- as the\- pertain to health. Bulletin of the .American Malacological Union for 1976:.5.5-57. van der Schalie, H. and H. D. Blankespoor. 1977. Potential use of solar energy for snail-control. The Biologist 59(1): 16-24. van der Schalie, H, 1978. Flukes that affect world progress. Wards Bulletin 17(l):l-,3 van der Schalie, H. 1979. Swimmers itch. Literature. Science, and .Arts (Uui\ersitv of Michigan) 2:-l— 7. van der Schalie, H 1981a. Is professional training too restric- tive, or can one earn a living working on snails? Spark (Cal\in College), September: 12-14. van der Schalie, H. 1981b. Fiftv years of malacologv' at the University of Michigan (1929-1979). BulleHn' of the American Malacological Union for 1980:1-5. van der Schalie. H. 1981c. A.swan revisited. BioScience 31(6): 420. van der Sch;ilie, H. 1981d. Pitst, present, ;md fiiture status of die MoUusca of the upper Tombigbee River. Sterkiana 71:8-11. van der Schalie, H. 1981e. Molhisks in the .Alabama River drainage: past and present. Sterkiana 71:2-1—40. van der Schalie, H. 1986. Mussels in the Detroit River op- posite Cobo Hall. Michigan Academician 18(l):31-.34. ACKNOWLEDGMENTS John B. Burch and Renee A. Sherman (MZUM), Walter R. Hoeh (Department of Biological Sciences, Kent State University', Ohio), and Fred G. Thompson (Florida Mu- seum of Natural Hi.storv, Unixersitv of Florida) provided much appreciated assistance and information during the preparation of this manuscript. M.G. Harase\v)'ch and Jose H. Leal, respectiveh' past and present editors of The Nautilus, and two anonvinous reviewers provided direction that improved the manuscript. Finally, this manuscript would have been considerably more difficult to prepare if it had not lieen for the large stack of re- prints and bibliographies given to me bv Van in 1979 during m\ first visit to the Museum of Zoologv', UM, and subsequent coninnmications. LITERATURE CITED .Abbott, R. T, ed. 1987. Register of American malacologists: a national register of professional and amateur malacologists and private shell collectors, second edition. .American Ma- lacologists, Inc., Melbourne, Florida. 168 pp. Atheani, H. D. 1964. Three new uniouids from .Alabama and Florida and a note on Lnmpsili.s /o/ics; The Nautilus 77(4):134-139. Hershler, R. and F. G. Thompson. 1992. .A review of the ai|U.it- ic gastropod subfamilv Cochliopinat' (.Prosobranchia: Hy- drohiidae). Makicological Review, supplement 5:1-140. |olinson. R. I, 1967, .Adtlitions to the nnionid fauna of the Gulf diaiuage of .Alabama, Georgia, and Floritla (Mollnsca: Bi- valvia). Breviora 270:1-21. Krai'Mier, L, R. and E. G. Beri-v 1986 Things trulv excel- lent. . .": Henn' van der Schalie. 1907-1986. The Nautilus 1()0(4):116-119. .Modell, H. 1943. Tertiare Na|aden III Archiv Inr .\Iolluskpe B, Shaskv Collection, 15.5X9.6 mm, same as Paratvpe A; Paratope C. SBMNH 144458, 10.4X5.9 mm, off Isla Vi- radores, Playas del Coco, Guanacaste, Costa Rica. 10°.34'N. 85°34'W, dredged 9-15 m, R. Koch coll.. 2-4 April 1986; Parat\pe D, Koch CollecHon, 10.2x6.0, same as Paratvpe C; Parat\pe E, AMNH 290728, 11. 5X7. .3 mm, S of Isla Chitre, Islas las Perlas, Panama, 8°36'N, 79°4'W, dredged 15-18 m, C. and R Skoglund and R. and W. Koch "coll., 15 April 1984; Parat\pe F, Skoglund Collection. 13.4X7.9 mm, same as Paratxpe E; Parat\pe G, Skoglund C'ollection, 12.2x7.8 mm, same as Paratvpe E; Paratype H, Skoglund Collection. 11.6X7.8 mm, same as Paratype E; Paratvpe I, Shask-\ Collection, 16.2X10.5 mm, Isla Venado, Panama, 8°52'30"N. 79°39'3(AV. J. McDaniel coll.. 8 Septemlier 1979; Paratype J, Shaskv Collection, 12.9X7.8 mm. N side of Isla Salango, Ecuador, 1°35'15"S, 80°52'52"W, D. R. Shasky coll., 15 Septemi)er 1978; Paratvpe K. Koch Collection, 11.5X7.2 nun. Isla Negritos Adentro. Golfo de Nicoya, Costa Rica, 9°57'N, 84%2'W, dredged 12-27 m, C. and P. Skoglund and R. and W. Koch coll., 8 Ma\ 1982; Paratype L, Koch Collection, 11.6X7.0 mm, Bah- liis de Iluatulco. Oaxaca. Mexico, 15°40'N, 96°08'W. dredged 9-30 m, R. and W. Koch , 2-4 June 1991; Para- type M, Koch (Collection, 15.5X9.6 mm, same as Para- tvpe E, 16 April 1984; Paratopes N-P; N, 14.6X9.0 mm; Paratvpe O. 12.7X8.2 mm; P, 12.5X7.3 mm, data for all same as Paratvpe E; Paratvpe Q. Skoglund Collection 10.1X6.9 mm,' off Isla Viradores Sur, Phivas del (loco. Guanacaste, Costa Rica. 10°.34'N, 85°34'W, dredged 9- 18 m. C. and R Skoglimd coll., April 1986; Paratvpe R. Skoglund Collection 12.1X8.2 mm, off Isla Rancheria, Golfo de Chiriqui. Panama, 7°35'N, 8r40'W, dredged 24-36 m. C. and P Skoglund coll., March 1986; Paratvpe S, Hertz Collection, 10.5X7.3 mm. Islas Tres Marietas, Navarit, Me.xico, 21°41'N, 105°36'W. diving in 13.7 m, K. L. Kaiser coll., 22 Febniarv 1995; Paratvpes T-U, K. L. Kai.ser Collection: T, 12.6x7.9 mm; U, 1().9X7.4 mm. Biiliia John Huston, Balii'a Banderas. Jalisco. Me.vico, 20°30.55'N, 105°21.11'W, 9 m. in sand under rocks. 2 Januar\- 1995. K. L. Kaiser coll. Type locality: Pedro Gonzales, Islas las Perlas, Pana- ma, 8°25'N, 79°05'W, 5.5 m depth. Distribution: Favaiiia (Muivxiclla) ptiulskafihiiuli is knowm to occur from Islas Tres Marietas, Na\arit, Mex- ico, its most northern locality, with an intermittent dis- tribution south to Isla Salango, Ecuador, in 5.5-36.0 m depth. Discussion: The new species was compared with the three s\iitvpes oi Fovaifia {Miircxiclla) cxi^iui (BMNH 19841227,' Figures 6, 7). Yokes (1988) considered F. (M) cxiiiuti to be a \alid species and referred to the illustra- tion of the largest SMitvpe figured in Yokes (1984. pi. 2, fig. 3), although she erroneousK' considered F (M.) rad- icfitn as a .s\Tionym of F. (M.) cxiguo in the 1984 paper. We have selected the largest specimen (16.2X9.4 mm) of the .sMitvpe series of F (A/.) cxi'^ua as the lectotvpe. Two smaller specimens (12.6X7.5 mm and 10.8X6.8 mm) are chosen as paralectotvpes. Favartia (M.) paul- skofjltiiuli differs from F. (M.) cxig^ua in having an indis- tinct suture and sloping shoulder with thickened varices, whereas F. (M.) cxigtia has an impressed suture and somewhat excavated slioukler with sliarpK' elevated var- ices. The new species has five strong major cords with one minor cord in each interspace, whereas F. (M. ) ex- C. M. Hertz and B. W. Myers, 1998 Page 97 Figures 6, 7. Favaiiin (MiircxicUa) exigiia (Broderip, 1833), BMNH 19S4122, sviihpe lot. Localin-: "PSalango". 6. Aper- tural view. 7. Dorsal \iew. Largest specimen, 16.2X9.4 mm, selected as lectot)pe. Two smaller specimens, 12.6x7.5 mm and 10.8x6.8 mm, selected as paralecto^y-pes. Specimens pho- tographed with kind permission of the trustees of The Natural Historv Museum, London. Figure 4, 5. Favaiiia (MiircxicUn) pniilskonliindi new spe- cies. 4. Paratvpe A, camera lucida drawing of protoconch (1.1. mm diameter). 5. Paratope A, camera lucida ilrawing of oper- culum (3.5 mm length). iiiua has fi\e strong cords with no minor cords in the deeply-cut interspaces, the edges of tlie cords somewhat overlapping the interspaces. Since Yokes (1970) stated that 'the tspe of A/. Itipjxi is no longer to be foimd," the new species was compared with the holot)pe of F. (M.) radicata (Hinds, 1844) (BMNH 1907.10,28.136, Figure 8), a junior synonym of F. (M.) lappa (Broderip, 1833) (Radwin and D'Attilio, 1976; Fair, 1976) and comparative material of K {M.) lappa in private collections. Favartia (M.) paulskoglundi differs from F. ( M. ) lappa in the number of spiral cords on the liod\- whorl, five major cords with strong minor cords between on the new species, and five major cords with two strong minor cords on the shoulder and no minor cords on the bodv whorl on F. (M.) lappa. In F. (M.) paulskog,hiiuli there are si.x to seven varices which do not obscure the indistinct suture, whereas in F. (M.) lappa the five varices project above and obscure the su- ture. The type species of F. (M. ) radicata had no re- maining protoconch. C'oniparisons between the new .species and F. (M.) vcnusfula (holotvpe, SDNHM 81610), show that F (M.) vcnii.stnla and F (A/.) cxigiia are conspecific. Both spe- cies are of comparable size with similar protoconchs ( F (M.) cxigua with slightly more than 2.5 smooth, convex whorls and F (M.) vcnusiula with 3 smooth, conve.x Page 98 THE NAUTILUS. Vol. 112, No. 3 Figure 8. Faiaiiia (Mitrexiella) rndicnta (Hinds, 1.S44). Ho- lotvpe, BMNH 1907.10.28.136, 19.8x11.7 mm. ApertunJ view. From: ". . .San Bias, west coast of Mexico. From 11 fms." Junior sviionym of Favartia (MiirexicUa) lappa (Broderip, 1833). Specimen photographed with kind permission of the Tnistees of The Natural Histon- Museum, London. whorls); both have teleoconchs with 5 whorls and 7 sharp varices crossini:; the suture to the preceding whorl. Both species have 2 spiral cords on the spire, 5 flattened spiral cords on the hodv whorl with incised threads along their length and deep interspaces lacking minor cords. The oval aperture with shallow anal sulcus and moder- ateK- long canal narrowh- opened distalK' are characters ot both species. Therefore F. (M.) vcnustula is consid- ered to be a junior 5)1100)011 of F. (M.) exigua. Etymology: The species is named for the late Paul Skoghmd of Phoenix, Arizona, who activeh' participated in the collection of deep-water Panamic species and de- signed a tube-pulley arrangement tor small boat dredg- ACKNOWLEDGMENTS Robert Koch, Donald R. Shaskv and Carol Skoglund made specimens available for stud\' and each donated paratxpes. Kirstie Kaiser also lent specimens for studv. The Natural Historx Museum, London, sent txpe ma- terial on loan. Henn' W. Chane\- (SBMNH) was helpful in providing coordinate information, David K. Mulliner photographed all the txpe specimens and Joxce Geni- mell made camera lucida drawings of details ot the new species. The San Diego Nattiral Histon Museum made their facilities available to us. Eniilx H. Yokes reviewed a draft of the manuscript and made helpful suggestions. To all ot them we e.xpress our gratitude. LITERATURE CITED Fair. Ruth H. 1976. The Murex Book: an Illustrated Catalogue ot Recent Muricidae (Muiicinae, Muricopsinae. Ocene- biinae). Author's edition, .\xi + 138 pp. Radwin, G. E. and A. D'Attilio. 1976. Murex Shells of the World an Illustrated Guide to the Muricidae. Stanford Universitx' Press, Stanford, 284 pp. Yokes, E. H. 1970. The west American species of Miircxiella. (Gastropoda: Mmicidae) including two new species. The Yeliger 12(3):325-329. Yokes. E. H. 1984. Comparison of the Muricidae of the eastern Pacific and western Atlantic, with cognate species. Shells and Sea Life 16(1 1):210-215. Yokes, E. H. 1988. Muricidae (Mollusca: Gastropoda) of the Esmeraldas Beds, northwestern Ecuador. Tulane Studies in Geologv' and Piileontology 21(l):l-50. INSTRUCTIONS TO AUTHORS THE NAUTILUS publishes papers on all aspects of the biolog)^ and systematics of mollusks. Manuscripts describing original, unpublished research as well as re\iew articles will be considered. Brief articles, not exceeding 1(M)0 words, will be pubUshed as notes and do not require an abstract. No- tices of meetings and other items of interest to malacolo- gists wall appear in a news and notices section. Manuscripts: Each original manuscript and accompanying illustrations should be submitted in triplicate. Text must be t\ped on one side of 8% X 11 inch white paper, double spaced throughout (including literature cited, tables and figure captions), with at least 1 inch of margin on all sides. All pages must be numbered consecutiveI>'. If printed on a word processor, the right margin should be ragged rather tlian justified. 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Leal The Bailey- Matthews Shell Museum 3075 Sanibel-Captixa Road Sanibel, PL 33957 EDITOR EMERITUS Dr. M. G. Harasewych Department of Invertebrate Zoology National Museum of Natural History Smithsonian Institution Washington, DC 20560 CONSULTING EDITORS Dr. Riidiger Bieler Department of Invertebrates Field Museum of Natural History Chicago, IL 60605 Dn Arthur E. Bogan North Carohna State Museum of Natural Sciences Raleigh, NC 27626 Dr. Philippe Bouchet Laboratoire de Biologic des Invertebres Marins et Malacologie Mu.seum National d'Histoire Naturelle 55, rue Buffon Paris, 75005 France Dr. Robert T. Dillon, Jr. Department of Biology College of Charleston Charleston, SC 29424 Dr. Wilham K. Emer.son Department of Living Invertebrates The American Museum of Natural History New York, NY 10024 Dr. Eileen H. Joldnen Instittite of Water Resources University of Connecticut Storrs, CT 06269-4018 Mr. Richard I. Johnson Department of Mollusks Museum of Comparative Zoology Harvard Universitv Cambridge. MA 02138 Dr. Douglas S. Jones Florida Museum of Natural Histor\' Universit)' of Florida Gainesville, FL 32611-2035 Dr James H. McLean Department of Malacology Los Angeles County Museum of Natural Histor)' 900 E.xposition Boulevard Los Angeles, CA 90007 Dr Arthur S. Merrill % Department of Mollusks Museum of Comparative Zoology Harvard Universitv Cambridge, MA 02138 Dr. Paula M. Mikkelsen Department of Lixdng Invertebrates The American Museum of Natural History New York, NY 10024 Dr Gustav Paulay Marine Laboratory University of Guam Mangilao, Guam 96923 Mr Richard E. Petit PO. Box 30 North Myrtle Beach, SC 29582 Dr Edward J. Petuch Department of Geolog)' Florida Atlantic University Boca Raton, FL 33431 Dr Garv' Rosenberg Department of Mollusks The Academy of Natural Sciences 1900 Benjamin Franklin Park\va\' Philadelphia, PA 19103 Dr. Ruth D. Turner Department of Mollusks Museum of Comparative Zoolog)' Harvard Universitv Cambridge, MA 02138 Dr. Geerat J. Vermeij Department of Geology Universit)' of Cahfornia at Davis Davis, CA 95616 Dr. G. Thomas Watters Aquatic Ecolog\' Laboratory^ 1314 Kinnear Road Columbus, OH 43212-1194 Dr John B. Wise Houston Museum of Natural Science Houston, TX 77030-1799 SUBSCRIPTION INFORMATION The subscription rate per volume is US $28.00 for incbviduals, US $45.00 for institutions. Postage outside the United States is an additional US $5.00 for surface and US $15.00 for air mail. All orders should be accompanied bv pavinent and sent to: THE NAUTILUS,' PO. Box 1580, Sanibel, FL 33957, USA. Chan'as- tellanos & Miquel, 1991), where the species inhabit rocky substrates covered bv moss and algae close to rap- ids and waterfalls. (.)iUiner (1937) described A, petricola from specimens collected at Nova Teutonia (27°16'S, 52°20'W), Santa Catarina, Brazil. Paraense and Deslandes (1959), based on radular moq:)holog\- and anatomy of the male genital system, treated A. petricola as a member of the genus Drcpanotrema, as previously suggested by Pilslirv (1938,1. Hvlton-Scott (1960,1 described a second species, Acrorbis odhneri, from specimens collected at Salto En- cantado (27°03'S, 54°50'W), located on the headwaters of Cunapini Brook, near Aristobulo del Valle, Depart- ment of Ciiinguas, central Misiones, Argentina, a site close to the Kpe locahtv of A. petricola. Moreover, Hvl- ton-Scott (1960) e.\cluded Acrorbis from the family Plan- orbidae and proposed a new family, Acrorbidae, to in- clude the 2 knowai species, based on a misinteipretation of radular moiphologv, especially that of the marginal teeth. After an analysis of shell and radular moiphologv of 23 specimens from Salto Encantado, the type-locality of Acrorbis odhneri, Paraense (1986) concluded that A. odhneri is a junior SMionvm of A. petricola, and provided evidence to support the inclusion of the genus Acrorbis in the family Planorbidae. Rumi (1986) reported the presence of the genus Ac- rorbis from Saltos de Apipe, Conientes, Argentina (27°27'S. 56°42'\\'), based on the collection of several dead shells, and noted a number of differences in shell moiphologv between the knowii species. The lack of soft parts did not allow her to determine the specific identity ot the reported specimens. This collection site, which is now under water and inaccessible due to the filling of the YacyTetii Dam empoundment, had environmental characteristics similar to those of previously knovvm lo- calities for the genus. In a review of Argentinean Plan- orbidae, Rumi ( 1991 ) considered A. odhneri to be a valid species, making no reterence to the specimens from Sal- tos de Apipe. The k-nown geographical range for the ge- nus Acrorbis had been limited to 1 locality in southern Brazil (Nova Teutonia, Santa Catarina State), and 2 lo- calities in northeastern Argentina (Salto Encantado, Mi- siones Province, and Saltos de Apipe, Corrientes Prov- ince). This paper reports a new record of Acrorbis pe- tricola from Salto Dos Hermanas, Iguazil Falls, Mi- siones, Argentina (25°35'S, 54°35'W), and includes data Page 104 THE NAUTILUS, Vol. 112, No. 4 oil ladular nioqihologx- ancl variahilitv in shell shape. Shell measurements and scannintj electron microiiraplis of the t\pe specimens of Acrorhis odhncri from the col- lection of M I. Ihlton-Scott, recently deposited in the Department of Invertebrates of" Museo de La Plata (MLP), are also given. MATERIAL AND METHODS Several specimens of Acrorbis petricola (MLP 5090) were collected on a basaltic cliff behind the waterfall known as Salto Dos Hermanas at the Iguazii National Park, Misiones, Argentina. The rocky w;i]l was densely covered by mosses and epilithic algae, whose rapid growth is favored by the permanent, localized sprav of water. The specimens were fixed in toto in a 10% for- malin solution immediately after being relaxed by brief immersion in warm water (50°C for 2-3 minutes). The specimens oi Acrorbis were collected together with oth- er gastropod mollusks such as Cliilina inc^asfoma H\l- ton-Scott, 1958 and Potainolithiis sp.; the samples were analyzed 3 years later. For this reason, the shells were serionsK- damaged by the corrosive action of the for- malin. However, 2 shells were preserved intact. Prior to chssection, specimens were rinsed for 24 hours in an aqueous solution of 5% formalin and 2% acetic acid. The radulae were extracted from the bucal mass under a ste- reoscopic microscope, treated for 2 hrs in a 10% potas- sium hydroxide solution, rinsetl in distilled water and mounted on stubs for observation under scanning elec- tron microscope (SEM). Shell measurements were taken following the criteria: shell length and shell width as the maximum and minimum diameters respectively, taken along the basal shell plane; shell height as the maximum perpendicular distance between the basal shell plane and the apex (or the most distant point on the shell sur- face); aperture length as the maximum chstance across the apertin-e and parallel to the maximum diameter axis, Sliells of 7 s\Tit)pes of A. odhncri (MLP 5089) from Salto Encantado, Caingiias, central Misiimes were mea- sured and photographed under the SEM for compara- tive purjioses. RESULTS Description of specimens from Iguazii, Mi.siones (Figures 1—4): Shell small, \vr\' low, depressed, not strictly turbinate or hehcoidal (Figures 1-3), with up to 2.5 rapidly expanding whorls. Two well-preserved shells measured: 2.6.5 and 2.(S unn in length; 2.15 and 2.2 mm in width; 1.2 and 1.1 nun in height. Apex lateralK dis- placed due to ontogenetic change in coiling angle, apex not highest point on spire. Shell wide, flat at base, with rapid expansion of body whorl, which occupies = % of spire. Aperture wide, D-shaped, with inner margin slightly curved. Aperture length excei'ds half shell length (65 and 69% of shell length). Inner and outer lips shaip Umbilicus only partially covered by small basal reffection o( inner li]i Shell wiill thin, with oliw-brown to yellowish periostracum. Shell-surface sculptured, with oblique striae and smooth spiral lines (Figure 4). External anat- om\': Foot with quadrangular base, rectangular or sub- rectangular anteriorly, wideh' rounded posteriorly, dif- fusely pigmented. Head uniform gray, pigmentation darker between tentacles. Mantle unpigmented over lung, darkly pigmented over albumen gland and distal end of xisceral mass. Genital ducts: Due to poor fixation and preservation, only male terminal genitaha is de- scribed. Penis, unarmed, with sharply pointed tip. Pre- puce about twice as long as penis sheath. Flagella paired, short, wide, somewhat romided, attached to base of vergic sac. Flagella half as long as vergic sac. Retractor and protractor muscles fused together before insertion into lateral wall of vergic sac (Figine 8). ,Alimentar\- sys- tem: Salivary glands arise dorsallv from buccal sac, run posteriorly, passing through nene ring. Pair of sahvary glands join to form sniiill loop behind cerebral comissure (Figure 9). Esophagus arises from dorsal wall of buccal sac, nms posteriorly, passing under cerebral commissure. Intestine passes over albumen gland. Jaw single, horse- shoe-shaped, somewhat enlarged at free ends (Figure 10), consisting of numerous, small, cylindrical segments, highest (= 50 |jLm) at central part of organ, lower highest (= 35 jxm) at ends. Radula folded along most of its length, radular sack tubular, \en long, extentling bcN'ond posterior end of buccal mass, reflected dorsally (Figure 9). Each row (jf teeth composed of 1 central tooth, 13- 14 teeth on each side (Figure 11). Central tooth bicus- pid, left cusp longer than right, with 1 spine-shaped den- ticle between cusps, 1 denticle along each outer edge of tooth (Figures 12, 13). Lateral teeth (5-7) each with 3 cusps (Figures 14, 15): mesocone, rhomboidal in shape, with major axis elongated, longer than entocone, ecto- cone, foi-wardly displaced with respect to their bases. Ectocone, entocone robust, prong-like, with outer mar- gin more curved than inner margin. Margin stndght in ectocone. Entocone generalK' slightK- longer than ecto- cone. Mesocone firmly attached to center of tooth plate by means of wide, median base (Figure 15). Mesocone tapering markedly towards base (Figure 15), entocone, ectocone with relatixelv wide bases. Small Init robust accessory cusps develop on either side of mesocone, out- side of ectocone and entocone. Other minor cusps are added to mesocone and ectocone, entocone in 2-3 in- termediate teeth that follow (Figure 16). Robustness of cusps decreases progressivcK' from first lateral to mar- ginal teeth. The marginal teeth (generally 4), with mul- tiple accesson cusps (usualK' 9-10") added to mesocone, ectocone, and entocone (Figures 16-17). Ectocone and entocone fade toward radular row margin, beconnng nearh' indistinguishable from ;iccessory cusps. Description of sMit\pcs of A. odhncri Hvlton-Scott, 1960 (MLP 5089) (Figures 5-7): Txpc'-senes of A odhncri composed of 7 well-presei-vcnl shells, 4 adults, 3 juveniles. Shells pale brown or amber, \er\- thin, nearly translucent. Suriace sculptured in several specimens, with weak s]Mr:il lines superimposed onto transverse. C. F. Ituarte, 199S Paee 105 Figures 1-7. Shells of Avivrliis pdrkola Odhner, 1937. 1—1. specimens from Igiiazii Falls. Vlisiones, Argentina {MLP5090). 5- 7. S)iitrvpes oi Acroil)ifi odhncri H\lton-Scott, 19fi() ( Anstohnio del N'alle, Misiones, Argentma) (MLP 50.S9). Scale bars tor all figures = 1 mm. Page 106 THE NAUTILUS, Vol. 112, No. 4 Figures 8-9. Some a.spect.s ot the siitt part aiiatoni\- of Ar- rorbis pctricola. 8. Penial comple.x. Scale bar = 0.5 mm. 9. Buccal .sac. Scale bar = 1 mm. Abbre\iations: bs, buccal .sac; eg, cerebral gauglion; dd, deferent duct; e, esophagus; fl, fla- gella; pe. penis; pp. prepuce; ps, penis sheath; rs, radular sac; sg, sali\ary glands (the arrow indicates the small loop behind the nerxe ring). ohhque striae that are present in all shells. Shell is t\p- ically helicoid. Spire relati\ely elevated, with up to 3V2 whorl.s separated by deep suture. Aperture nearly cir- cular in shape, ver\- ol)li(jue. vvath inner niartjiTi straisjht. Umbilicus covered h\ weaklv developed lip reflection. Measurements of 4 adult shells shown in Talili- 1. DLSCUSSION The .specimens oi Acrorbis pdricolu collected at Iguazil r'alls .show several differences in shell morjiholog)' and radular tooth shape from those of published descriptions for this species. The shell ol A. pctricohi trom Iguazii is low and depressed, its apex laterally displaced and not prominent. The plane of the aperture is coincidcMit with the ba.se of the shell (Figures 1, 3). These characters differ from the t\pic;il, helicoidal shell-shape of A. pc- tricola described and figured from Nova Tcutonia, Brazil and Aristobulf) del Valle, Argentina (Figures 5, fi). In specimens from Itiuazii. the spire is less (Icvdopcd than mm length X in specimens from the type locaHty, the shell does consist of more than 2.5 whorls, and the aperture is markedlv D-shaped (Figure 3) rather than circular or subcircular as in other localities. The Iguazi'i Falls specimens also have a wide umbilicus that is covered by a small reflec- tion of the inner lip. With respect to the shell shape, the specimens from Iguazij are similar to those described bv Rumi (19S6) from Saltos de Apipe, Corrientes, which were diagnosed as having: a shell that was helicoidal in shape but with a flattened base; a teleoconch of not more than 2.5 whorls; a rapid whorl expansion rate: an umbilicus not covered by a reflected lip; a shell surface sculptured with weak spiral Unes; small adult size (1.12 0.91 mm height). The variabilitv in shell moiphology observed in A. pc- tricola seems correlated with habitat characteristics. The 4 localities at which this species was recorded are all associated with high-energy water courses (Hylton-Scott, I960; Paraense, 1986: Rumi, 1986). The spray and or How of water from waterfalls or high energv' streams onto rock^ banks contributes to the rapid growth of moss and algae on which A. pctricola feeds. However, these hvdrocKiiamic forces mav also be capa- ble of dislodging specimens. This is particularlv evident in the case of the specimens collected at Salto Dos Her- manas, Iguazu Fails. Here, the population of A. petricola is e.xposed to powerfid water forces that intermitently sweep the vertical rock-\- cliff behind the water fiill (> 20 m high). Tnissell ct al. (1993) studied a population of Littorina ohtusata living on a wave exposed intertickil rocky shore, and found that exposure to high wave action mav induce intraspecific variation in the shell and foot moiphologv so as to reduce the shells coefficient ot drag. Similarly, there seems to be selection in A. petricola for flat shells and increased basal area (apertural plane tan- gential t-o the outer edge of the shell) to decrease hy- drodviiamic lesistance in response to the force of the water flow. This is likelv' to occur in specimens of A. pctricola from Iguazii Falls (and Saltos de Apipe, Cor- rientes) were water flow is verv swift. The radular moiphologv of A. pctricola from Iguazii is similar to that previouslv ilescribed (Paraense & Des- landes, 1959; Paraense, 1986). However, the shape of the teeth differs slightly but constantly, particularly along the mesocone of the lateral teeth, which is rhomboidiil rather than havin" lonii dasjijer-hke cusps, as described aTid figured by Paraense (1986). The cusps tend to be shorter and more robust in the Iguazu F;ills specimens. The gross anatomical features of the Iguazu Falls specimens generallv agree with the detailed decriptions given by Paraense & Deslandes (1959) and Paraense (1975). However, several nimor differences were ob- seived in the Iguazu Falls material. The posterior ends of tln' salivan glands, which pass through the nene ring, fusi- together to form a small loop, which seems to be significantiv shorter than previouslv figured. The flagella appear to be shorter and more rounded than previously described. These difference's, as well as those reported C. F. Ituarte, 1998 Page 107 Figures 10-17. Scanning electron niicrojjraphs of the jaw and radula ot Acrorhis pctncola tiom Ignazii Falls, Misioiies Argentina (MLP 5090), 10. Jaw. Scale bar = 100 \x.m. 11. Dorsal \iew of half radnla. 12-1.3. Central and first lateral teeth. Tlie central teeth in figvn'e 12 are somewhat abnoniial. 14-1.5. Lateral teeth. 16. Internietliate (nght) and niargnial (left) teeth. 17. marginal teeth. Scale bars tor figures 11-17 = 10 |jirn. Page 108 THE NAUTILUS, Vol. 112, No. 4 Table 1. Measurfinents of adult shells in the tvpe-series of Acrorlm odhwri Hvlton Scott, 1960 (MLP 50S9) {=Acrorbis petricoUO (all measurements are given in millimeters) Shell-length Sliell-widtli Shell-height Aperture length 3.6 3.0 2.0 2.0 3.65 3.0 1.9 2.0 3.7 3.1 1.9 1.95 .18 3.3 ■l.\ 2.1 for the length ratios calculated tor the vergic .sac aiul prepuce, mav be due to post mortem changes during fixation of muscular organs and structures without skel- etal support. It is the autlior's opinion, given the evi- dence here presented and re\aewed, that Acrorbis pe- tricola and A. odhncri are conspecific taxa. As previously reported, A. pctricola is known onlv spe- cific habitats always associated with high-energv' water courses such as rapids and water tails (Paraense & Des- landes, 1959: Hylton-Scott, I960; Rumi, 1986). Such particular emironniental requirements ma\ be the tactor responsible tor the Hniited geographical distribution of this species. Given the abundance of rapids and small waterfalls in Misiones Pro\ince, a survev of the mala- colofflcal tauna of the resjion would most likelv result in the extension ot the kiiown geographical range of A. pc- tricola. ACKNOWLEDGMENTS The assistance of Dr. Gustavo A. Darrigran and Santiago Ituarte during the field trip and sampling is here ac- knowledged. LITERATURE CITED Ageitos de Castellanos, Z. J. and S. E. Miquel. 1991. Distri- bucion de los Pulmonata Bassoniniatophora. Fauna de agiia dulce de la Republica Argentina 15(9):.3-11. Baker. F. C. 1945. The niolluscan famiK Planorbidae. Uni\er- sit\ ot Illinois Press, Urbana, .\xxvi -I- 530 pp. H\lton Scott, M. I. 1960. Nueva familia de pulmonado baso- matoforo. Neotropica 6(21):65-69. Hubendick, B. 1955. Phvlogeny in the Planorbidae. Transac- tions ot the Zoological Society- t)f London 2:45.3-.542. Odhner, N. H. 1937. Acrorbis pctricola n.gen. n.sp., eine nierk- liclie Siis.swasserschnecke aus Brasilien. Arldv tiirZoologie 29B:l-8. Paraense, W. L. 1975. Estado atual da sistematica dos planor- bideos brasileiros. Arqui\os do Museu Nacional 55:105- 12. Paraense, W. L. 1986. The radula o'i Acrorbis pctricola (Pul- monata; Planorbidae). The Nautilus 10()(3):1()9-112. Paraense, W. L. and N. Deslandes. 1959. The Brazilian species of Drepanotrema. Ml. D. pctricola (Odhner, 1937). Re- vista Brasileira de Biologia 19(3):319-.329. Pilsbrv, H. A. 19.38. Acrorbis pctricola. Review. The Nautilus .51:107. Rumi, A. 1986. Estudio mortologico, taxinoinico \ bioecologico de ios planorbidos argentinos. Tests Doctoral n° 461, Fa- cultad de Ciencias Naturales v Museo, Universidad Na- cional de La Plata. Rumi, A. 1991. La familia Planorbidae Rafinesque, 1815 en la Republica Argentina. Fauna de agiia dulce de la Repi'ib- lica Argentina 15(8):.3-51. Tnissell, G. C A. S. Johnson, S.G. Rudolph and E. S. Gilfillan. 1993. Resistance to dislodgement: habitat and size-specific ditferences in niorphologv and tenacitv in an intertidal snail. Marine Ecolog)- Progress Series 100:135-144. THE NAUTILUS 112(4):l()y-112, 199^ Page 109 A New Species of Land Snail of the Genns Georissa (Gastropoda: Hydrocenidae) from the Phihppine Islands Kurt Auffenberg Division ot Invertebrate Paleoiitolog)' Florida Museum ot Natural History Uuiversitv ot F"lorida Gainesville, FL 32611 USA kauffe(S'HMuili.uH.edn ABSTRACT Gforissa cavini new species is described from an isolated lime- stone outcrop in the northeastern extremitv ot Panav Island. Philippine Islands. The new species is characterized by its rel- atixelv large size and sliell sculpture of weak spiral threads which become increasingly oblicjue below the peripheiy. It is most similar to Georissn coccinra Quadras and Moellendortt. 1895 from Ma.sbate Island. Philippine Islands. Georissa cveri'tii E. A. Smith, 1895 from Sarawak has a similar shell, but much stronger shell sculpture. Key Words: Hydrocenidae. Georissa. new species, Philippine Islands, Panav Island, Ma.sbate Island. INTRODUCTION The author conchicted a .survey of the terrestrial niol- kisks of Panay Island, Visavan Islands, in the central Phihppines, during April-May, 1992. Fiftv-five sites were sampled, with emphasis on the small species found in the widespread limestone areas of Panay and its satellite islands. Onl\ al)out 52 species and sulispecies ot terrestrial mollusks have been recorded from Panay (Bartsch, 1919, 1938, 1939, 1942; Moellendorff, 1898; Smith, 1932a, b). This is a small mmiber, when compared to landmasses of similar size in the Philippines. However, much of the island consists ot low, rolling hills (noyv mostly converted to agriculture), which do not provide the varied habitats of other areas of the Philippines. Substantial elevations and a greater dixersitv ot habitats ta\()ralile to terrestrial mollusks occur only in the extreme western portion of the island. Most t;L\a previously recorded from Panay are still extant, but their chstributions are now restricted to small plots of remnant forest, except those of the com- parativeh' pristine higher elevations (> 8()()m) in the west. Few undescribed species were encountered. Geo- rissa is a common faunal component throughout the Phihppines, vet all specimens of this genus found in Pa- nay belong to a single, undescrilied species. This new species is here named and described. The type-series and other specimens e.xamined are de- posited in the Philippine National Museum (PNM) and Florida Museum of Natural Histon', Universitv of Flor- ida (UF). SYSTEMATICS Class Gastropoda Cuvier, 1797 Superlamily Neritoidea Rafinesque, 1815 Family Ilvdrocenitlae Troschel, 1856 Genus Georissa Blantord, 1864 Georissa cavini new species (Figures I, 3, 5, 6) Description: Shell (Figure 1) medium-sized for genus (mean = 2.4 mm length, 1.8 mm width), turbinitorm, ratio length/width about 1.33. Shell with 3.6 whorls. Em- bryonic whorl bulbous; subsecjuent whorls convex. Su- tures deeply impressed; sutural channel thstinct on early whorls, occasionidlv becoming very shallow on last hall whorl. Sculpture of embryonic yvhorls a fine mesh oi pits. Teleoconch glossy, with growth threads of variable strength, crossed by e.xtremeh' fine obh(jue spiral threads that become increasingly oblique below periphery. Groyvth striations more distinct on body whorl; spiral threads stronger on upper whorls (Figure 3). Aperture subovate, ratio aperture lengtli/shell length about 0.42. Parietal wall King at an angle of about 27-38° (mean = 34°) to iLxis of shell; plane of aperture in lateral profile at about 28-34° (mean = 31°) to shell axis. Umbilical area indented; basal area with y\ide concave columellar shield; edge of basal fip extending to left in an arc, form- ing thin, but prominent ridge terminating in umliilical resiion, borderinti columellar shield on left. Umbihcal callus concave to very slightly- c(3n\ex. Peristome thick- ened internally, usually more so near base ot columella and upper insertion (Figure 1). Outer lip slightly sinuose in lateral profile. Shell color Imck-red to orange, usually- more intense on spire; inner edge ot fip and insertion area with rose red border in live-collected specimens. Operculum (Figure fi) calcareous, concentric with sub- central nucleus. Inner surface with relatively long, slen- Page 110 THE NAUTILUS, Vol. 112, No. 4 Figures l-(>. Cforissa cavini new .species and G. coccinea Quadras and Moellendorff, LS95. 1, 3. C. cavini. holuhpe, 2.3 mm length, PNM-CO 39481. 2, 4. G. coccinea, 1.8 mm length. UF 87296. 5. G. cavini. parahpe, jinenile, 1.3 mm length, UF 250000. 6. G. cavini, internal surface of operculum, parat\pe, 0.9 mm opercnhnii length, UF 267482. K. Auffenberg, 1998 Page 111 Tabic 1. Adult .shell measurements (mm. converted from oc- ular iincronieter nnjt.s) of .selected .specimens ofGcorissa cavini new species (holot\pe in parentheses) and Georissn corciiwn (UF ST29fiV L = lenijth, W = width. Ap = aperture G cmiiii G. coccuwa Number of specimens 1 3 L 2.2-2.8 (2.3) 1.7-1.8 \y 1.8-2.0 (1.8) 1.4 \\7L 0.70-0.82 (0.77) (),77_0,84 ApL 0.9-1.0 (1.0) 0.7 ApW ()..SU0.9 (0.8) 0.6 ApW'/ApL 0.76-0.84 (0.76) 0.83-0.91 ApL/L 0.37-0.47 (0.44) 0.37-0.44 Number of \Miorls 3.2-3.9 (3.8) 3,2 der peg along columellar margin. Peg King at low angle to plane of operculum; laterally compressed, interior surface of peg narrower than base; base extending to columellar edge of operculum. Inner smface of oper- culum Hat with slightly raised callus along edge. Outer surface covered with thin chitinous sheet e.xtending well be\ond edge of operculiun. T>pe material: Holot\pe, PNM-CO 394.S1, 2.3 mm length X 1.8 mm width, (coated for scanning electron microscop\); Paratopes: UF 250000. 12 paratopes; UF 267482, 2 parat)pes (shells anil opercula coated for scan- ning electron microscopx ). The holot\pe and parahpes were collected alive. Type locality: Phihppine Islands, Panav Island, Iloilo Pnnince, ca. 10 km NE ot Balasan. shaded limestone cliff face, limestone outcrop about 5 m above sea level on east side of the road between Balasan and Carles (GPS: 11°29'15"N, 123°()()'27"E). Other material examined: UF 267481, 53 shells, col- lected dead at t\pe locality, in soil sample taken at the base of the hmestone outcrop. EtymologA': I take great pleasure in naming this spe- cies in honor of Mr. Edward W. Caxin of Jacksonville, Florida. Mr. Cavin not onlv assisted in the collection of the t\pe series, but has shared with me the rigors of fieldwork on several occasions, usually under spartan liv- ing conditions and with far less than adequate means of transportation. Comparati\e remarks: The shell sculpture of Gco- rissa cavini new species is most similar to G. coccinca Quadras and Moellendorff, 1895 (Figures 2, 4), only re- ported from Palanoc, Masbate (Quadras and Moellen- dorff, 1895:88), probably currently known as Palanog, a village located in the lowlands about 8 kilometers SSE Masbate Citw Ceorissa coccinca has a smaller, broadK' conical shell with a proportionately shorter spire than G. cavini (Figure 2, Table 1). The former usualK' has about 0.4 fewer whorls (Table 1), which are decidedK more convex in outline (Figure 2). The prominent ridge bor- dering the columellar shield in G. cavini is \eiy thin and erect. When present in G. coccinca this ridge is weak and low. Juvenile G. cavini and adult G. coccinca of equivalent shell size are superficially similar, but the for- mer are more globose, ha\e a siinple, unthickened peri- stome and a proportionately larger aperture (Figure 5). Opercular characters of G. coccinca are unknown. Geo- rissa cvcrctti E. A. Smith, 1895 of Sarawak has a similar, but much stronger shell sculpture. The shell of G. cv- crctti is smaller (1.95-2.20 mm length), with tleepK' im- pressed sutures and a more broadly conical shell shape (Thompson and Dance, 1983:120-121, figs. 55-57), Large (> 2.4 mm length) individuals of G. cavini have more protruding protoconchs and whorls which descend more rapidlx' than those ot typical adult shells, devel- oping a longer spire and a more elongate shell. A con- tiniuun is presented in the type series and, besides these differences, the shells are identical to the other exam- ined material. DISCUSSION The H\drocenidae are wideK' distributed in southeast- ern Europe, Africa, Madagascar, much of southern Asia, northern Australia, New Zealand, and some Pacific Is- lands. Some taxonomic confusion exists at the genus lex- el, but most Asian species have been assigned tradition- ally to Gcorissa. Most hvdrocenids are obligator\- inhab- itants of limestone rockfaces and man\" ha\e extremely limited distributions. Collectors, because of the small size of h\drocenids, may easih' owrlook these snails. Specimens can be most successfully found through close inspection of hmestone rock faces and soil samples. Un- doubtedK', many undescribed species e.xist throughout the vast range of the group. Available information indi- cates that the highest diversitx' of Gcoris.so occurs in the Philippines (11 species; this paper and Moellendorff, 1898, but 3 of these species are better assigned to the Assimineidae, see below) and Borneo (14 species; Thompson and Dance, 1983). Gcorissa cavini was the onl\' h\'drocenid collected during the sur\'e\' of Panav, despite thorough searches in many limestone areas throughout the island. Dead shells were not foimd even though soil samples were taken at most collecting sites. Several series of Gcorissa snbjila- hrata Moellendorff, 1887, which has been recorded from numerous locahties throughout the Philippines (Moellendorff, 1898), were collected. However, this spe- cies and probabK- hvo other \er\' similar taxa, G rcffi- laris Quadras and Moellenck)rif, 1895, and G turritclla Moellendorff, 1893, are properly assigned to the Assi- mineidae (Auffenberg, pers. obs.). Sur|)risingly, Gcorissa (Icnsclirata Moellendorff, 1894, which is known from other Visavan Islands, including Negros. Cebu and Rom- bl(jn (Zilch, 1973), was not encountered on Panav. ACKNOWLEDGMENTS Pedro C. Gonzales, Jaime J. Cabrera and Patemo Com- intan of the Philippine National Museum assisted during Page 112 THE NAUTILUS, Vol. 112, No. 4 all phases of the survey. Robert S. Kennetlv, Cincinnati Museum of Natural Historv, initiated the Philippine Bio- cli\ersit\' Iu\enton Project, which was luntled tliroutjli generous support troni the John D. and Catherine T. Mac-Arthur Foundation. Luis A. Ruedas, Museum of Southwestern Biolog\', University' of New Mexico, facil- itated many aspects of the fieldwork. Many persons in the Philippine Department (A En\ironnient and Natural Resources, Protected Areas and Wildlife Bureau deserve acknowledgment for their assistance with logistics and collecting permits. SEM micrographs were made with an Hitachi S-41.5A scanning electron microscope in the Department of Zoologv, Uni\ersit)' of Floriila Tins field- work was conducted with financial support from the Thomas L. McGintv Endowment Fund, University of Florida Foundation. LITERATURE CITED Bartsch, P. 1919, Critical remarks on Pliilippine landsliells vvitli descriptions ot new forms. Proceedmgs of the Biological Sot'iet\- of Washington .32:15-20. Bartsch, P. 19.38. A synopsis of the Philippine land moliusks of the subgenus Rijssota. Proceedings of the Biological So- ciety of Washington 51;101-120. Bartsch, P 1939. A synopsis of the Philippine land moliusks of the subgenera Lamarckiella and Pararyssota of the genus Rtjssota. Proceedings of tlie Biological Societ\' of Wasli- ington 52:41-56. Bartsch, P. 1942. A synopsis of the Philippine land Uioliusks of the genus Hcmitrichia. Proceedings of the Biological So- ciety of Washington 55:27^4. Moellendorff O. v. 1898. Verzeichniss der auf den Pliilippinen lebenden Landmollusken. Abhandlungen der natnrtor- schende Gesellschaft zu Gorlitz 22:26-208. Quadras J. F. and O. F. \\ Moellentlorff 1895. Diagnoses spe- ciennn no\anmi e.\ insniis Philippinis. Nachrichtsblatt der deutschen Malakozoologischen Gesellschaft. 27(.5-6):7.3- 88. Smith, M. 19.32a. Notes upon Philippine land shells with de- scriptions of new forms. Tlie Nautilus 45(3): 102-104. pis. 7-8. Smith, M. 19.32h. New Philippine Island land shells. The Nau- tilus 46(2):62-65, pi. 4. Thompson, F. G. and S. P. Dance. 1983. Non-marine moliusks of Borneo. II Pulmonata: Pupillidae, Clausiliidae. Ill Prosobranchia: Hydrocenidae, Helicinidae. Bulletin of the Florida State .Vluseum. Biological Sciences 29(3):I01-152. Zilch. A, 1973, Die T\pen and Tvpoide des Natur-Museums Senckenberg, 52). Mollusca: Hydrocenidae. Archi\' fiir Molhiskenkunde 103(4/6):263-272. THE NAUTILUS 112(4):113-116, 1998 Page 113 CanceUaria (EucUa) hiiirettae, a New Species of Cancellariidae (Mollusca: Neogastropoda) from Western Panama Richard E. Petit M. G. Harasewych Departineiit of Invertebrate Zoology' National Museum of Natural History Smithsonian Institution Washinsiton, DC 20560-0118 USA ABSTRACT CanceUaria (Eiiclia) laurettae new species, is described from eleven specimens dredged in tiie Golfo de Chiriqui, Panama at depths of 270-360 meters. This species is assigned to the subgenus Eiiclia based on the moipholog\' of its colvmiellar plications, spiral sculpture, and o\erall shell shape. It is readiK distinguished troni the other Recent members ot the subgenus b\ its lack ol a distinct shoulder vvnth spines or nodes and finer spiral sculpture. Key words: Cancellariidae. Panamic Pro\ince, eastern Pacific, Neogene, batlnal. INTRODUCTION This paper describes a new species of cancellariiil that was recenth' trawled from bath\al depths in the Golfo de Chiriqui. Panama. Based on conchological features, it is assigned to the genus CanceUaria Lamarck, 1799. subgenus Eiiclia H. & A. Adams. 1854, which, in its Recent geographical range, is endemic to the tropical eastern Pacific. While similar to the Recent species Can- ceUaria {Eiiclia) cassidifonnis Sowerby, 1832, and C. (£. ) halboae Pilsbrv, 1931, this new species is more similar to the Miocene C. (E.) dinota Woodring, 1970, from the Gatiin Formation of Panama and especially the Late Miocene/Earlv Pliocene C. (E.) maldonacloi Olsson, 1964 of the Angostura Formation of Ecuador than to either of the Recent species. The taxonomic historv of the subgenus Eiiclia is briefl\' reviewed, and a table of included species provided. ABBREVIATIONS USED AMNH American Museum of Natural Historw New York ANSP Academy of Natiual Sciences, Philadelphia LACM Natural Historv Museum of Los Angeles Coimtw Los Angeles USNM National Museum of Natural HistoiT, Smith- sonian Institution, Washington, DC. SYSTEMATICS Family Cancellariidae Forbes & Hanley. 1S51 Subtamily Cancellariinae Forbes & Hanley, 1851 Genus CanceUaria Lamarck, 1799 Cancrllaria Lamarck. 1799. T\pe species: Vohda reticulata Linne. 1767, b\ monotvpy. Subgenus Eiiclia H. & A. Adams, 1854 Eticlia H. & A. Adams, 1S54; 277. T\pe species: CanceUaria cassidifonnis Sowerby. 1S32. by subsequent designation of Cossmann, 1899). Table I. Species assigned to the subgenus Eiiclia. together with their geographical and geological ranges. Panamic CanceUaria {Eiiclia) cassidijonnis Sowerby. 18.32 — Gulf ot Cahfonna to Peru, RECENT C. {E.)balboae Pilsbrv', 1931— Mexico to Panama, RECENT C. (£. ) laurettae. new species — Gulf of Panama. RECENT C. (£.) harpifonnis Pilsbrv & Olsson. 1941 — Ecuador. Canoa Fonuation, PLIOCENE C. (£, ) larkinii Nelson, 1870 — Peni. Tunibes Formation, LATE MIOCENE C. (£. ) triaujiularis Nelson, 1870 — Peru, Tumbes Fonuation. LATE .MIOCENE C- (£. ) dinota Woodnug, 1970 — Panama, Gatiin Formation. LATE MIOCENE C. (£. ) maldonadoi Olsson. 196-1 — Ecuador. Angostura For- mation, LATE MIOCENE Caribbean C. (£. ) codazzii .Anderscjii, 1929 — Northern Colombia, Tii- bara Group. L.-^TE MIOCENE-EARLY PLIOCENE C. (£. ) venezuelana Hodson, 1931 — Falcon, \'enezuela, UP- PER MIDDLE MIOCENE C (£. ) montserratensis .Vlaur\', 1925 — Trinidad, Springvale Fonuation, EARLY PLIOCENE C. (£. ) tcerenfelsi Jung. 196.5 — Wnezuela. Cantaure Fonua- tion. LATE EARLY MIOCENE Page 114 THE NAUTILUS, Vol. 112. No. 4 Figure.s 1-2. Q'ancdlarUi {Eiirlia) laurctiat' new species. 1. H()l()t)pe. USNM 'ossmann's ( 1S99) design;i(ion lA Cancellaria cassidifor- R E. Petit and M. G. Harasexweh, 1998 Paet- 115 Tabic 2. Shell nieasiirenients of Caucellaria (Euclia) Inurrttae lieu species. All linear measurements in mm. The measure- ments are hased on the entire t\pe series, consistiiiij on le holot\pe and 10 paratopes (N = II). Figures 3-4. Cancellaiia [Euclia ) laurettde new species. 3. Lateral and 4. apical \iews of protoconch of holotspe. Scale bars = 500 jam. mis as type species of Euclia to he an unfoitmiate nus- inteqiretation of H. & A Adams intent, because, of the 4 species uriginallv included m Euclia by these authors, it ciifters most from their description, .\lthough EucUa is restricted to the Panamic Pro\ince in the Recent fauna, its range extended into the C'anhhean during the Neo- gene. Species presentK mcluded in this suhgemis are hsted in Table 1. Cancellaiia (Euclia) laurcttac new species (Figures 1—4, Table 2) Description: Shell (figures 1-2) to 46 nuu, thin, bi- conic, lacking umliilicus or weaklv pseudoumbilicate. Spire high (.spire angle 68-71°) conical, comprising about 1/3 shell length. Protoconch (figures 3—4) coaxial Character Mean Range (j Shell length 43.0 38.6-46.2 2 ■' Aperture length 30.6 28.0-31.8 1.5 No. of whorls, protoconch' 2.5 2.2-2.6 0,1 No. of whorls, teleoconeh 5.6 4.6-6.0 0,4 No. of axial ribs, bod\- whorl 24.6 20-30 3.0 No. apeitural lirae 13.1 0, 12-14- 0.9 ' N = 9 for this character, - Apertural lirae were not set developet! ni 2 specimens and thus scored at 0. When present, apertural lirae numbered 12- 14. The mean and standard dexiation tor this character is based on 9 specimens. xsith teleoconeh, paucispiral, of approximateh' 2V2 low, eveniv inflated, glassv whorls, separated from teleoconeh li\ flaring lip. Teleoconeh of up to 6 evenly convex whorls. Shoulder inconspicuous. Suture deepK' im- pressed. i\-\ial sculpture of 20-30 ribs, triangular in pro- file, narrower than intenening spaces. Ribs prosocline on hodv whorl, less so on siphonal canal, with conspic- uous inflection at stromboid notch. Spiral sculpture of fine, e\enl\' spaced prinian' cords (3 on eark whorls, 6- 9 on penultimate whorl, 56-62 on body whorl and si- phonal canal) that form weaklv cancellate sculpture at intersections with axial ribs. Weaker, secondary' cords ap- pear betx\een adjacent priman- cords on bodv whorl. Fine threads (0-1) present between priniaiT and sec- ondar\- cords in most specimens. Aperture deflected from coiling axis bv 20-22°, large, brtjad and evenly rounded or with outer eilge straight and neark' parallel to coifing axis. Specimens with o\ate apertures lighter in weight, with thinner parietal calluses that specimens with stniight-sitled outer fips. Outer fip finelv crenulated, weaklv outwardly flared, with broad, shallow "stromboid notch". Inner surface n"ia\' be smooth or bear 12-14 re- cessed spiral lirae, diminishing l/6th whorl into the ap- erture. Inner Up with 2 columeUar and 1 siphonal fold, each with single, sharp keel. Most posterior fold largest, o\'erla\ing weak siphonal fasciole, anteritjr siphonal fold weaker, strongly deflected anteriorly. Pustules (3—4) may be present on inductura in region lietween columellar folds. Siphonal canal narrow, nearly axial. Base color w^hite to light beige, with 3 broad, spiral bands of ginger to dark browii (darkest along sutiu'e, I at mid-whorl. 1 anterior to stromboid notch). Aperture with light tan overglaze, with darker band along (JUter lip of specimens with strong denticles. T>pe locality: Off Isla Montuosa, W of Punta Her- mosa, Isla de Coiba, Golfo de Chiriqui, Panama, trawled in 270-360 meters. T\pe material: Holotvpe, USNM 880277 , 44.9 nun: P'aratxpe 1, USNM 88()278: Paratxpe 2, LACM 2815; Parat\pe 3, ANSP 401180; Paratvpe 4, AMNH 292527; Page 116 THE NAUTILUS, Vol. 112, No. 4 Parahpes 5-6, R. Petit (Collection; Parat\pes 7-8, E. Garcia Collection; Parat)pes 9-10, J. Ernest Collection. All from type locality. Etymology: This species honors Mrs. Lauretta Marr of Midland, Texas, for her contributions to the collection and studv of Panainic niollusks. Comparative remark.s: Caiurllaria laitrctfae occurs in 2 fonns as indicated in the above description. The ovate form (figure 1) is much lighter in weight and slightly larger than the squarer form (figure 2). As no anatomical material is a\';iilable, it is not clear if these differences represent sexual dimoiphism. This new spe- cies differs from the 2 previously known Recent species included in the subgenus Euclia bv Keen, 1971, bv its lack of a well-defined tuberculate shoulder and by its more dehcate sculpture. Canccllaria laiirctfac is closest in overall appearance to the Late Miocene C. (E.) mal- donadoi Olsson, 1964 (which Woodring, 1970, incor- rectlv SMionvmized v\ith C. (E.) codazzii Anderson, 1929) than to the other Recent or fossil species. How- ever, it differs from C inahlonadoi in lacking a periph- eral cord on the shoulder, in ha\ing more numerous and finer spiral sculpture and in lacking folds along the in- ductural region of the parietal shield. ACKNOWLEDGMENTS We thank Mr. James Ernest of Panama and Dr. Emilio Garcia of Lafayette, Louisiana for making specimens of this new species available antl for donating the hpe specimens. LITERATURE CITED Adams, H. and A. Adams, 185.'S-.58. The Genera of Recent Molhisca; arranged according to their organization. 2 \ol- umes. John \an \'oorst. London, 60 pp., 138 pis. Cossmann, M. 1899. Essais de Paleontologie Coniparee. Tro- i.sieme livraison. Paris, 201 pp., 8 pis. Jousseaume, F. P. 1887. La famille des Cancellariidae (Moll- usques gasteropodes). Le Natnraliste, Annee 2e Serie [An- nee 1] pp. 155-157, 192-194, 21:3-214, 221-223. Keen, A. M. 1971. .Seashells ot Tropical West America. Second edition. Stanford Uni\ersih Press, Stanford, 1064 pp. Olsson, A. A. 1932. Contribntions to the Tertiary Paleontology of northern Peni: Part 5, The Peni\-ian Miocene. Bulletins of American Paleontolog)' 19(68)4-272, pis. 1-24. Olsson, A. A. 1964. Neogene Mollusks from Northern Ecua- dor Paeontological Research Institution. Ithaca, 256 pp., 38 pis. \\bodring, W. P. 1970. GeologN' and paleontologs ot Canal Zone and adjoining parts of Panama Description of Ter- tiar\- mollusks (Gastropoda: Eulimidae, Marginellidae to Helminthoglvptidae). United States Geological Sur\ey Professional Paper 306-D;299-452. plates 48-66. THE NAUTILUS 1 12(4): 117-1 19, 1998 Pac;e 117 Leucozonia poncJerosa, a New Fasciolariid Gastropod from Brazil Geerat J. Vermeij Department ot Geologv ami Center for Population Biologs' Uni\ersit\- of California at Davis One Shields Avenue Davis, CA 95616 USA vermeij(S'geoIog\'.ucdavis,edu Martin Averj' Snyder 745 Newtown Road Villanova, PA 19085 USA ABSTRACT A new species of fasciolariid gastropod, Leucozonia ponderosa. is described. The new species is apparentlv endemic to Trin- dade Island, which is located 1140 km east ot the Brazihan mainland. This large, tliick-shelled species is cjiaracterized by three spiral rows of tubercles, three basal cords on the last whorl, and a distinct labral tooth. Uike other described and undescribed Trindade endemics, L. pondcrosa appears to be derived from a western Atlantic ancestor INTRODUCTION The fasciolariid genus Leucozonia is a tropical American and eastern Atlantic group of shallow-water predators. In the course of a review of the species of this genus, we have discovered a distinct, imnamed species that ap- pears to be endemic to Trindade Islanil, Brazil. Here we introduce this new species as Leucozonia pondcrosa. and comment hrieflv on the western Atlantic affinities of this anil other Trindade endemics. Abbreviations used are: MNHN, Museum national d'Histoire naturelle, Paris, France: MORG, Museu Oceanografico, Funda^ao Univ- ersidade do Rio Grande, Rio Grande, Brazil. SYSTEMATICS Family Fasciolariidae Gray, 1853 Genus Leucozonia (Jrav, 1847 Type species: Murex nassa Gmelin, 1791, bv original designation. Leucozonia /lojif/cnwfl new species (Figures 1—4) Leucozonia nnssa (Gmelin, 1791). — Leal, 1991. p. 157, pi. 20. figs. B. C (in part). Diagnosis: A Leucozonia with five or six primary- spi- ral cords on the last whorl, the adapical three rows being sculptured by low, rounded tubercles; distinct labral tooth present at end of third cord. Description: Shell large (maximum length 67 mm), moderately sijuat (length:v\idth 1.56-1.72), moderately high-spired (last whorl length:length 0.53-0.59), heavy and solid. Color light-brovvii. Spiral sculpture ol last whorl consisting of six (sometimes five) cords, one at shoulder, a second below the shoulder, and a third cen- tral, and three (sometimes two) basal cords, all separated bv fine threads. Axial sculptiu-e oi last whorl consisting of eight strong, roimded ribs, forming low, roundeil nodes where thev are crossed bv each of the three adap- ical spiral cords. Outer fip with distinct atlapical sinus, convex metlial sector, and weaklv concave abapical sinus. Third spiral cord enihng in well-developetl labral tooth. Adaxial side of outer lip with 11-12 strong, beaded firae. Aperture relatively broad (aperture length:aperture width 2.4—2.7), color fight-peach. Siphonal fiisciole well developed; umbilical sfit absent. Siphonal canal relatively long (siphonal canal length:aperture length 0.38-0.42). Type locality: Enseada dos Portugueses, Trindade Is- land, Brazil, 20°.30'S, 29°20'VV. depth 10 m, rocky bot- tom. Type material: HolotApe, MORG 39298. length 47.3 mm, width 28.8 mm, aperture length 28.0 mm, aperture wiilth 10.5 mm, siphonal canal length 11 mm; Paratvpe 1, MORG 39299, length 66.8 mm,\vidth 40.0 mm, ap- erture length 35.5 mm, aperture width 14.2 mm, si- phonal canal length 15 mm; Paratvpe 2, MNHN, length 62.8 mm, width 36.6 mm, aperture length 35.7 mm, ap- erture width 15. 1 mm, siphonal can;d length 15 mm, collected by J, H. Leal and R Bouchet, SCUBA, 10 m, 22 May 19S7 (all from type locafitv). DISCUSSION Two species of Leucozonia occur on Trindade Island. One of these is a thick-shelled, obsoletelv' spirally sculp- tured form of the widespread western Atlantic species Leucozonia nassa (Gmefin, 1791) (figures 5-6). This f(jrm, which also occurs at Fernando de Noronha Ar- Page 118 THE NAUTILUS. Vol. 112. No. 4 Figures 1-2. Lciicozoiiit: jxindcrosn new species, Tnndade Lsland. Brazil, length 66.8 nun, parahpe 1, .\1(JRG 39299. Figures 3—4. Lciicozonia pcinilcrosd. Trindade Island, Brazil, length 47.3 mm. liolot\pe, MORG 39298. Figures 5-6. Lericozonin nnssa. Trindade Island, Brazil, length 52.1 mm, MNIIN nnnnmbered, Enseada dos Portugueses, Trindade Island. Brazil. J. H, Leal and P. Bouehet, SCUBA, 10 ni depth, 22 May 1987 chipelago, Atol da.s Roca.s. and B;iliia (all in Brazil), is characterized b\- .spiral sculpture con.si.sting of weak threads or sometimes very weak spiral cords. If cords are present, they number six or seven from the shoulder to tlie central, tooth-hearing cortl. The single specimen we have seen from Trindade has a dark-peach colored aperture and strongly granulated lirae on the adaxial side of the outer Up. The second species from Trindade. which we distinguish as the new species L. poiulcrosa. is characterized by three distinct, nodose adapical spiral cords, one at the shoulder, a central cord that bears the labral tooth, and a cord l)etween these two. In addition to tliese adapical cords, there are three (sometimes two) major cords below the tooth-bearing cord. The aperture is of a hght-peach color, and the Urae on the ailaxial side of the outer lip are weakly beaded. Some specimens of L. luissa from Cma^ao and else- where in the southern and eastern (Jaribbean superfi- cially resemble /. poiulcrosa in having three nodose adapical cords, but tlu'y differ from L. poiulcrosa in hav- ing the two adapical cords so close together that the shoulder appears to have a double keel. In L. pouilcrosa. the .second row of nodes lies farther from the shoulder Moreover, the aperture of Caribbean specimens of L nassa is white instead of peach-colored. Two other species of Lciicozonia have three adapicid cords sculptured by nodes or tubercles. L. triscrialis (La- marck, 1822) from the Cape Verde Islands chffers from L. poiulcrosa in having a much more tuberculate shell, one basal cord instead of three, and a white instead of a peach-colored aperture. L. tiihcrculata (Broderip, 18.33) from the Galapagos and Cocos Islands in the east- ern Pacific differs from L. poiulcrosa h\ having two in- stead of the usual three basal cords, by being smaller in size, and by having a white aperture. Moreover, the third (tooth-bearing) cord of L tuhcrculata is not nodose, whereas that of L poiulcrosa bears nodes. BIOGEOGRAPHY OF TRINDADE Trindade Island, located 1140 km east ol tlu- l^razilian mainland at 2()''3n' S, 2y°2()' W, is a small (4.8 km m length), high island in the South .\tlantic. Its gastropod fauna was first systematically studied by Leal (1991). Species in the islands fauna that are not endemic are widely distributed in the tropical western Atlantic. Leal ( 1991 ) reported that about 169f of the gastropod species in the launa of Trindade are endcTuics. Thus far. only two ol these, the lottiid Patelloida inarcusi (Righi, 1966) and the neritid \crita asccnsiouis triiuladccusis \'ermeij. G. J. Vermeij and M. A. Snyder, 1998 Page 119 1970, have been de,scribed Both .species also occur at Martin Vaz, a tinv archipelago 48 km east of Trindade (Leal, 1991). In addition. Leal (1991) reports unnamed species endemic to Tiinilade in the genera Diodoro. Cal- liostoma. S